October 2020 Wetland Science & Practice

Wetland Science
&
Practice published by the Society of Wetland Scientists
Vol. 37, No. 4 October 2020
ISSN: 1943-6254
SWS - Celebrating Our 40th Anniversary in 2020

FROM THE EDITOR’S DESK CONTENTS
Vol. 37, No. 4 October 2020
Greetings to all. As we wrap up our issues for 2020, the ISSN: 1943-6254
corona virus still affects everyday life and fires continue
to burn in the western United States and in South Ameri- 214 / From the Editor’s Desk
ca’s Pantanal, claimed to be the 216 / President’s Message
world’s largest tropical wetland. 218 / SWS News
These fires are devastating and 219 / Awards
cause for great concern not 220 / SWS Events
only for residents but also for
222 / SWS Webinars
natural resource managers. For
perspective, in California alone, 223 / Wetland Practice:
Passing of a Leader in Wetland Policy Development
an area the size of Rhode Island
has burned so far this year. In
224 / INTRODUCTION TO THIS ISSUE: EMPHASIS
one week, more land burned
ON LATIN AMERICAN WETLANDS
in California than in all of
231 / Statement on Pantanal Fires
Ralph Tiner 2018 and twice as much as had
WSP Editor burned in 2017 according to
the state’s forestry department. ARTICLES
In the Pantanal, the number of Wetland Research
fires this year are over 10 times the number that occurred 232 / Wetland Science in Latin America and the Caribbean
in 2018. While lightning is responsible for the California Region: Insights into the Andean States
blazes with a couple of exceptions, the combination of K.S. Navarro and others
lightning and intentional human-set fires (e.g., to con- 241 / Wetlands of the Coast of Lima: Patterns of Plant
vert forest to cattle ranches) are the causes for those in Diversity and Challenges for their Conservation
the Pantanal (see some articles listed in “Wetlands in H. Aponte
the News” for details). On September 15 and 16, I even 246 / Andes, Bofedales, and the Communities
witnessed the effect of the U.S. western wildfires on our of Huascarán National Park, Peru
skies in Massachusetts at sunset (see images below; pho- R.A. Chimner and others
tos by Barbara Tiner). In one image you’ll see the sun 255 / Peatlands of the Central Andes Puna, South America
appears as a red dot! Didn’t smell any smoke as it was E. Oyague and D.J. Cooper
thousands of feet above the land surface. 261 / What is the Flora of the Pantanal Wetland?
Along the U.S. East and Gulf Coasts, hurricane A. Pott and V.J. Pott
season is in full swing with AccuWeather predictions for 267 / Connectivity of River Floodplains – the Case of Ibera Wetlands
as many as 28 names storms and 13 hurricanes. To date after 10,000 Years of Isolation from Parana River
we’ve had eight landfalls and we’ve still got a couple J.J. Neiff and others
of months to go before the season ends (normally we 283 / Urban Wetland Trends in Three Latin American Cities
get 3 or 4 landfalls each year). All this plus more news during the Latest Decades (2002-2019): Concón
(Chile), Barranquilla (Colombia), and Lima (Peru)
on melting glaciers and permafrost. Climate change is
C. Rojas and others
real and affecting both wildlife and human populations.
Anything positive? For sports lovers, there is something
Wetland Conservation/Education/Outreach
to watch and for all, there are more opportunities for din-
294 / Wetland Conservation Concerns in Southern Mexico
ing out, especially while the weather is good. Interesting T. Lobato de-Magalhães and others
times indeed. Also we’ve got a great issue for you!
302 / Propagation of Endangered Aquatic Plants: An Experience that
This issue of Wetland Science & Practice has a theme Promotes ex situ Conservation and Environmental Education
– Latin American wetlands. Last fall in speaking with S.N. González Mateos
Tatiana Lobato-de Magalhães about her article for the
January 2020 issue, I mentioned that it would be great if
we could have an entire issue emphasizing ongoing wet-
land work in South and Central America. She agreed and
took the lead in contacting potential contributors while I
contacted a couple of colleagues who I knew were work-
From the Editor’s Desk, continued on page 223
214 Wetland Science & Practice October 2020
NOTES
Wetland Science
&
Practice
PRESIDENT / Loretta Battaglia, Ph.D.
PRESIDENT-ELECT / Gregory Noe, Ph.D.
Wetland Research IMMEDIATE PAST PRESIDENT / Max Finlayson, Ph.D.
308 / Bark Traits: A Predictor for Recognition of Successional SECRETARY GENERAL / Leandra Cleveland, PWS
TREASURER / Lori Sutter, Ph.D.
Groups in Riparian Forest Species
EXECUTIVE ADMINISTRATOR / Suzanna Hogendorn
J. Rodrigues da Silva and others CONSULTING DIRECTOR / Michelle Czosek, CAE
310 / Mangrove Ecological Restoration Inside Pantanos WETLAND SCIENCE & PRACTICE EDITOR / Ralph Tiner, PWS Emeritus
de Centla Biosphere Reserve (Tabasco, Mexico)
R.A. Betancourth and others CHAPTERS
ALASKA / Emily Creely
312 / Diversity of Temporary Ponds from the Guajira, Colombia ASIA / Wei-Ta Fang, Ph.D.
C.E. Tamaris-Turizo and others CANADA / Gordon Goldborough, Ph.D.
314 / Treatment Wetlands – Experiences in Mexico CENTRAL / Katie Astroth
Armando Rivas CHINA / Xianguo Lyu
EUROPE / Matthew Simpson, PWS
INTERNATIONAL / Ian Bredlin, Msc; Pr.Sci.Nat and
Wetland Conservation/Education/Outreach Tatiana Lobato de Magalhães
316 / An Ecosystem-based Approach to Managing Fish, Cattle MID-ATLANTIC / Jennifer Slacum
NEW ENGLAND / Dwight Dunk
and Forests on the Amazon Floodplain NORTH CENTRAL / Christina Hargiss, Ph.D.
D.G. McGrath and others OCEANIA / Phil Papas
319 / Strengthening Goverance in the Monterrico Multiple Use Natural PACIFIC NORTHWEST / Josh Wozniak
Reserve: Planning for Conservation with a Bottom-Up Approach ROCKY MOUNTAIN / Ryan Hammons, PWS
A. Silvia Morales SOUTH ATLANTIC / Brian Benscoter, Ph.D.
SOUTH CENTRAL / Scott Jecker, PWS
321 / Amphibious Colombia: A Country of Wetlands WESTERN / Richard Beck, PWS, CPESC, CEP
R. Ayazo Toscano and others
323 / Amphibian Territories in Transition: Socio-ecological SECTIONS
BIOGEOCHEMISTRY / Lisa Chambers, Ph.D.
Rehabilitation of Wetlands EDUCATION / Derek Faust, Ph.D.
R. Ayazo Toscano and others GLOBAL CHANGE ECOLOGY / Wei Wu, Ph.D.
325 / Guardians of Wetlands (Los Guardianes de los Humedales): PEATLANDS / Bin Xu, Ph.D.
Young Peruvians Committed to Wetlands PUBLIC POLICY AND REGULATION / John Lowenthal, PWS
H. Aponte RAMSAR / Nicholas Davidson
WETLAND RESTORATION / Andy Herb
326 / We Are Wetlands (@Somos_humedales) WILDLIFE / Andy Nyman, Ph.D.
K. Paz Arteaga WOMEN IN WETLANDS / Carrie Reinhardt Adams, Ph.D.
327 / Urban Wetlands Interactive Platform STUDENT / David Riera
C. Teutsch Barros
COMMITTEES
AWARDS / Siobhan Fennessy, Ph.D.
328 / Wetlands in the News HUMAN DIVERSITY / Kwanza Johnson and Jacoby Carter, Ph.D.
MEETINGS / Yvonne Vallette, PWS
331 / Weltand Bookshelf PUBLICATIONS / Keith Edwards
333 / What’s New in the SWS Journal - WETLANDS MEMBERSHIP / Leandra Cleveland, PWS
WAYS & MEANS / Lori Sutter, Ph.D.
334 / About WSP/Submission Guidelines SWS WETLANDS OF DISTINCTION / Roy Messaros, Ph.D.
Bill Morgante and Jason Smith, PWS
REPRESENTATIVES
COVER PHOTO: PCP / Scott Jecker, PWS
Peatland (bofedal), Ayacucho, Peru. Photo by Héctor Aponte. STUDENT / David Riera
WETLANDS / Marinus Otte, Ph.D.
WETLAND SCIENCE & PRACTICE / Ralph Tiner, PWS Emeritus
ASWM / Jill Aspinwall
RAMSAR / Nicholas Davidson, Ph.D.
AIBS / Dennis Whigham, Ph.D.
SOCIETY OF WETLAND SCIENTISTS

1818 Parmenter St., Ste 300,
Note to Readers: All State-of-the-Science reports are peer Middleton, WI 53562
reviewed, with anonymity to reviewers. (608) 310-7855
www.sws.org
Wetland Science & Practice October 2020 215
PRESIDENT’S ADDRESS
REFLECTIONS, CONNECTIONS AND THE CHANGING SEASONS
I’ve been watching intently and enjoying the seasons change

in southern Illinois, from summer to autumn, and thinking
about other lovely parts of the world
now shifting from winter to spring.
With all of the challenges and
disappointment 2020 has brought,
it has also been a time to slow
down and reflect on past travel,
fieldtrips, and good times with
colleagues from all over the world.
The love of wetlands has led many
of us to travel far and wide and
Loretta Battaglia, Ph.D. also brought us together. Just as
our precious waters and wetlands
Southern Illinois
connect landscapes, so too do they
University connect us through shared experi-
SWS President ences, passion for conservation
and restoration, and the desire to
discover their intricacies and ecological secrets. These
connections run deep. If there has ever been a time to sus-
tain and nurture these connections, it is now.
The virtual platforms that have enabled conversations,
collaborations and the ongoing business of our Society are
admittedly not as much fun as slogging through a swamp or
bog together or meeting up in a cozy corner at our annual
meeting to brainstorm projects and initiatives. They have
kept us going, however, and it is likely they will remain a
part of the way we operate long after the Covid-19 pandemic
is over. Virtual tools have and will continue to be valuable
complements to our conventional mode of operations. They
will also help lessen the Carbon footprint of our events while
better supporting the needs of a global wetland community
that is geographically broad but aims to be inclusive.
I hope you will join us in celebrating the 40th anni-
versary of SWS by participating and tuning in to our first
Society-wide virtual meeting (December 1-3, 2020), aptly
named Wetland Connections over 40 years. n

SWS NEWS
WORLD’S LEADING AQUATIC SCIENTIFIC SOCIETIES PUBLIC POLICY UPDATES

URGENTLY CALL FOR CUTS TO GLOBAL GREENHOUSE GAS County of Maui v. Hawai‘i Wildlife Fund amici brief
EMISSIONS featured in PBS NOVA podcast
In an unprecedented statement released September 14, 110 SWS' Royal Gardner and his team of attorneys, as well as
aquatic scientific societies, including SWS and representing many of our members and members of our aquatic soci-
more than 80,000 scientists across the world, joined forces ety partners, played a role in preparation of an amici brief
to sound a climate change alarm. The societies call for dras- cited earlier this year in the U.S. Supreme Court’s opinion
tically curtailed global greenhouse gas emissions to avoid in County of Maui v. Hawai‘i Wildlife Fund. On April 23,
the worst impacts of man-made climate change to fish and 2020, SCOTUS ruled 6-3 that the Clean Water Act (CWA)
aquatic ecosystems. Unless urgent action is taken to reduce covers the functional equivalent of direct discharges of pol-
emissions, scientists predict catastrophic impacts to com- lutants to navigable waters.
mercial, recreational, and subsistence fisheries and human The amici brief was cited by Justice Breyer, who wrote
health and global economies. n the opinion for the majority, “Virtually all water, polluted
or not, eventually makes its way to navigable water. This
is just as true for groundwater. See generally 2 Van Nos-
WETLANDS OF DISTINCTION
trand’s Scientific Encyclopedia 2600 (10th ed. 2008) (defin-
The newly-designated Wetland of Distinction Quakertown
ing “Hydrology”). Given the power of modern science,
Swamp (Pennsylvania) received some favorable press from
the Ninth Circuit’s limitation, “fairly traceable,” may well
the Pennsylvania Intelligencer (https://www.theintell.com/
allow EPA to assert permitting authority over the release
story/news/local/2020/07/22/quakertown-swamp-honored-
of pollutants that reach navigable waters many years after
as-ldquowetland-of-distinctionrdquo/112701110/), as
their release (say, from a well or pipe or compost heap) and
well as in this news video (https://www.wfmz.com/news/
in highly diluted forms. See, e.g., Brief for Aquatic Scien-
area/southeastern-pa/quakertown-swamp-gets-wetland-
tists et al. as Amici Curiae 13–28.”
of-distinction-title/article_f6c60f74-e7ea-11ea-838d-
Steph Tai, a member of Royal's legal team, was re-
4707c0188c10.html)!
cently interviewed for the PBS NOVA podcast Science in
And, the first Wetland of Distinction outside the US
the courtroom: https://www.pbs.org/wgbh/nova/podcast/.
has been designated, as featured in Restoring a gem in the
The discussion on the Maui case and the brief begins about
Murray-Darling Basin: the success story of the Winton
14:30. Our work is clearly making a difference!
Wetlands (https://theconversation.com/restoring-a-gem-in-
the-murray-darling-basin-the-success-story-of-the-winton-
wetlands-140337). SWS Past President Max Finlayson WOTUS Rulings
co-authored this article about the Winton Wetlands in the Colorado
Murray-Darling Basin of Australia. In addition to being a Some good news: the rule appears to be enjoined in Colora-
story about SWS' first Wetland of Distinction designated do (where judge did not permit amici briefs): https://www.
outside of the United States, this article details a success bloomberglaw.com/public/desktop/document/StateofColo-
story about a community-based wetland restoration venture. radoThevUSEnvironmentalProtectionAgencyetalDocket-
Visit https://www.wetlandsofdistinction.org/ for infor- No120/4?1592664339
mation about this impoartant SWS initiative, and to nomi-
nate your favorite wetland to be recognized as a Wetland of California
Distinction today! n The Court denied the motion for a preliminary injunction,
and the Navigable Waters Protection Rule will go into ef-
fect on Monday. Here is the order: https://sws.org/images/
MAKE A DIFFERENCE FOR SWS THIS HOLIDAY SEASON pdfs/CA_v_Wheeler_PI_decision.pdf n
Do your Amazon shopping at smile.amazon.com/ch/48-
1146960 to generate donations for SWS, at no additional
cost to you. n

AWARDS
Introducing the Society of Wetland Scientists 40th Anniversary Award

PURPOSE: On the occasion of the 40th Anniversary of the statement by each of two referees in support of the nomi-
Society of Wetland Scientists the Executive Board would like nation against one or more of the categories for the award
to recognize high level and sustained contributions to wetland (wetland research, practice, education, communication,
research, practice, education, or communication, or service to or support to the Society). These letters should outline the
the Society. These awards are additional to the awards offered reasons for the nomination and provide tangible exam-
annually by the Society and covers the breadth of professional ples or evidence. This information should accompany the
activity normally undertaken by members. Nominations for the nominations forwarded from the Chair of each Chapter,
Award are open to all members in good standing through the Section or Standing Committee to the Chair of the Selec-
Chairs of the Chapters, Sections and Standing Committees. tion Committee (fennessym@kenyon.edu) and clearly
marked “SWS 40th Anniversary Award Nomination”.
PROCESS:
4. In making nominations please consider the outcomes
1. Each Chapter, Section and Standing Committee is invited and impact on wetland research, practice, education,
to nominate 1 or 2 of their members (in good standing) communication, or support to the Society, and the period
for consideration as a recipient of the Society of Wetland over which these have occurred in addition to any indi-
Scientists 40th Anniversary Award for high level and vidual or one-off high value contributions. The emphasis
sustained contributions to wetland research, practice, is on outcomes and impact, not just outputs (that is,
education, communication, or support to the Society. while the quality of research publications is an important
This message is being sent to all Chairs and also to all indicator, it is not the sole indicator in the assessment of
members via the Society’s web page news service. importance and impact). The nomination should sum-
2. The Chair of the Chapter, Section or Standing Commit- marize the evidence in support of the high level and
tee, or their nominee, is asked to seek nominations from sustained contributions made by the nominee.
their members and arrange for the supporting informa- 5. The award will comprise a certificate with an inscrip-
tion for their agreed 1 or 2 nominations to be forwarded tion reading “Society of Wetland Scientists 40th
to the Chair of the Awards Committee by November 13, Anniversary Award for high level and sustained
2020. In a cover, message the Chair (or nominee) should contributions to [wetland research, practice, educa-
describe in 3-4 sentences the reasons for the nominations tion, communication or service to the Society]” and
along with a separate statement about how they avoided be announced at the 2020 Virtual Conference.
any potential conflicts of interest in choosing their nomi-
6. The decision on the number of awards and awardees
nees. The Executive Board will also be invited to make
will be made by the Selection Committee. Late nomina-
nominations with a particular emphasis on sustained con-
tions cannot be accepted and the Committee will not
tributions, including for support to the Society through
enter into any discussion with other parties about the
contributions to good governance of the Society.
award processes. The Selection Committee will be head-
3. Each nomination should consist of a short (1-2 pages) ed by the Chair of the Awards Committee, along with
Curriculum Vitae of the nominee and a succinct 1 page the President and two Past Presidents of the Society. n
Student Section Virtual Conference SWS Grants International Travel Awards

Award Winners The Awards Committee awarded two 2020 International Trav-
The SWS Student Section held a virtual conference in June el Grants, which will be deferred to 2021. Congratulations:
2020, and have announced award winners. • Rajashree Naik, Rajasthan, India
Abstract: “Ecological Status of Largest Saline Wetland-
LIGHTNING TALK WINNERS scape of India: A Study of Sambhar Lake under Driver-
• First Place: Amanda Loder, Nova Scotia, Canada Pressure-State-Impact- Response (DPSIR) Framework”
• Tie for Second Place: • Pankyes Datok, Occitanie, France
Andrea Stumpf, Massachussetts, United States Abstract: “Investigating the role of the Cuvette Cen-
Chelsea Duball, Wyoming, United States trale wetlands in the hydrology and Organic carbon
• Third Place: Clay Tucker, Louisianna, United States fluxes of the Congo River basin”

SWS EVENTS
2020 Virtual Meeting

December 1-3, 2020
The Society of Wetland Scientists (SWS) is excited to an- There are limited sponsorship opportunities remaining:
nounce our inaugural virtual meeting on December 1-3, https://static1.squarespace.com/
2020, themed “Wetland Connections Over 40 Years.” We static/5967a224725e258a852d731e/t/5f848650b21edc3a8a
listened to your feedback and decided that a smaller, virtual cd6790/1602520658016/Sponsor+opportunties+SWS+2020-
meeting is in order to celebrate the SWS 40th anniver- updated+10.12.pdf n
sary and to showcase our members’ outstanding work in
wetlands. Our goal is to offer an abbreviated format for
the SWS community to connect, to share research, and to
continue expanding our Society’s global network. Be sure
to visit our event website for all the details! swsvirtual-
meeting.org
Thank you to sponsors:

2021 Annual Meeting
June 1-4, 2021
The Society of Wetland Scientists is planning the 2021 an- Although we cannot predict what June 2021 will look like
nual meeting, to occur in Spokane, Washington from June for the ever-evolving world pandemic, we are planning a
1-4, 2021. Our theme next year is Wetland Science 2021: compelling program of speakers, research and presentations
Adaptation Drives Innovation. We have selected the very that will have a voice and platform for discussion. If all goes
industrious and innovative North American beaver (Castor well, we plan on hosting the conference at the Davenport
canadensis) as the conference mascot. Their current role Grand, in the heart of the great town of Spokane. There are
and profound historical influence on watersheds, wetlands extensive parks and open spaces at the doorstep of the hotel,
and hydrology is a growing field of study and restoration. and wildlife refuges and wilderness a short drive away. We
They also provide a model of wetland management and are also planning for contingencies, including more, smaller
integration of habitats with relevance to our work. We are session gatherings and opportunities to remotely share infor-
developing program symposia now and are compiling local mation with folks who may have difficulties traveling.
and international research topics that tackle the world’s
more challenging wetland management issues. We hope you can join us at the SWS 2021 Annual Meeting!
https://www.swsannualmeeting.org/ n
STUDENTS - APPLY FOR SWAMMP PARTICIPATION

Deadline: Monday, November 16, 2020
The SWS Multicultural Mentoring Program (SWaMMP) works to increase diversity within the Society of Wetland Scien-
tists and throughout the environmental sciences. SWaMMP enables undergraduate students from underrepresented groups
to attend and receive full travel benefits to the SWS Annual Meeting, held in Spokane, Washington, June 1-4, 2021. The
Annual Meeting offers students valuable career guidance and opportunities to network with leading wetland science pro-
fessionals from around the world.
SWaMMP provides:
• Conference registration, lodging and all travel expenses to the SWS Annual Meeting
• Individual career mentoring
• Postgraduate and career workshops
• Networking opportunities to meet professionals from diverse fields
• Opportunity to present a research poster
More info: http://sws.org/Awards-and-Grants/sws-undergraduate-mentoring-program-swammp.html n

WEBINARS
SOCIETY
WETLAND
SCIENTISTS
Monthly webinars are offered by the Society of Wetland Scientists (SWS) as a benefit of membership. Once each quarter, in
March, July, September and December (marked with an asterisk below), the monthly SWS webinar is open for non-members
to attend, at no cost, as well. Spanish language webinars are always free for both members and non-members.
ENGLISH: SPANISH:
sws.org > Events > Upcoming Webinars sws.org > Events > Spanish Language Webinars
11/19/2020 | 1:00 pm ET 12/9/2020 | 1:00 pm ET

Resurrecting ‘ghost ponds’ and other approaches Urban Wetlands, an opportunity
in pond restoration and conservation to make sustainable cities *
Presenter: Emily Alderton Presenter: Carolina Rojas Quezada
12/17/2020 | 9:00 am ET 3/24/2021 | 1:00 pm ET

SWS History - 40 Years of Globalization * The history of biodiversity origin and humanity
Presenters: Kathy Ewel, Gillian Davies, Fred Ellery, future through Cuatro Ciénegas wetlands
Wai-Ta Fang, Max Finlayson, Luisa Ricaurte Presenter: Valeria Souza
2/18/2021 | 1:00 pm ET
Assessing Vegetative Species Re-colonization
of Commercial Cranberry Bogs
Presenter: Kate McPherson
ARCHIVES
Did you miss a webinar?
All webinars are recorded and archived for complimentary viewing by members on our Past Webinars web page.
Thank you to our 2020 Webinar Series Sponsors

For more information on sponsoring the SWS Webinar Series, contact membership@sws.org
WWW.IN-SITU.COM WWW.FACEBOOK.COM/ WWW.WHITENTONGROUP.COM/ HTTPS://WILDNOTEAPP.COM/

WATERRESOURCESHYDROLOGY
HYDRAULICSEDUCATION/
#SWSWebinars #SWS #WetlandScience #WetlandResearch

IN REMEMBRANCE
Passing of a Leader in Wetland Policy Development

Dr. Jon Kusler passed away on October 8, 2020. Jon was mover and a shaker for wetland policy and wetlands now
a leader in helping guide national wetland and floodplain receive more attention in the policy arena in large part due
policy since the 1970s. He wrote hundreds of papers on a to his efforts and those of his collaborators at ASWM and
variety of topics ranging from wetland regulation, restora- elsewhere. We'll miss Jon, may he rest in peace.
tion, functional assessment to floodplain management. He The ASWM has established a memorial page on its
was the founder of the Association of State Wetland Manag- website for folks to post sentiments and share stories about
ers (ASWM), an organization that will continue Jon's legacy Jon (https://www.aswm.org/aswm/10210-jon-kusler-memo-
and press for improvements in wetland and floodplain con- rial). They have also established the Jon A. Kusler Student
servation. In 1990 he received the first National Wetlands Scholarship Award to fund student attendance at ASWM's
Awards Lifetime Achievement Award from the Environmen- annual State/Tribal/Federal Coordination meeting (for de-
tal Law Institute in recognition of his dedication to wetlands tails, see https://www.aswm.org/aswm/donate). n
(http://elinwa.org/awards/recipients/jon-kusler). He was a
From the Editor’s Desk, continued from page 214

ing with South American scientists on high-altitude wetlands. I hope that we can get leaders of various SWS sections
What a fantastic job she did - thanks to Tatiana this issue con- and chapters, or others to help coordinate contributions
tains 9 articles and 11 notes on various research, conservation, so that we can produce other thematic issues of WSP. If
and public education/outreach efforts in Latin America (LA). interested in taking on the challenge, please contact me at
Needless to say, this kept me quite busy over the past few ralphtiner83@gmail.com. The Wetlands of Distinction team
months. While the collection is not intended to be a compre- has plans to provide profiles of individual Wetlands of Dis-
hensive overview of the ecology and conservation of LA wet- tinction for publication in WSP on a quarterly basis.
lands, the texts should provide readers with a good introduction Meanwhile, I hope all is well and that a vaccine and ef-
to these wetlands and the challenges they face, and what some fective vaccination program will arrive in 2021 well before
folks are doing to improve the status of wetlands across the our next annual meeting in Spokane, Washington.
region. I again want to especially thank Tatiana and all of the Happy Swamping and Stay Safe. n
contributors for their efforts in making this issue possible.
Sunset, Leverett, MA on September 15, 2020. Sunset, Leverett, MA on September 16, 2020.

INTRODUCTION
This Issue: Emphasis on Latin America Wetlands

Tatiana Lobato de Magalhães, Co-chair International Chapter of the Society of Wetland Scientists
n behalf of the International Chapter and representing Latin America and the Caribbean, I am proud to announce
O that we now have the first region-focused Wetland Science and Practice issue - the October 2020 Latin America
issue. This SWS effort is an important step to strengthen the Society’s internationalization as well as to spread the word
on the Society across the Latin American countries. We invited several wetland scientists and practitioners to publish
their research or summaries of their research or public outreach projects. This resulted in publication of nine articles and
11 notes in this issue. Here, we can travel across the huge Latin American region, from Mexico to Patagonia, addressing
both coastal and highland wetlands and wetlands in between. This issue provides an introduction to the region’s wetlands
and examples of ongoing Latin America wetland science and wetland education/public outreach efforts. Such activities
include remarkable wetland biodiversity research in Argentine Parana delta and Brazilian Pantanal; challenges for man-
agement and conservation in the Brazilian Amazon, Chile, Colombia, Guatemala, and Mexico; wetland organizations’
database in the Andean States; education and awareness in the Chilean Patagonia and Coastal Peru, among others.
Thanks to all contributors and the editor Ralph Tiner for supporting this innovative idea and reviewing/editing the
contributions. We hope you enjoy reading about Latin American wetlands and learn something about them and the chal-
lenges they face. n
PICTORIAL OVERVIEW OF LATIN AMERICAN WETLANDS
These images are intended to provide readers with a view of some of the wetlands found across Latin America. While
they do not show all the types, they reflect the beauty and diversity of types that occur in this species-rich region. We
thank the photographers for their contributions. More images are included in the articles and notes that follow.
Estuarine salt marsh (Rocuant-Andalién), Talcahuano, Chile by Christopher Momberg and URBANCOST.

Mangroves, Ceará, Brazil by Tatiana Lobato de Magalhães.
Ephemeral wetlands, Lençóis Maranhenses, Brazil by Tatiana Lobato de Magalhães.

Amazon rainforest (location unknown) by Jonathan Lampel (courtesy of unsplash.com).

Freshwater wetland, Los Lagos, Los Rios Region, Chile by Alejandro Jara.
Lacustrine marsh, Panguipulli, Los Rios Region, Chile by Karina Arteaga.

Highland emergent wetland (temporarily flooded meadow), Guanajuato, Mexico by Tatiana Lobato de Magalhães. (Note: Eleocharis densa is the promi-
nent tall plant in the foreground.)
Cushion plant bofedales, Cordillera Apolobamba, Natural de Manejo Integrado Nacional Apolobamba, and Ula Ula National Fauna reserve (for vicuña),
4695 m elevation, Bolivia by David J. Cooper.

Highland wetland (Páramos), El Cajas National Park, Ecuador by Tatiana Lobato de Magalhães.
Inland salt flat (white strip) along Laguna Colorada, Bolivia’s “Red Lake” by Miguel Navaza (www.flicker.com; Creative Commons CC BY-NC-SA 2.0).
Its red color is due to red sediments and algae. This area is part of Eduardo Avaroa Andean Fauna National Reserve and Los Lípez - a Ramsar Wetland
of International Importance. Three species of flamingos can be found here.

Lago Junin (also called Lago Chinchaycocha), in Peru’s Reserve Nacional de Junin (the largest peatland in the Andes, 45 km long and 16 km wide,
covering more than 500 km2) by David J. Cooper.
Sphagnum magellanicum cushions on the edge of a ombrotrophic bog near Ushuaia, Argentina by David J. Cooper.

Fen in Colombia’s Park Nacional Sumapaz by David J. Cooper. The large plants are characteristic of the paramos region, and
feature species of Puya and Espeletia.
PANTANAL
Fire in Paradise Declaration: A Call for Your Participation
Scientists of the world, coming from across all disciplines If you agree with it, you can sign the Declaration at:
and working across a wide range of subjects and themes, https://forms.gle/r7ArbQvoQYvGk6tk9 and express your
are very concerned with the extensive fires which have opinion about a problem which is taking place in Brazil,
been taking place in the Amazon Rainforests and in the but which concerns the whole world. Please feel free to
Pantanal Region (shared by Brazil, Bolivia and Paraguay), pass this note on to other colleagues. n
the planet's largest tropical wetland. In order to express our
concern, a document titled "Fire in Paradise: Declaration
of World Scientists" has been prepared, asking for a set of
actions aimed at addressing the problem. The Declaration
can be seen at:
https://www.haw-hamburg.de/en/university/newsroom/
news-details/news/news/show/fire-in-paradise-declaration-
of-world-scientists/

WETLAND RESEARCH
Wetland Science in Latin America and the Caribbean Region: Insights into the Andean States
Karol Salazar Navarro1,2, Alexandra Dallely Olortigue Tello2, Héctor Aponte2, and Tatiana Lobato-de Magalhães3,4
Abstract: Through a mentoring initiative of the Society of zil), Mesoamerica, and the tropical Andes, which require
Wetland Scientists International Chapter, including volun- greater effort for their conservation since endemic species
teer scientists, students, and early-career professionals, we of plants and vertebrates are being affected by habitat loss
created a database of 283 organizations focused on wetlands and fragmentation (Myers et al. 2000). Across the exten-
for the Andean States (Argentina, Bolivia, Chile, Colom- sive LAC region, 350 Ramsar Sites are protecting 700,000
bia, Ecuador, Peru, and Venezuela). This review includes km2 of wetlands and a variety of wetland ecosystems such
data about organizations spatial distribution among seven as mangrove forests, swamp forests with palms, flooded
countries, their sectors, and their principal research areas as savannah and forests, marshes, and peatlands (Junk and
well as Andean Ramsar Sites data. The most representative Piedade 2011; Ramsar 2020). Despite the high ecosystem
research areas were hydrogeology, biogeochemistry, and bio- services and economic values estimated for wetlands,
diversity (represented by fauna, flora, algae, phytoplankton, especially for mangroves and peatlands, LAC wetlands are
and zooplankton). We observed a lack of wetland restoration, extremely threatened and at risk of disappearing. In the last
mapping, and management research. The academic sector four decades, the estimated wetland losses are around 60%
(universities) had the largest number of organizations (176 in LAC and worldwide (Junk 2013; Davidson 2014; Darrah
organizations), followed by government (51), non-profits et al. 2019; Davidson et al. 2019).
(46), and the private sector (10). The Andean States harbor One of the aims of the International Chapter of the
a total of 94 Ramsar Sites covering 300,000 km2 distributed Society of Wetland Scientists (SWS) is to increase wetland
in seven countries. Through this review, we highlight the science and conservation in the LAC region. Through an
magnitude of wetland science in the Andean States and hope integrated database for LAC, we seek to link scientists and
to support a future wetland scientist network for the Latin wetland professionals at an international level, as well as to
American and Caribbean (LAC) region and allow LAC sci- engage young people in wetland activities. To achieve this
entists to connect internationally. objective, the SWS International Chapter in collaboration
with volunteers of Universidad Científica del Sur in Lima,
INTRODUCTION Peru, including students and early-career professionals,
The Latin American and Caribbean (LAC) region contains created a database of organizations focused on wetlands
46 countries and non-independent territories (e.g., Aruba, for the Andean States. Here we present the results for the
Curaçao, French Guiana, and Puerto Rico) (FAO 2020). Andean States, a region that encompasses seven countries
Overall, LAC countries harbor a high number of species of (Argentina, Bolivia, Chile, Colombia, Ecuador, Peru, and
fauna and flora, as well as unique neotropical ecosystems. Venezuela) with unique biodiversity associated with the
Countries such as Brazil, Colombia, Costa Rica, Ecuador, Andes Mountains and its varied topography. Our main
Mexico, Peru, and Venezuela are considered “megadiverse objectives were to answer these four questions: 1) how
countries” because they are home to a large part of the many organizations are focused on wetlands?, 2) how are
planet’s biodiversity (Cancun Declaration 2002). Addition- they spatially distributed among the countries?, 3) what are
ally, the Neotropics support one of the greatest aquatic the principal wetland research areas?, and 4) how many
plant biodiversity and endemism in the world (Murphy Ramsar Sites were designated and how they are spatially
et al. 2019). The LAC region contains important and distributed? Through this review, we highlight the mag-
threatened ecosystems known as “biodiversity hotspots” nitude of wetland science in the Andean States and offer
including the Caribbean Islands, Atlantic Forest (Bra- support for a future wetland scientist network for LAC that
allows exchange and connection of Latin American and
1 Universidad Nacional Tecnológica de Lima Sur, Peru. Caribbean scientists regionally and internationally.
2 Universidad Científica del Sur, Peru.
3 ECOSUR (El Colegio de la Frontera Sur), Mexico.
4 Correspondence author contact: tatilobato@gmail.com.

FIGURE 1. Andean States wetlands: a) Coastal wetland, Albufera de Medio Mundo, Peru, b) Flamingos in a mangrove, San Pedro de Vice, Peru, c)
Páramo La Culata, Sierra Nevada, Mérida, Venezuela, d) Bofedal Guitarrachayoc, Peru, e) Laguna Colorada, Ramsar Site, Bolivia. Photo credits: (a, b, d)
Hector Aponte; (c, e) Tatiana Lobato de Magalhães.

FIGURE 2. Heat maps of wetland organizations distribution in the Andean States BACKGROUND
and distribution of the major Science fields studied in this region. The Andean States
The Andean States of Argentina, Bolivia, Chile, Co-
lombia, Ecuador, Peru, and Venezuela have a variety
of ecosystems and climates from 0 to 6,962 m a.s.l.,
including dry deserts, tropical rainforests, and the
snowy highlands (Guerrero et al. 2011). The varied
hydrogeomorphology across this region with the
different conditions of humidity, precipitation, and
temperature determines the characteristics of diverse
wetland types. Overall, there are five major Cow-
ardin system types of wetlands in these countries:
marine (coastal wetlands, including lagoons, rocky
shores, and coral reefs; Figure 1a), estuarine (deltas,
tidal marshes, and mangroves; Figure 1b), palustrine
(swamps, marshes, and bogs), lacustrine, and riverine
types, plus human-made wetlands. In the palustrine
type there are many highlands wetlands (e.g., pára-
mos and bofedales; Figure 1c, d), that have been
strongly affected by climate change (Junk 2013).
Data Search and Classification
To identify wetland organizations, we reviewed
proceedings of wetland scientific meetings such
as congresses, workshops, and symposia held in
countries of the Andean region from 2010 to 2020.
Due to the lack of wetland-specific scientific events
in many of these countries, we considered meetings
of closely related science fields (e.g., limnology,
ecology, and botany).
We followed these four steps:
Step 1: Identification of scientific events held
between 2010-2020 to search for organizations
focused on wetlands, except for Ecuador, which
we included a 2001 wetland event (Table 1).
Step 2: Detailed analysis of the title, abstracts,
and keywords of each article found in the
proceedings of the scientific events. This step
allowed us to filter out papers on wetland issues
FIGURE 3. Principal sectors of the Andean States Wetland Organizations. and the respective authors’ affiliated organiza-
tions.
Step 3: Production of a list of organizations that
conduct research on wetlands for each country.
Step 4: Classification of organizations through
an online search, considering the following
aspects: (a) general: name, acronym, and sector
of the organization (i.e., private, governmental,
non-profit, and academic); (b) research areas:
identified major research areas through review
of the website of each organization and scien-

tific event publications keywords; (c) location: country, In some countries, we observed a higher number of
state, city, geographical coordinates of each organiza- organizations in the country’s capital region than in the
tion; and (d) point of contact: website link, name of an interior (Figure 4), especially in Argentina and Peru. In the
associated researcher, and email address. latter country, the difference between the capital (57 organi-
zations) and the interior of the country (16) is most evident.
In addition, we listed the Ramsar Sites for each country,
On the other hand, we observed the opposite in Bolivia,
including site code and name, total area, and geographical
Chile, Colombia, and Venezuela. Bolivia did not even pres-
coordinates for each country (Ramsar 2020).
ent any organization located in the constitutional capital
Data Analysis and Mapping (City of Sucre). The highest number of organizations in Bo-
We created descriptive graphs to represent total numbers of livia was found in La Paz (13), which is the main political
organizations per sector, per each country’s regions (orga- center of the country. The largest number of organizations
nizations located in the country capital and country interior in Colombia was located in the regions of Cundinamarca
cities), and Ramsar sites for each country, using GraphPad and Antioquia, particularly in cities of Bogota (Colombia’s
Prism v. 8.4.2. We created heat maps (density maps) to capital with 13 organizations) and Medellín (11), which are
represent the spatial distribution of the organizations, major the main political and economic centers of the country. It is
research areas, and Ramsar Sites in the Andean States, us- worth noting that a similar case occurred in Ecuador where
ing the Kernel density analysis tool of the ArcGIS v. 9.3. 14 organizations were observed in the capital Quito and
To highlight the wetland research areas developed by each eight in the city of Guayaquil, located in the Guayas State,
country we created “word cloud” graphs with the keywords Southern Ecuador.
of all publications selected in Step 2 of the data search, us-
Principal Wetland Science Fields
ing an online free-tool (wordclouds.com). This tool works
We identified 13 major wetland research areas developed
with a list of words, which are displayed in different dimen-
by the 283 organizations (Table 2). The most representative
sions and randomly distributed, according to the frequency
ones in the heat maps were hydrogeology, biogeochemistry,
that each word is found in the list in question.
and biodiversity (represented by fauna, flora, algae, phyto-
FINDINGS plankton, and zooplankton). We observed a lack of wetland
Distribution of Wetland Organizations in the Andean States restoration, mapping, and management (Figure 2). Overall,
We identified 283 wetland organizations in the Andean the word cloud graphs highlighted keywords as ‘macroinver-
States, with the most located in Peru (26%) followed by tebrates’, ‘bofedales’, ‘rivers’, ‘biodiversity’, ‘conservation’,
Colombia (17%), Chile (16%), Argentina (12%), Bolivia and ‘mangroves’ (Figure 5a). The hydrology field is highly
(10%), Venezuela (10%), and Ecuador (9%) (Figure 2). The studied in South American countries (Kandus et al. 2018)
academic sector (universities) had the largest
number of organizations (176 organizations), FIGURE 4. Wetland organizations location in the country’s capital and interior of the
followed by government (51), non-profits Andean States.
(46), and private sector (10) (Figure 3). The
university sector encompasses research groups,
laboratories, institutions, museums, facul-
ties, graduate programs, research centers, and
consequently are more representative in the
number of studies presented in the scientific
events we analyzed in this review. Usually,
in developing countries the academic sector
represents a higher contribution to scientific
research than other sectors (Kumar 2017).
Some of the more important Andean universi-
ties include Pontificia Universidad Católica
del Perú, Universidad San Francisco de Quito,
Universidad Central de Venezuela, Universi-
dad Nacional de la Plata, Universidad de Los
Andes, and Pontificia Universidad Católica de
Chile (World University Ranking 2020).

TABLE 1. Proceedings of scientific meetings of the Andean States.
Country Scientific meeting Source
Argentina VI Congreso Argentino de Limnología CAL, 2014 http://sedici.unlp.edu.ar/handle/10915/69018
https://www.researchgate.net/publication/262876439_Memo-
IV Congreso Boliviano de Ecología, 2014
Bolivia rias_del_IV_Congreso_Boliviano_de_Ecologia ,
III Congreso Boliviano de Botánica, 2016
https://issuu.com/iasa-usfxch/docs/memoria_congreso_botanica ,
http://sociedadchilenadelimnologia.cl/libro-de-resumenes-xiv-
Chile XIV Congreso Sociedad Chilena de Limnología, 2017
congreso-sociedad-chilena-de-limnologia/
Colombia IX Seminario Colombiano de Limnología, 2012 https://revistas.udea.edu.co/index.php/actbio/article/view/331656
Humedales Marino - Costeros Continentales, 2001 https://biblio.flacsoandes.edu.ec/libros/digital/56577.pdf ,
Ecuador Segunda Reunión de la Iniciativa Regional para la Con- https://www.ramsar.org/sites/default/files/documents/library/
servación y Uso Racional de Manglares y Corales, 2011 informemanglarescoralesguayaquil2011.pdf
https://copehu2017.wixsite.com/copehu2017/libro-de-re-
I Congreso Peruano de Humedales, 2017,
Peru sumenes ; https://es.scribd.com/document/429166928/LIBRO-
II Congreso Peruano de Humedales, 2019
RESUMENES-COPEHU-2019-1-pdf
https://revistamipensamiento.files.wordpress.com/2015/10/
Venezuela X Congreso Venezolano de Ecología, 2012
libro-de-resumenes-x-cve-final.pdf
TABLE 2. Principal research areas of the Andean States’ organizations.

Field of Science Sub-topics
Wetland wise-use and Sustainable aquaculture, Fishery, Economic valuation, Ecological valuation, Ecosystem processes, Car-
services bon stock, Management of wetlands, Sustainable development, Ecotourism.
Focus on ecosystem Mangroves, Marine and coastal ecosystems, Estuaries, Coastal wetlands, Highland Andean bogs, “Bofe-
dales”, Peatlands, Rivers, Basin, Brackish lagoons, Bolivian Amazonia, Pantanal.
Hydrogeology Sediments, Groundwater, Geomorphology, Geological characterization, Hydraulic parameters, Reser-
voir, Hydrology, Physicochemical parameters, Hydro systems, Tropical river fluvial, Tributaries, Lake
and river water bodies, Fluvial ecosystem dynamics, Water drainage of rains, Watersheds, Hydrobiologi-
cal resources, Hydrodynamics, Management of water resources, Model ecohydrology of biodiversity,
River valleys, Lotic environments, Riversides, Pampas streams, Geological evolution.
Climate Change and Climate change, Particulate organic carbon, Adaptation to climate change, Carbon storage, Climate vari-
Carbon Storage ability.
Conservation and Biodiversity, Conservation, Endangered species, Ramsar Sites, Taxonomy.
Biodiversity
Fauna Birdlife, Ichthyofauna, Macroinvertebrates, Benthic Macroinvertebrates, Wildlife, Macrobenthos, Ento-
mofauna, Ornithology, Mastozoology, Herpetology, Bivalves.
Botany Flora, Aquatic plants, Vascular macrophyte index, Vascular flora.
Algae and Zooplankton, Biological communities, Phytoplankton composition, Algal blooms, Microalgae, Taxo-
Zoo-phytoplankton nomic composition of microalgae, Cyanobacteria, Protozoa, Perifiton, Diatoms.
Biogeochemistry Water quality, Water pollution, Physicochemical parameters.
Monitoring and Risk Environmental impact, Indicators of physical habitat, Bioindicators, Water quality, Vulnerability, Wa-
analysis ter contamination, Biological evaluation of wetlands, Species monitoring, Toxicity, Bioaccumulation,
Heavy metals, Ecohydrological indicators, Limnological status index, Microplastics.
Social and Research dissemination, Environmental education, Socio-economic evaluation, Environmental policy,
Educational Traditional management, Sustainable development.
Ecology Ecological state, Trophic state, Ecosystem processes, Lotic communities, Evapotranspiration, Plant
ecology, Phenology, Population dynamics, Eutrophication, Limnology, Migration.
Restoration, Management Reforestation and Restoration, Environmental legislation, Wetland inventory,, Territorial management,
and Mapping Remote sensing, Environmental enforcement and policy.

as well as macroinvertebrates, especially in Argentina and Ramsar wetlands are mostly distributed in the Argentinean
Colombia (Ramírez and Gutiérrez 2014). For Argentina, and Ecuadorian territory. Another large portion of these
Chile, and Colombia we observed that the principal words sites are distributed in Northern Peru and Southern Bolivia
of the word cloud graphs were ‘macroinvertebrates’, ‘algae’, (Figure 6). The majority of the Andean Ramsar Sites are
‘zooplankton’, ‘phytoplankton’, and ‘water quality’, which is beautiful and scenic places with tourism importance (e.g.,
coherent with the heat maps’ results (Figure 5 b, d, and e). the Laguna Colorada in Southern Bolivia) (Figure 1e).
Wetland biodiversity and conservation is probably the
most relevant research area for the Andean States since CONCLUSIONS AND INSIGHTS INTO WETLAND SCIENCE IN
this region harbors ecosystems such as the tropical Andes THE ANDEAN STATES
(an important ‘hotspot’ considered to be a highly diverse Even though the Andean region is extremely important for
ecosystem) and a large area of tropical forests with diverse wetland biodiversity due to its diversity of ecosystems as
wetland types (Myers et al. 2000; Junk 2013). Several orga- well as endemic flora and fauna (Junk and Piedade 2011;
nizations have been focusing on specific ecosystems such Marreno and Rodríguez 2017; Murphy et al. 2019) through-
as mangrove forests and coastal wetlands, particularly in out the region, there is a lack of wetland-focused scientific
Ecuador, Peru, and Venezuela. Despite the widely recog- events, associations, and societies. We can highlight the fol-
nized importance of the coastal wetlands and mangroves lowing wetland scientific local meetings: Argentinean Con-
for carbon storage (Hamilton et al. 2020) and protection gress of Limnology, Chilean Society of Limnology Con-
against natural disasters, they remain highly threatened gress, Colombian Seminar of Limnology, Marine Coastal
(Davidson 2014; Davidson et al. 2019). For example, Ecua- and Continental Wetlands Meeting, Regional Initiative for
dor reported 80% mangrove forest loss in the last decade as the Conservation and Wise-use of Mangroves and Corals
a result of aquaculture development (Hamilton et al. 2020). Meeting, and the Peruvian Congress of Wetlands (Table 1).
Yet, Venezuela has focused mainly on wetlands associated We carefully analyzed their proceedings, as well as some
with the Orinoco River, coastal deltas and lagoons, which local scientific events related to wetlands such as the Boliv-
have suffered high deterioration, as well as flora studies ian Congress of Ecology, Bolivian Congress of Botany, and
(Lárez and Prada 2015; Suárez 2016; Marrero and Rodrí- Venezuelan Congress of Ecology to highlight the efforts of
guez 2017). researchers involved with local organizations. The creation
Bofedales, a highland Andean wetland ecosystem is of this Andean wetland organizations database with 283 or-
highlighted by the word cloud graphs in Bolivia and Peru ganizations is a first step to recognize this wetland science
(Figure 5 c, g). This ecosystem is important due to water community and promote a LAC wetland-network.
and human-food resources, biodiversity, and for livestock This review allowed us to identify research areas that
activities as well as for climate change and carbon accumu- stand out in the Andean States such as biodiversity (mainly
lation studies (Maldonado 2014). However, the bofedales avifauna, macroinvertebrates, botany, phytoplankton, and
are one of the most vulnerable ecosystems in the world that zooplankton), ecology, hydrogeology, and water quality.
are being affected by climate change, as high temperatures As emerging areas that have the potential to be developed
will alter peatland distribution and extension (Dangles et in the Andean region, we believe research in education,
al. 2014; Huamán et al. 2020). Peru also stands out in the climate change, sustainable use, and ecosystem services
study of the use and valuation of ecosystem functions, as valuation could be expanded. Yet, we consider that res-
well as for studies of avifauna, mainly in coastal wetlands toration, mapping, management, and wetland policy are
that shelter both local and migratory birds (Rivas and Cuel- less represented in the Andean States and extra efforts are
lar 2017) (Figures 2 and 5). needed to develop and research these important issues in
the region. Additionally, identifying wetlands research
Ramsar Sites in the Andean States
areas and topics in Venezuela was a challenge due to the
All countries of the Andes are Ramsar Convention sig-
scarcity of information about wetlands on digital platforms,
natories. Overall this region harbors a total of 94 Ramsar
probably because the political scenario of the last decades,
Sites covering 300,000 km2 distributed in seven countries,
where wetland science and conservation were not a priority
Argentina (23 sites), Ecuador (19), Chile (14), Peru (13),
(Lobato-de Magalhães et al. 2016).
Bolivia (11), Colombia (nine), and Venezuela (five) (Fig-
Overall, we observed a high number of wetland orga-
ures 6 and 7) (Ramsar 2020). The first Andean Ramsar
nizations and Ramsar Sites in the Andean States, which
Site was designated in 1981 (Carlos Anwandter Sanctuary,
reinforces the countries’ government commitment to pre-
Chile), while the most recent one was designed in 2018
serving wetland biodiversity and ecosystem services. The
(Tongoy Bay Coastal Wetland, Chile) (Ramsar 2020).
observed number of wetland research organizations reflects

FIGURE 5. Word cloud graphs using the keywords from research presented on Scientific meetings’ proceedings in the Andean States:
a) Overall in Andean States, b) Argentina, c) Bolivia, d) Chile, e) Colombia, f) Ecuador, g) Peru, h) Venezuela.

the potential of scientific production in the region, FIGURE 6. Ramsar Sites distribution in the Andean States.
the human capital available to generate it, and efforts
to bring this wetland scientific-knowledge closer
to the population and decision-makers locally. This
review is an approach to understanding the potential
of the LAC region to address wetland science and
contribute to the international scientific wetland
community. It is also important that, in the coming
years, alliances are sought between research and
conservation groups of LAC and other regions (e.g.,
Global Environmental Facility “GEF Humedales”,
Wetlands International, and World Wildlife Fund).
A good example of such joint work is what is cur-
rently happening with the Coastal Wetlands Initia-
tive (humedalescosteros.org), which seeks to protect
the Pacific desert wetlands corridor and to generate
synergies between researchers and partners in Chile,
Ecuador, and Peru.
Finally, the elaboration of this review reflects the
role of the SWS in creating opportunities to integrate
scientists from diverse countries. More efforts are
needed to further wetland science and conservation
in the LAC region. However, review paper initia-
tives focused on wetland organizations and research
topics including other LAC regions can create the
appropriate scenario that will allow successful
coordination of wetland science and conservation in
the LAC region. Additionally, through an SWS-LAC
network, we could connect wetland scientists and
allow them to disseminate their efforts on wetland FIGURE 7. Ramsar Sites in the Andean States, number of sites and total area (km2).
research to other Latin American and Caribbean
scientists and internationally. n
ACKNOWLEDGMENTS
We thank D. Faust (SWS Education Section) and
D. Lobato for reviewing this manuscript. The cor-
responding author holds a postdoctoral fellowship
from ECOSUR (El Colegio de la Frontera Sur)
at CONACyT (National Council of Science and
Technology) in Mexico. We appreciate the valuable
comments of the editor R. Tiner.
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10.1007/s11747-016-0496-3

WETLAND RESEARCH
Wetlands of the Coast of Lima: Patterns of Plant Diversity and Challenges

for their Conservation
Héctor Aponte1
ABSTRACT (2013) for just six coastal wetlands of Lima reported the pres-
he wetlands of the coast of Lima are ecosystems very ence of 123 vascular plant species. When we add birds (Tello
T close to the city. This makes them particularly important
wetlands in composition and function. In recent years, mul-
and Engblom 2010), mammals (Pacheco et al. 2015), reptiles
(Icochea 1998), spiders (Paredes 2010) and protozoa (Guil-
tiple investigations have been carried out on plant diversity lén et al 2013; Guillén et al. 2015) to the plant species, more
of these wetlands. This allows us to know various aspects of than 300 species of organisms occur in Lima’s wetlands. This
their composition and structure. This article compiles the most diversity alone testifies to the importance of these ecosystems
important information on these ecosystems (mainly at the as shelters for the diversity of life on the desertlike coast of
vegetation level), trying to establish some patterns regarding Peru. How much do we know about the patterns that diver-
their diversity at the alpha, beta and gamma levels. As a result, sity of species follows in this region? And, if known, can this
we found that alpha and gamma diversity is intimately related help us make better conservation decisions?
to human activities, which triggers, for example, the presence
FIGURE 1. Map showing the location of the seven wetlands studied.
of invasive species that now represent approximately 50% of MM=Albufera de Medio Mundo, CAR=Humedal de Carquín – Hualmay;
the plants. Plant richness is independent of the state of con- PAR=Laguna El Paraíso; SR=Humedales de Santa Rosa; VEN=Humedales
servation of the areas (having protected areas of high and low de Ventanilla; PAN=Pantanos de Villa; PV=Humedales de Puerto Viejo. Scale
diversity compared to the non-protected) and size (the small- bar =100km.
est wetland is the one with the highest values of richness per
area unit). At the regional level, no patterns of beta diversity
have been found, which suggests that we should conserve each
wetland along the corridor. Some of the challenges for the con-
servation of these ecosystems are raised.
INTRODUCTION
Peru is rich in wetlands including coastal, Andean, and
Amazonian types. The country has more than 12,000 la-
goons: 3,896 in the Pacific basin, 7,441 in the Atlantic basin,
841 in the Titicaca basin and 23 in the closed basin of the
Huarmicocha System that contribute to this diversity (Min-
isterio de Agricultura, INRENA 1996). On the coast of Peru,
wetlands are part of the migratory route of birds along the
Pacific Corridor and as such are especially important for the
conservation of biodiversity. To date, 91 coastal wetlands,
56 natural, 11 artificial, and 14 river mouths are known and
of these, 25 are in Lima: 16 natural, 4 artificial and 5 river
mouths (Pronaturaleza 2010). More than 80 species of vas-
cular plants and 180 species of birds been recorded in these
wetlands by preliminary studies (Pronaturaleza 2010). More
detailed work of the different biological groups produce
more accurate figures of the diversity that is housed in these
ecosystems. For example, the work of Aponte and Cano
1 Professional Wetland Scientist. Associate Researcher, Universidad Científica del

Sur, Lima- Perú; haponte@cientifica.edu.pe.

STUDY AREA example, the alpha diversity indices (such as Shannon-Wie-
In the present study, we worked with data from seven ner Index) indicate diversity at the habitat level, while beta
wetlands of the central coast of Peru: Wetlands of Puerto diversity estimators (such as Whittaker and similarity indi-
Viejo, Pantanos de Villa, Ventanilla Wetlands, Wetlands of ces) allow us to evaluate the exchange of species between
Santa Rosa, El Paraíso Lagoon, Carquín-Hualmay Wetland sites (Halffter and Moreno 2005). Available information on
and Albuferas de Medio Mundo. They are located between alpha diversity (mean Shannon-Wiener value from different
10°58`05.15 “S and 12°34`16.77” S, and occur from sea habitats from Aponte and Ramírez 2011) and beta diversity
level to altitudes no higher than 25m (Figures 1 and 2). (Whittaker and Cody indices from Aponte 2017a, 2017b)
Most of these wetlands are associated with coastal lagoons. for these wetlands was added. To evaluate the correlation
While most of the water of these wetlands comes from the between beta diversity and distance between wetlands, the
Andes, some water enters the subsoil in the highlands and similarity between plant composition in these wetlands
flows into coastal lagoons. In many cases, this water cross- was calculated using the Jaccard index. The equations and
es urban, livestock or agricultural areas before entering the detailed interpretation of all the mentioned indices can be
subsoil and upon reaching the coast results in euthrophic found in Moreno (2001). The distance between the wetlands
ecosystems. In some other cases, it enters through tempo- was measured using Google Earth.
rary connections (directly or by the underground) between
RESULTS AND DISCUSSION
the lagoons and the sea.
Alpha and Gamma Diversity: Their Relationship with
METHODS Anthropic Processes
To answer the questions raised about diversity patterns, The data obtained to date show that alpha (habitat level)
several sources of vascular plant richness data were ex- and gamma (wetland level) diversity is high in both pro-
amined: Aponte and Cano (2013) for the wetlands of tected wetlands and non-protected wetlands (Table 1). For
Paraíso (not protected), Santa Rosa (not protected), Medio example, Los Pantanos de Villa (a protected wetland) has
Mundo (protected by the regional government) and Puerto the greatest historical richness reported to date while the
Viejo (not protected); Aponte and Ramirez (2014) for the Santa Rosa wetland (non-protected) is the second-ranked
Ventanilla wetlands (protected by the regional govern- wetland with the highest number of plant species and alpha
ment); Ramirez and Cano (2010) for Los Pantanos de Villa diversity, despite receiving minimal protection and being
(protected, Ramsar site and a National Protected Area) and located where human activities have considerably inter-
Aponte and Cano (2018) for Carquín-Hualmay Wetland vened the landscape. In the latter case, livestock seems to
(not protected). Complementary to the species richness of play an important role in increasing diversity at the alpha
each wetland (which can be an estimator of gamma diversi- level (with high-value plant communities where this activity
ty) are other measures of diversity on a different scale. For is practiced). Finding an adequate level of human activity is
essential to achieve an adequate use of space and the conse-
FIGURE 2. The lagoon and wetlands of Paraíso (Huacho, Lima). Wetlands quent conservation of the ecosystem.
of the coast of Lima are ecosystems formed by the runoff from the water One of the key challenges for the conservation of biodi-
that comes from the Andes frequently interacting with salt water brought in versity has been the need to conserve the “natural” diversity
underground from the ocean. The freshwater entering these wetlands often of ecosystems, so that not only the species found in ecosys-
crosses agricultural fields and urban areas.
tems are conserved, but also those that historically have a role
in that place (Santana 2019). To achieve this in the wetlands
of the Peruvian coast, it is essential to carry out historical
studies that allow us to understand the temporal composition
of these ecosystems, in order to identify the natural diversity
of plants while maintaining the ecosystem processes and
services that these species provide. Likewise, it is important
to identify the level of contribution that non-native species
have in the ecosystem and in the ecosystem services they
provide in order to adequately decide what steps to follow in
their management. It has been suggested that non-native spe-
cies should be considered part of the diversity of ecosystems,
at least initially (Schlaepfer 2018). So we must consider their
distribution in ecosystems such as Peruvian coastal wetlands

where non-native plants represent a considerable percentage of plant species is governed by a set of processes where
of the species in the plant community. human activities intervene. These interactions are often
The most recent study of plant diversity carried out complex and little studied. Consequently, it is not possible
in the Carquín-Hualmay wetland revealed the importance to easily make a decision on which ecosystem to protect
of small wetlands (which are often little studied) (Aponte and makes it is necessary to study the processes that occur
and Cano 2018). Although this wetland was not the most within and between these wetlands in order to make a good
diverse of those along the coast of Lima, due to its size, it conservation decision. For example, little is known about
hosts the greatest number of species per area. This makes it the role of connectivity in these ecosystems and between
one of the most important ecosystems in the region, and by these wetlands and coastal hills. Recently the presence of
this alone, worthy of conservation. Moreover, its location an aquatic plant has been reported in the lomas formations
at the center of the coast of Lima makes it vital as the con- – Lemna minuta Kunth (duckweed). This plant most likely
nection in the diversity corridor along the coast. came from a nearby wetland (Aponte 2016).
Finally, the origin of the flora of these wetlands shows The absence of patterns among these wetlands, as well as
that approximately 50% of the plants are invasive species their high complementarity, is an indicator that the best way
(see the tables in Aponte and Cano 2013 for more de- to protect them is through the creation of large-scale pro-
tails). This situation is the result of the interaction of these tected areas. There is experience of this type in Peru (i.e., the
ecosystems with adjacent populations and human activities island and guano islands protection system), so these propos-
(e.g., garbage disposal, livestock grazing, and agriculture; als are not unfamiliar to decision-makers. Currently there are
Young 1998; Ramirez et al. 2010). The presence of inva- initiatives to protect the wetlands of the Pacific coast (for ex-
sive species is now a characteristic of these ecosystems
and occurs both in protected and unprotected wetlands. FIGURE 3. Diversity of birds and plants in the coastal wetlands of Lima,
While the presence of these species is typically interpreted evaluated with the beta indices of a) Whitaker and b) Cody, and c) at the
richness level.
as a symbol of the deterioration of wetlands, it might be
interpreted it as an integral part of surviving in a human-
influenced landscape that accomplish the function as a
reservoir for urban flora. So our conservation goal must
reflect the reality of invasive species being a component of
the diversity of these wetlands, but the challenge is how to
manage them.
Regional Beta Diversity: Patterns in Plants?
The studies carried out at the beta level show that, although
birds have a greater richness of species in Lima, the turn-
over between sites (beta diversity) is greater in plants than
in birds (Figure 2). This means that the complementarity
between wetlands at the regional scale is greater in the case FIGURE 4. Scatter plot of the distance between wetlands and the similar-
of plants than in birds. Considering this indicator, it is very ity (measured with the Jaccard index) between them. The figure shows no
correlation between beta diversity and distance between sites which means
difficult to decide which wetland to conserve and which
that there is no spatial pattern of beta diversity at the regional scale. The cor-
one not to conserve, or rather where do we prioritize con- relation analysis for these variables resulted in p> 0.05.
servation initiatives? Also, by analyzing the diversity from
this second perspective (considering complementarity), we
can understand the importance of each of these wetlands
for the conservation of the region’s flora.
When analyzing the correlation between beta diversity
and distance between sites, no pattern of beta diversity
at the regional scale was found. Normally, the greater
the distance between localities, the greater the difference
between them and as a consequence there is a greater beta
diversity, but this pattern was not observed in the study area
(Figure 3). This indicates that the generalization of known
patterns for diversity is not the best route to make conserva-
tion decisions. In this case, the structure and composition

ample the Initiative for the Conservation of Coastal Wetlands for those ecosystems. In addition to these, Lima’s wet-
and Shorebirds in the Arid Coast of the South American lands have several common characteristics (e.g., proximity
Pacific), where these proposals would fit perfectly. to the city, land ownership or management problems in
Challenges for the Conservation of the Coastal perimeter areas, presence of human activities for the use of
Wetlands of Lima its resources, absence of permanent monitoring programs
The study of diversity patterns identified research needs for both biotic and abiotic components, and ignorance of
that are important to make adequate conservation decisions. the existence and importance of these ecosystems by the
There are small wetlands that have not yet been studied general public) that have allowed us to identify situations
(e.g., Laguna la Encantada) and whose importance as requiring immediate attention for conservation (Aponte
shelters for diversity should not be underestimated. Like- 2017c; Aponte et al. 2018). At least three activities have
wise, the role of invasive plant species (considering that, been identified as fundamental for the protection of these
in some cases, they represent around 50% of the species) ecosystems in the long term:
in these wetlands is hitherto unknown, so it is necessary to • Legally recover the spaces. Many of the areas have
study them before indicating that they are harmful species owners who could legally eliminate the wetland.
NGOs and the state must work to reverse the land
TABLE 1. Plant mean alpha diversity, historical gamma diversity, area and tenure processes and be able to properly manage
number of species per area for the studied wetlands. Protected wetlands these spaces. This will require public education and
are marked with a * outreach and dialogue with the owners to help them
understand the environmental value of the wetlands
Wetland Mean Gamma Area Species/ and how use of these lands can be done in a way to
Alpha (ha) area minimize adverse effects.
Puerto Viejo 0.32 32 200 0.16
• Media movement. It is necessary to involve local
Pantanos de media (radio, television) and social networks to help
Villa* - 72 276 0.26 improve the public’s understanding and appreciation
Ventanilla* - 37 265 0.14 of these wetlands. Surprisingly, many people who
Santa Rosa 0.72 67 60 1.12 live in Lima do not know about the existence of these
Carquín-Hualmay - 41 11.64 3.52 ecosystems. This outreach must include information
(e.g., pamphlets and videos) and education through
Paraíso 1.05 33 690.42 0.05
field trips by school students and encouraging uni-
Medio Mundo* 0.48 21 261 0.08 versities to use wetlands as a subject for ecosystem
FIGURE 5. A pelican in the coastal wetland Poza de la Arenilla (La Punta,
research.
Callao), an artificial wetland that is enriched by nutrients (probably from • Working together. It is essential that all people
urban origin plus natural organic matter) causing the growth of thick involved in the protection of these wetlands work
mats of green algae (in the background). Controlling the growth of these together. In recent years there have been several
algae and the sources of eutrophication is very important, and will only personal and isolated initiatives, but I think it is more
be achieved with research and appropriate land management. To date,
no studies of the flora of this wetland have been published. productive to work cooperatively on outreach activi-
ties. Environmental professionals need to work with
economists, lawyers, sociologists and communica-
tors on improving public awareness of wetlands and
conservation needs. In this way, the fruits of inter-
ventions and research can be analyzed from various
perspectives, considering possible gaps in the design
of management and management plans that allow for
solutions (Figure 5).
Along with the lomas formations where we also find
hundreds of species (Dillon et al. 2011), wetlands are an
important refuge for the flora and fauna of this region.
Due to their proximity to human populations, both lomas
formation and wetland ecosystems have suffered reduc-
tions in their extent and degradation by various impacts.

Therefore, management plans must be applied in an inte- Guillén, G., E. Morales, and R. Severino. 2013. Adiciones a la fauna de
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Biología 10:175–182. doi: 10.15381/rpb.v10i2.2500
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achieved thanks to the support of the Museum of Natural SEA, CONABIO, Grupo DIVERSITAS & CONACYT, Zaragoza. pp.
History (Universidad Nacional Mayor de San Marcos), the 5 - 8.
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azados del departamento de Lima. Los Pantanos de Villa: Biología y
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grateful. Conservación de Humedales en el Perú.
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WETLAND RESEARCH
Andes, Bofedales, and the Communities of Huascarán National Park, Peru

Rodney A Chimner1, Randall Boone2, Gillian Bowser2, Laura Bourgeau-Chavez3, Beatriz Fuentealba4, Jessica Gilbert5,
Javier A. Ñaupari6, Molly H. Polk7, Sigrid Resh1, Cecilia Turin8, Kenneth R. Young7, and Melody Zarria-Samanamud2
ABSTRACT INTRODUCTION
ountain wetlands are abundant in the high elevations The tropical Andes are rich in biological diversity and have
M of the tropical Andes. Wetlands occupy ~11% of the
total park area and are mostly found in the large mountain
been utilized for millennia by human communities (Young
and Lipton 2006). When the Spanish conquistadors arrived
valleys. Wetlands occur up to 5000 m asl, but most occur in the 16th Century, bringing horses and cattle and a differ-
between 4,000–4,700 m asl. The highest elevation wetlands ent land management scheme with them, cultural resources
are typically dominated by cushion plants, while lower dramatically changed and many unique ecosystems were
elevation wetlands are more commonly occupied by grami- transformed by the introduction of new grazers and brows-
noids. About 60% of all wetlands are peatlands and the ers and the abandonment of maintenance of water systems
remainder are mineral soil wet meadows. The peatlands are and much of the original land use practices shifted and
up to 11 m deep and 12,000 years old, storing an average of adapted. Dramatic transformations are once again occurring
2,101 Mg C ha-1, which is comparable to lowland tropical with rapid climate change and glacial retreat in the Andes
peatlands. Our work in Huascarán National Park in Peru of Peru that is further transforming water and biological re-
is also showing the importance of wetlands in a coupled sources, changing biodiversity as species shift, livelihoods
natural-human system. These wetlands and alpine land- adapt by changing agriculture, livestock husbandry, and
scapes are shaped in part by legacies of past human land other economic activities, and new species are introduced
use, including ancient pastoralism and farming, and are also (Anderson et al. 2011; IPCC 2019; Hock et al. 2019). In
affected by millions of downstream users dependent upon Peru, these problems are exacerbated by growing demands
wetlands and glacier-fed streams for water and energy pro- for water and other mountain natural resources associated
duction. Biodiversity and endemism is high among taxo- with economic growth (Mark et al. 2017; Hock et al. 2019).
nomic groups such as plants, birds, fish, amphibians and The high Andes of Peru have received worldwide atten-
insects. Currently the tropical Andes are in ecological flux tion due to rapid glacial retreat and socioeconomic changes
due to rapid land cover changes caused by both biophysi- to the traditional Andean societies. These changes could
cal and socioeconomic drivers. In addition, the high Andes cause the collapse of traditional societies altering human
are experiencing warming and rapid glacial retreat that is well-being and cultural heritage, coupled with ecological
resulting in hydroecological changes and socioeconomic collapse in one of the most biologically diverse landscapes
changes to the traditional Andean societies that feed back to in the world (IPCC 2019). Such dramatic shifts may be
changes in wetland sustainability. mirrored in other ecological systems and traditional societ-
ies throughout the world especially where wild pollination
is tied to traditional food crops and small-scale agricultural
1 College of Forest Resources and Environmental Science, Michigan Techno-
logical University, Houghton, MI, USA; Corresponding author: rchimner@ systems (IPBES 2018; Villagra et al. 2020). A multidisci-
mtu.edu. plinary research team including U.S. and Peruvian institu-
2 Department of Ecosystem Science and Sustainability, Colorado State Uni-
tions, Peruvian National Park managers, agropastoralist
versity, Fort Collins, Colorado, USA.
3 Michigan Tech Research Institute, Michigan Technological Univer- communities and a local non-governmental organization
sity, Ann Arbor, Michigan, USA. with funding from National Science Foundation-Coupled
4 Instituto Nacional de Investigación en Glaciares y Ecosistemas de Natural Human Systems are examining these issues in
Montaña, Huaraz, Ancash, Peru. Huascarán National Park (HNP) as a case study of how
5 Department of Wildlife and Fisheries Sciences, Texas A&M Uni-
versity, College Station, TX, USA. communities, biodiversity, protected area management, and
6 Animal Production Department, Universidad Nacional Agraria La rapid climate change all intersect in a wetland rich moun-
Molina. tain environment.
7 Department of Geography and the Environment, University of
Texas at Austin, Austin, Texas, USA.
8 Instituto de Montaña. Lima, Peru.

Natural Protected Areas (NPAs) provide critical envi- Bofedales (singular bofedal) is a commonly used term
ronmental services in an increasingly anthropogenic world. in Peru to refer to mountain wetlands (Maldonado Fonken
High elevation NPAs protect both the alpine species and 2014). One of the unique aspects of bofedales in the Andes
essential watersheds needed by local communities within is the dominance of cushion plants (Figure 1). The compact
and below alpine systems. Because the tropical Andes is growth form of cushion plants is an adaptation to arctic and
one of the global biodiversity hotspots (Myers et al. 2000) alpine conditions as it traps heat and minimizes wind shear
and high elevation endemism among taxonomic groups (Billings and Mooney 1968). In the Andes, cushion plants
such as plants, birds, fish, amphibians and insects, a series are mostly in the family Juncaceae, Asteraceae and Plantag-
of protected areas were created to attempt to conserve that inaceae (Cooper et al. 2010).
biodiversity (Rodríguez and Young 2000). HNP (340,000
ha) was created in 1975 to conserve the ecosys-
tems of the Cordillera Blanca, which is in the FIGURE 1. High elevation cushion plant peatland in Pastoruri, HNP. (Photo by Rod
central Andes of Peru with high peaks ranging Chimner.)
from 5,000 to 6,768 masl (including Peru’s high-
est peak, Huascarán Sur). The United Nations
Educational, Scientific and Cultural Organization
declared HNP a Biosphere Reserve in 1977 and
included it in the list of Natural Heritage of Hu-
manity in 1985. As of 2016, the Cordillera Blanca
contained approximately 755 glaciers (25% of all
tropical glaciers) and 830 lakes of glacial origin
(Mark et al. 2010; Autoridad Nacional del Agua
2016). These mountain glaciers are an important
source of water, and they regulate water quantity
by buffering the temporal precipitation variabil-
ity that supplies water for domestic, agricultural
and industrial uses during the dry season for both
locally and in the arid coast of Peru (Rabatel et
al. 2013). HNP is surrounded by 30 agropastoral-
ist communities that existed before park creation.
Cattle and sheep are numerous and free rang-
ing in HNP, with limited numbers of horses and
FIGURE 2. Cattle and horses graze in cushion plant wet meadows and peatlands in Ulta
burros allowed to graze in peripheral park areas. Valley, HNP. (Photo by Rod Chimner.)
Some communities continue to have access rights
to HNP land, and their animals can use grass-
lands and wetlands under specific agreements
and arrangements with the HNP.
WETLANDS
Mountains are often areas of high wetland abun-
dance due to excess water from high rates of oro-
graphic precipitation, and these wetlands provide
many benefits, including high-quality habitat, nutri-
ent sinks and transformations, carbon and water
storage, and areas for pasture (Chimner et al. 2010,
Cooper et al. 2012). The Andes are no exception,
with wetlands common across the entire range from
the northern páramo region of Colombia, Venezu-
ela, and Ecuador through the humid and dry puna
of Peru and Bolivia all the way down to southern
Argentina (Chimner et al. 2011).

Cushion plant dominated bofedales are common in similar (Figure 2), there can be confusion in the literature
HNP, and many of them are organic soil peatlands or fens and with management as they are both called bofedales
(Figure 1). However, other cushion plant bofedales are ac- (Chimner et al. 2019).
tually mineral soil wet meadows (Figure 2: Chimner et al. While cushion plants are common, there are many
2019). Cushion plant peatlands can have high biodiversity other plant species found in wetlands. The most recent flo-
for species assemblages of endemics such as anurans (Les- ristic analysis that focused on bofedales in HNP document-
cano et al. 2020) and wet meadows often support mixed ed relatively low alpha diversity, but potentially high beta
avian flocks that include rare and new-to science species diversity. Polk et al. (2019) sampled three valleys (Llanga-
(Schulenberg et al. 2020). Because cushion plant-domi- nuco, Quilcayhuanca, and Carhuascancha) and identified
nated wet meadows and peatlands can look superficially 112 vascular plant species in 29 families. The most species
rich families were Poaceae, Asteraceae, and Cyperaceae.
The most frequent species were Plantago
FIGURE 3. Graminoid peatlands and graminoid wet meadows in Rio Negro, HNP. (Photo tubulosa Decne., Eleocharis albibracteata
by Rod Chimner.)
Nees & Meyen ex Kunth, Juncus ebracteatus
E. Mey., Gentiana sedifolia Kunth, and Cala-
magrostis rigescens (J. Presl) Scribn. Rarity
was common in the inventory - 37% of all spe-
cies occurred only once. Mean alpha diversity
was 12.6, which is what might be expected for
high mountain vegetation. The survey design
used by Polk et al. (2019) allowed for a valley-
to-valley analysis, which showed that vegeta-
tion in the three valleys are more dissimilar
than they are similar, sharing only 35% of the
species on average. This finding suggests that
there is high beta diversity among plant species
in bofedales in HNP and more vegetation sur-
veys should be completed to further document
plant diversity.
Previous research identified four wetland
types based on hydrophytic plant species that
co-occur in Peruvian bofedales (Weberbauer
FIGURE 4. High elevation clear water lake in Rio Negro, HNP. (Photo by Rod Chimner.) 1945; Maldonado Fonkén 2014). Polk et al.
(2019) identified five plant assemblages or
groups with one cushion plant-dominated as-
semblage: 1) Plantago tubulosa - Oreobolus
obtusangulus, and four graminiod dominated
assemblages: 2) Werneria pygmaea - Per-
nettya prostrata, 3) Juncus ebracteatus - Carex
bonplandii, 4) Eleocharis albibracteata -
Calamagrostis rigescens - Lachemilla pin-
nata, and 5) Werneria nubigena - Oritrophium
limnophilum - Phlegmariurus crassus These
five groups could be conceived of as subgroups
of the system published previously by Weber-
bauer (1945) and Maldonado Fonken (2014).
Cation exchange capacity, organic matter, bulk
density, and elevation were factors associated
with structuring bofedal vegetation in the three
valleys sampled. At the scale of HNP, subse-
quent mapping work by Chimner et al. (2019)

showed that elevation and latitude are additional FIGURE 5. Interdisciplinary team in the field with Peruvian and US students and local
organizing factors and they classified bofedales partners in Ulta, HNP. (Photo by Molly Polk.)
as cushion plant peatlands, cushion plant wet
meadows (Figure 2), graminoid peatlands (Figure
3), and graminoid wet meadows (Figure 3).
There is normally little information about
mountain wetland abundance, because they are
often small and located in remote and rugged set-
tings that make them difficult to map (Bourgeau‐
Chavez et al. 2017). Mapping work by Chimner
et al. (2019) in HNP has found that wetlands are
numerous and occupy ~11% of the total HNP
area, mostly found in the large mountain val-
leys. For context, grasslands comprise 17% and
woodlands 8% of the HNP area. Cushion plant
peatlands were the most abundant wetland type
occupying 6.2% of HNP, followed by graminoid
wet meadows (3.5%) and cushion wet meadows
(1.3%), and graminoid peatlands (0.1%).
Wetland type and abundance varied with el-
evation. Wetlands occurred up to 5,000 masl, but
were most abundant between 4,000–4,700 masl FIGURE 6. Research team collaborating with community members at The Mountain
with about three-quarters of all mapped wet- Institute in Huaraz, Peru. (Photo by Molly Polk.)
land area occurring within this elevation zone.
At the lower elevations, wetlands were mostly
graminoid wet meadows and at higher elevations
wetlands were mostly cushion plant peatlands.
Cushion plant wet meadows were most common
in mid-elevations. Wetland type and abundance
also varied north to south, with more wetlands
found in the south (mostly cushion plant peat-
lands) compared to the northern part of the park
(cushion wet meadows) (Chimner et al. 2019).
Wet meadows and peatlands have large
differences in soils, hydrology, chemistry, and
often respond very differently to grazing. Wet
meadows are often preferred for grazing because
they are seasonally wet and have stable soils
compared with peatlands, which makes it easy
for livestock to utilize. Wet meadows can still be
overgrazed leading to vegetation loss, erosion,
nutrient losses and soil carbon declines (Enriquez
et al. 2014, 2015). In contrast to wet meadows, mountain of native species with abundant litter, high soil infiltration
peatlands are very susceptible to grazing impacts due to and water quality, high plant diversity, no bare soil, and no
their thick and soft organic soils, which are easily trampled sign of erosion. Whereas bofedales in poor condition had <
(Chimner et al. 2010, 2011; Cooper et al. 2012) and can 25% of native species, poor soil infiltration and water qual-
significantly reduce or reverse carbon storage and increase ity, high percent cover of bare soil, and signs of erosion.
emissions of the potent greenhouse gas methane by an Peatlands in HNP range in age from 3,200 to 12,000
order of magnitude (Sánchez et al. 2017). In HNP, Calvo years (Hribljan unpublished data) and have an average
(2016) assessed bofedal condition in Quilcayhuanca Valley depth of ~ 5 m, with maximum depths reaching ~11 m
and found that bofedales in good condition had >70% cover (Hribljan unpublished data; Chimner unpublished data).

The thick peat deposits combined with dense peat (aver- Biodiversity research has been limited to a few local-
age bulk density = 0.26 g cm-3) result in these peatlands ized studies (Fjeldså 2002; Yensen and Tarifa 2002; Lloyd
containing very high belowground carbon stocks (average and Marsden 2008; Renison et al. 2018; IPCC 2019). The
= 2,101 Mg C ha-1) compared to most peatlands globally. lack of biodiversity assessments has left gaps in under-
For instance, high elevation peatlands in HNP have carbon standing how change and disturbance will affect the bio-
stocks on an areal basis that are greater than average carbon logical communities that inhabit these systems in the future
stocks of 1,421 Mg C ha-1 from peatlands in the Peruvian (Fjeldså 2002; Lloyd and Marsden 2008; Dangles et al.
Amazon Pastaza-Marañon basin, which is the second larg- 2020). For instance, macroinvertebrates may have reduced
est continuous peatland complex in the tropics (Lähteenoja functional diversity as glaciers retreat in high elevation
et al. 2012). systems. Most wild pollinators are poorly studied with little
Although little water table data exists, peatlands in known about high elevation bombus species, which are of
HNP appear to be similar to other peatlands in that they particular concern due to the introduction of non-native
require continuous high water tables to slow decomposition bombus species such as Bombus terralis. The impacts of
rates (Planas-Clark et al. 2020). For example, even though non-native bombus introductions may be linked to the
there were large differences in total precipitation between declines of the IUCN listed species such as B. dahlbomii,
the wet and dry seasons, water table levels were relatively one of the largest bombus species (Singer and Sanguinetti
stable throughout the year and stayed near the soil surface 2014), and the Andean Bumble bee (B. coccineus). In ad-
in reference peatlands (Planas-Clark et al. 2020). This dition, several species of honey producing bees such as
stability of water also provides fresh water during the long Apis mellifera have been introduced in many high elevation
dry season for local agropastoralists (Maldonado Fonkén communities, leading to the introduction of parasites and
2014). In addition to providing water, wetlands in HNP are pathogens that appear to impact native pollinators especial-
also being used to improve water quality from heavy metal ly, the rarer bombus species such as B. dahlbomii (deLanda
pollution from mines and glacial melting exposing geologic et al 2020).
formations that are weathering metals.
WETLANDS AS COUPLED NATURAL AND HUMAN SYSTEMS
BIODIVERSITY Wetlands of the high Andes Mountains can be conceptu-
The high Andean landscape is considered one of the alized as coupled natural-human systems because their
world’s hotspots of biodiversity but much of that diver- characteristics and dynamics are only in part regulated by
sity is largely unknown and includes taxa that provide biophysical processes pertaining to the water cycle and
critical ecosystem services. High elevation wetlands and to the dynamics of ecological succession. An important
clear water lakes (Figure 4) have high diversity levels of additional component is the conspicuous human dimen-
aquatic macroinvertebrates (Nieto et al. 2020), endemic sion, such that wetland ecosystems cannot be understood
avian species (Schulenbuerg et al. 2020), amphibians without reference to interactions among the various bio-
(Lescano et al. 2020), and wild pollinators (IPBES 2018). physical and social processes. For instance, many Andean
Endemic species are still poorly known with new species bofedales have been managed since pre-hispanic times by
of birds (Schulenberg et al. 2020) and insects still be- using water management technology (Morlon et al. 1996;
ing discovered (Figure 5). On the landscape level, HNP Lane 2006, 2014).
provide habitat to over 780 vertebrate species, many of In the case of our study areas this is true even for the
which are considered threatened or near threatened by most remote and “pristine” parts of HNP. The human influ-
the IUCN Red List of Threatened Species, including the ences are ancient: one of our study sites is located along a
larger animals like the puma (Puma concolor), Andean hiking trail used by trekkers going to the Chavin archaeo-
fox (Lycalopex culpaeus), and Andean bear (Tremarctos logical site for tourism - they move along routes where
ornatus) (Fjeldså 1993; Yensen and Tarifa 2002; Lloyd pilgrims came to participate in rituals 3,500 years ago. Ag-
2008; Gareca et al. 2010). Many of these species are ropastoralists have also created and modified wetlands for
impacted by current park management. For example, the livestock by diverting surface water to maintain or enlarge
South American deer or taruka (Hippocamelus antisensis), wetlands (Lane 2006). Similarly, farmers use canals to
a high altitude specialist listed as vulnerable by the IUCN extend the irrigation of their crops. The sizes of these water
Red List of Threatened Species (Barrio et al. 2017), shares reservoirs are 10 - 15 m in diameter and hold between 100 -
habitat with domestic livestock leading to the displace- 300 m3 of water (Lane 2014). The expansion of the wetland
ment of taruka (Merkt 1987; Barrio 1999, 2006; Gazzolo area not only improves the water provision service, but also
and Barrio 2016). regulates water flow during the dry season, increases water

quality, and contributes to soil carbon sequestration (Lane managers, and as negotiated based on past land tenure. Our
2014). Understanding the deep timeline of human influ- research suggests that the park-people interactions involve
ences will be an important complement for understanding two archetypes of land use: a seasonal rotational livestock
the complex couplings in this system. system that brings in cattle to the wetlands and sheep to the
Inside the park today, it is not uncommon for cattle to hillslopes of the park during the dry season, and a system
navigate steep slopes above 4,000 m elevation. Livestock that keeps cattle in the park year-round. The former appears
are far more visible in HNP than large wild mammal spe- to be less environmentally damaging (and more productive
cies, which are more commonly seen in landscapes outside economically), but the latter is still common because cattle
HNP boundaries. The study of the pastoral systems not only owners who remove their livestock risk losing access rights
provides needed information for park managers hoping in the future. The wetlands thus are viewed as critical dry
to lessen damaging environmental impacts, but provides season production areas in some valleys, but can be sources
information to local communities who rely on wetlands for of mortality for cattle that free range year-round, with only
their economic and subsistence needs (Figures 6 and 7). weekly or biweekly visits by the livestock owners. Our
This human-nature interaction exemplifies the historical research has included detailed informant interviews by
cultural exchange between Western and Andean cultures, as an anthropologist fluent in Quechua, the first language of
they borrowed cattle from Western culture and incorporated many local people, to get their perspectives on the costs
and transformed their Andean human landscape. Cattle are and benefits of wetlands for their livelihoods.
so integrated into the local cultures now that local commu- It is impossible to do research in this part of Peru and
nities cannot understand life in the mountains without these not be struck by social injustices having to do with past
animals. For local people, cattle are part of their natural environmental and land tenure concerns. The Santa River,
landscape. It is like this in many parts of the Andes, how- fed by glaciers within the national park, is littered by
ever, there are still some remaining places where original unmitigated mine tailings and contaminated with heavy
livestock (camelids) are the main element of the landscape. metals. More than half of the high elevations of this part
Areas within HNP has been heavily used for tours by of Peru are within exploratory mining claims and several
school groups; the view of glaciers and other natural fea- large mines are in active exploitation. The park provides
tures was often the most direct experience Peruvian school an important protected space for natural environments, but
children had of nature as seen in real life rather than on
television. The glaciers pro-
vide critical runoff, especially FIGURE 7. Project leaders in the field with community partners in Rio Negro, HNP. (Photo by Molly Polk.)
during the five month long dry
season, for maintaining surface
water flows and for drinking
water for the city of Huaraz.
So, the park provides numer-
ous educational and ecosystem
services locally, regionally, and
nationally. The park’s wetlands
provide unique biodiversity
habitat, function in ways that
store water and carbon, and are
conspicuous landscape features
visible to park visitors.
There are also important
social roles filled by HNP, oc-
cupying the headwaters of val-
leys used by farmers for their
crops downslope. The high
elevations within HNP are ac-
cessible to traditional land use
by the park’s neighbors through
agreements made with park

must also accommodate legitimate concerns about land example). Mountain wetlands are most directly affected by
tenure that date back to pre-park years when lands were their hydroperiod, set by surface and groundwater hydrol-
controlled by large landowners in the form of haciendas. ogy, overlying biodiversity and ultimately by rainfall and
Some of the concerns mentioned by local people predate runoff from glaciers in glaciated watersheds. Because of
the park, but still drive emotions and actions today. their importance for ecosystem services, they are often
HNP is emblematic of global change impacts. Its considered valuable by local people, who often utilize wet-
boundaries include the longest extension of glaciers lands in their pastoralism systems, or by conservationists
anywhere in the tropics. Glacier retreat is about 30% on as keystone ecosystems with high endemism in protected
average but on the ground, there are some places where areas such as national parks. Yet most of the ecosystem
the valley-head ice is already gone, meaning that the val- characteristics unique to the high elevation remain un-
ley downslope is now “postglacial,” with hydrology now known, such as rates of endemism in high elevation birds,
regulated by precipitation and groundwater, and no longer shifts in anuran composition in wetlands and the decline
supplemented by glacial runoff. In valleys under the highest of wild pollinators such as Andean bombus species. The
peaks, the ice is predicted to last for at least another cen- number of endemic species such as wild pollinators or birds
tury. So, glacial retreat is adding additional heterogeneity as is threatened by shifts in the wetlands systems coupled with
to how wetlands function, and implications for human land the introduction of invasive pollinator species as a second-
use also vary from valley to valley depending on the size ary stressor. The lack of knowledge of species diversity and
and height of glaciers’ upslope. We expect to find that this overall endemism is profound especially as high elevation
heterogeneity will be an important part of explanations for habitat patches shift along with local livelihoods - a perfect
how and why wetlands change from place to place. storm for the collapse of many ecosystem services appears
The use of a coupled agent-based model (DECUMA) to be brewing yet unseen in research inquiry.
and ecosystem model (L-Range model) is being used The monitoring of glacial retreat, stream discharge, and
to help us understand the complexities of these coupled depth to water table would all provide insights into controls
natural-human interactions (Boone and Galvin 2014; Boone on hydroperiod. The monitoring of the extent and spatial
et al. 2018). We are simulating how landscape features such configuration of the wet meadows and peatlands can be
as topography, wetland location and cover, primary produc- done through mapping with remote sensing (Chimner at
tion, and the location of water sources influence land-use al. 2019). In particular, our additional use of active remote
decisions of agropastoral families. The coupled model is sensing data through satellite-borne radar can map both sur-
also being used to analyze scenarios of climate change and face and subsurface features. Vegetation shifts of interest in
their impacts on the landscape, wetland sustainability, and those wetlands include the shift from dominance by grami-
households and communities’ actions. Management sce- noids at lower elevations to cushion plants nearer the peaks.
narios such as changes in livestock population and rotation The human dimensions involved are multifaceted
of grazing lands will be established using a participatory given that individual valleys in the park have different
approach. The use of a coupled-modeling simulation ap- histories on land uses that are permitted, and especially in
proach will provide us information about the role of natural regards to the degree of control that park neighbors have
and human drivers on the ecosystem, decision-making (or do not have) in access rights that allow them to use
processes, and explore the impact of short vs. long-term park lands for livestock grazing and firewood collection.
management options. Since wetlands are key ecosystems in Some valleys have virtually no human impact, except for
this landscape, the model will provide insights of their role the passing of occasional hikers, while others are over-
for land and livestock management, as well as information grazed with extensive trampling, an especially damaging
of how these ecosystems could be impacted due to climate impact on wetlands. The monitoring and evaluation of
change and management decisions. these anthropogenic influences is hence more complicated
and may require nuances in mediating park-people inter-
FUTURE ISSUES actions that may in some cases be antagonistic or may
Mountain wetlands act as important sentinels of global have histories of such antagonism.
changes, mediated by shifts in climate, associated ecologi- Although COVID-19 was not a consideration when
cal/hydrological alterations, and a suite of interactions with we developed our research plans for the park, now it is
people. The global pandemic highlighted the intersections and will require us to think about wetlands, socioeconomic
between large-scale landscape changes due to climate drivers such as tourism, livestock and biodiversity in an
conditions coupled with changes in socioeconomic stability entirely new context. For instance, COVID-19 has sharply
for local communities (loss of tourism due to COVID for reduced tourism and much of the regional interchanges of

highland agricultural products for consumer goods from the Calvo 2016. Marco conceptual y metodológico para estimar el estado
de salud de bofedales de alta montaña. Tesis de maestría Universidad
coastal cities of Peru. At HNP, the local communities have Nacional Agraria La Molina
problems with their incomes since they do not have oppor-
Chimner R.A., D.J. Cooper and J.M. Lemly. 2010. Mountain fen distri-
tunities to provide guiding services, transport, hostels, and bution, types and restoration priorities, San Juan Mountains, Colorado,
food, as well as to sell souvenirs. New approaches to food USA. Wetlands 30: 763-771.
security, such as the localized production of honey, which Chimner, R.A., G.L. Bonvissuto, M.V. Cremona, J.J. Gaitan, and C.R.
has increased 100-fold in recent years, may also increase Lopez. 2011. Ecohydrological conditions of wetlands along a precipita-
tion gradient in Patagonia, Argentina. Ecologia Austral 21: 329-337.
due to the pandemic, modifying the current land manage-
ment. The local economy has been decreasing and presents Chimner R.A., L.L. Bourgeau-Chavez, S. Grelik, J.A. Hribljan, A. Maria
Planas Clarke, M.H. Polk, E.A. Lilleskov, and B. Fuentealba. 2019.
an economic slowdown. The infrastructure for communica- Mapping extent and types of wetlands in the Cordillera Blanca, Peru.
tions, drinking water, and public health is especially poor Wetlands 39: 1057-1067.
in smaller and more isolated settlements, perhaps mean- Cooper, D.J., E.C. Wolf, C. Colson, W. Vering, A. Granda, and M.
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more important for the people living as neighbors to the Antarctic, and Alpine Research 42: 19–33.
park. We do not know what this will mean for park-people Cooper, D.J., R.A. Chimner, and D.M. Merritt. 2012. Western Mountain
Wetlands. In: D.P. Batzer and A.H. Baldwin (eds.). Wetland Habitats of
relations, and in particular for the ongoing dynamism of North America: Ecology and Conservation Concerns. University of Cali-
the mountain wetlands, but suggest that it be an important fornia Press, Berkeley, CA. pp. 313–328.
focus of monitoring for the next decade. n Dangles, O., C. Ibarra, R. Espinosa, P. Andino, V. Crespo‐Pérez, D.
Jacobsen, and S. Cauvy‐Fraunié. 2020. Functional structure and diversity
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WETLAND RESEARCH
Peatlands of the Central Andes Puna, South America

Eduardo Oyague1 and David J. Cooper2
ABSTRACT the world’s most biodiverse region, and on the west by

he Andean puna is the highest elevation mountain and the Atacama Desert, the world’s driest region. The region
T plateau region in the western hemisphere, extending
from northern Peru to central Chile and Argentina, with
has complex geodynamic and volcanic processes geologic
origin (Strecker et al. 2009) Above 3500 meters asl, these
vast areas above 4,250 m elevation. The Atacama Desert geodynamic factors have combined with glacial processes
on the western side of the Andes is the world’s most arid to shape the landscape.
region, and this aridity extends across the mountains. The The central Andes alpine region is termed “Puna”,
upland vegetation is mostly continuous in the more hu- a treeless windswept area of the higher Andes, a word
mid north but becomes more and more patchy and barren derived from the Quechua language. Its climate is cold and
composed only of bunch grasses in the central and southern
region. Extensive upland talus, moraines, and colluvial FIGURE 1. Location of Central Andes Puna ecoregions in western South
deposits of the mountains produces perennial ground water America (Olson et al. 2001). A and B: Current study areas in central and
flow systems that support thousands of peatlands and southern Peru. A: Nor Yauyos – Cochas Landscape Reserve, Cordillera Cen-
other wetlands that are regionally termed “bofedales” or tral. B: Vilacota – Maure Regional Conservation Area, Cordillera del Barroso.
“vegas”. The peatlands are dominated by several species
in the family Juncaceae, most famously Distichia muscoi-
des, that forms dense clonal cushions that characterize the
alpine peatlands and landscapes. Being close to the equa-
tor, the growing season extends across the entire year, and
continuous growth of the cushion plants have produced
high carbon accumulation rates and peat bodies 7-10 m
thick in many areas. The region supports unique plants and
animals, and ecosystems, and the wetlands are threatened
by overuse for livestock grazing, and climate changes that
could reduce water provision may lead to some bofedales
completely drying.
INTRODUCTION
The Andes form the world’s longest mountain system
extending more than 7000 km along the entire western edge
of South America, and continuing the American Cordillera
from the Arctic to the Antarctic. The mountain region from
central Peru through Bolivia, northern Chile and Argen-
tina is the highest alpine region in the western hemisphere
with many peaks over 6000 meters and extensive plateaus
at 3200 to 5000 meters asl. The only region on Earth with
similar extensive high elevation is the Tibetan Plateau. The
central Andes is bordered on the east by the Amazon basin,
1 Universidad Nacional Jorge Basadre Grohmann, Tacna, Peru; Corresponding

author: eoyaguep@unjbg.edu.pe
2 Department of Forest and Rangeland Stewardship, Colorado State University,
Fort Collins, Colorado, USA.

FIGURE 2. Expansive bofedale in the highlands of southern Peru, at 4,500 m elevation. Surrounding peaks all 5200-5500 m high. (Photo by D.J. Cooper.)
semi-arid, with annual precipitation ranging from 200 to elevation forest types, for example on Nevado Sajama, the
700 mm per year. It is influenced by the movement of wet highest mountain in Bolivia. On the eastern edge of the An-
air masses from the Amazon, the temperature and currents des cloud forest forms the upper treeline. Many wetlands
in the Pacific Ocean, and the presence of Lake Titicaca in the valleys and basins are supported by groundwater dis-
(Falvey and Garreaud 2005)precipitation over the high- charge and are peat accumulating fens (Cooper et al. 2010,
altitude plateau of the South American Altiplano exhibits a 2019). Those peatlands are termed ‘bofedales’ in Peru,
marked intraseasonal variability which has been associated Bolivia and northern Chile, and ‘vegas’ in central Chile and
with alternating moist and dry conditions observed at sur- Argentina (Rojo et al. 2019) (Figure 2). They have some of
face stations near the Altiplano western cordillera. In this the most rapid known rates of peat accumulation, up to 1.4
study the characteristics of humid (wet. It is well known for - 2.2 mm/yr and a long term carbon accumulation rate of
having “summer every day and winter every night”, with 37 - 47 g C/m2/yr (Hribljan et al. 2015). Carbon storage is a
freezing or near freezing temperatures in all seasons (Run- key ecosystem service provided by bofedales, but they also
del et al. 1994). The puna is climatically variable, with “wet represent important refugia for many organisms, particu-
puna” in central Peru, “puna” in southern Peru and northern larly during the prolonged dry seasons.
Argentina, and “dry puna” on the Peruvian-Bolivian Pla- Bofedales are critical pastures for indigenous pastoral
teau called “altiplano” and in northern Chile (Figure 1). communities (Yager et al. 2019) whose inhabitants live at
The vegetation is composed of open to continuous extreme high elevations. Aymaran highlanders, who live in
bunch grasses and shrubs on upland slopes with wetlands in the puna of Bolivia, are often cited as examples of human
the valley and basin bottoms. Stands of Polylepis spp. (Ro- adaptation to extreme high elevations, with blood hemoglo-
saceae) forest can occur to over 4800 m elevation on west- bin oxygen levels, and lung capacity, higher than any other
ern slopes of the Andes forming one of the world’s highest people. Bofedales are under intensive pressure due to direct
FIGURE 3. Intact peatland during the dry season with water table near the FIGURE 4. Heavy stocking rates of alpaca in bofedale in southern Peru,
surface and cushions and pools separated in most areas; Apolobamba, Vilcanota Mountains (4570 m elevation). (Photo by D.J. Cooper.)
Bolivia, 4350 m elevation. (Photo by D.J. Cooper.)

uses including grazing by the domesticated Andean live- dewater bofedales. Cushion plants are sensitive to grazing.
stock llama and alpaca, and also by European domesticated Llama and alpaca use their chisel like teeth to scrape the
livestock particularly sheep and cattle. Many bofedales hard Distichia shoots from the cushions, and with heavy
have been ditched and drained or irrigated to “improve” the use the cushions are killed and replaced by short-lived and
pastures. shallow-rooted plants (Figure 5).
Other factors also stress these wetlands. Some indus-
PEATLAND TYPES AND CHARACTERISTICS tries, such as metal mining, can have significant environ-
‘Bofedales’ and ‘vegas’ are terms used to denotate many mental effects due to ground water extraction and pollution.
types of Andean wetlands, including fens, wet meadows The climate is also changing, as is well known from the
with mineral soils, and stream floodplains (Chimner et al. rapid loss of glaciers throughout the tropical Andes (Ra-
2019). However, fens are the most conspicuous and char- batel et al. 2013). Some bofedales in small watersheds have
acteristic wetland in the puna. From central Peru to north- completely desiccated for unknown reasons (Figure 6),
ern Chile fens are dominated by cushion-plants, particularly while others persist but have dry season water level de-
Distichia muscoides and Oxychloe andina in the family clines of nearly one meter (Figure 7).
Juncaceae. In central Chile and northern Argentina fens The hydrological functioning of Andean peatlands has
are dominated by cushion plants with a higher presence of been discussed and many conflicting viewpoints provided.
bunch grasses, such as Deyeuxia spp. In southern Bolivia Some have proposed their importance as reservoirs and
and northern Chile and parts of Peru highly saline closed water providers to downstream communities. Others have
basin lakes and flats are common, including the famous suggested they are dependent on glacier meltwater. How-
Salar de Uyuni in Bolivia the world’s largest salt flat and ever, glaciers cover < 5% of the Peruvian highlands and
important habitat for several species of flamingos that also most fens in the puna occur in regions lacking glaciers. In
inhabit many bofedales.
Natural bofedales have a distinctive cushion and pool
FIGURE 6. Dessicated bofedale with bare exposed peat near Huaytire, Peru
structure (Figure 3). The cushion plant shoots grow so (4570 m elevation). (Photo by D.J. Cooper.)
densely packed that they form an almost waterproof dam
that limits lateral water movement, and when the cushions
touch they isolate pools which prevents rapid drainage.
Overgrazing is common as the uplands provide little
forage. In addition, because there is no long lasting winter
snow grazing occurs every day of the year. Stocking rates
can exceed 500 animals per pasture in some districts (Fig-
ure 4). Heavy grazing can break the cushion margins al-
lowing the pools to connect creating drainage paths that can
FIGURE 5. Highly degraded cushion plant peatland with most cushions

destroyed and bare ground exposed. (Photo by D.J. Cooper.)

the puna, as elsewhere in the world, fens are groundwater discharge, the only water resources available during long
dependent ecosystems, with discharge from hillslope and dry periods (Falvey and Garreaud 2005)precipitation over
other aquifers and their evapotranspiration consumes water the high-altitude plateau of the South American Altiplano
resources. The remarkable aridity of the puna ecoregion, exhibits a marked intraseasonal variability which has been
with only 200 - 500 mm of total annual precipitation and associated with alternating moist and dry conditions ob-
strong seasonality with a 7 to 9 month- long dry season, served at surface stations near the Altiplano western cordil-
reinforces bofedale dependence on perennial groundwater lera. In this study the characteristics of humid (wet.
The stability of water provision is a key element for
FIGURE 7. Dry season water table decline in peatlands dominated by Dis- the ecological functionality of these ecosystems. The
tichia muscoides cushions, Nor Yauyos – Cochas, Peru (4630 m elevation).
hydrological stability combined with the relatively con-
Surrounding peaks are up to 5750 m in elevation. (Photo by D.J. Cooper.)
stant year-around thermal regime and daylight intensity
due to proximity to the equator, allows peat to accumulate
(Cooper et al. 2014; Hribljan et al. 2015). These peatlands
form potentially important but usually neglected natural
carbon storage systems in tropical and sub-tropical regions
where extensive lowland peatlands such as the Amazonian
palm swamps in the Pastaza Marañon, the Congo Basin, or
Indonesian swamp forests were considered more important.
The importance of the alpine peatlands is accentuated when
considering the modelled thermal future under climate
change because carbon emissions for the tropical lowland
areas are predicted to rise due to higher water demands
while less significant changes are predicted for the high-
lands (Gallego-Sala et al. 2018).
PEATLAND USES AND CONSERVATION

Andean peatlands are intensively used by pastoralist com-
FIGURE 8. Interpolated groundwater isolines and dominant flow patterns munities as grazing fields for native and introduced live-
in Paucarani an Oxychloe andina dominated peatland. Vilacota – Maure stock and these wetlands are the only ever-green fields.
Regional Conservation Area, Tacna Region, southern Peru. Ground water That capacity for permanent use combined with the low-in-
is flowing (white arrows) from ancient glacial moraines that function as
come of local communities mean that they must constantly
aquifer storage.
increase their flock size and expose the wetlands to over
use, affecting the native vegetation cover and its ecologi-
cal functioning. In some native communities in central and
southern Peru, Bolivia and northern Chile, which preserve
traditional knowledge practices, the wetlands are managed
by artificially spreading water through irrigation with the
aim of increasing the grazing areas. But in the northern
range of the central Puna as in Cordillera Blanca, or in
the more humid eastern slopes, peatland drainage is more
common than irrigation, especially in communities where
traditional camelid-based livestock are replaced by intro-
duced sheep or cattle.
The peatlands are key nesting, forage and water provi-
sion habitat for native wildlife including vicuna (Vicugna
vicugna), small mammals, amphibians and numerous birds
including the South American species of Rhea (R. pen-
nata and R. americana). In central Peru, cushion-plant
peatlands dominated by Distichia muscoides are the only
known feeding habitats for Cinclodes palliatus, a critically
endangered bird species whose total population has not

surpassed 250 adult individuals in recent decades. The Pe- Cordillera, one of the highest mountain ranges in Peru with
ruvian government is the only central Andean country that tropical glaciers, a relatively humid climate with rain that
requires priority protection for wetland ecosystems (article varies between 400 to 700 mm/year and a seven-month dry
99 of the General Environmental Law - Law No 28611) period from May to November. Vilacota – Maure located
and considers bofedales to be fragile ecosystems needing in Tacna province, the most southern administrative region
special protection and compensation measures if disturbed. in Peru including the Atacama Desert and the dry puna
But the threats to those peatlands are varied and continue: includes the completely deglaciated Cordillera del Barroso
overgrazing, illegal peat extraction, drainage, and mining as the main mountain range. The climate in Vilacota Maure
and gas activities. is semi-arid, with precipitation varying from 200 to 400
Current and future conditions of changing climate also mm/year and a long dry season from April to December.
can affect the functionality of these peatlands changing In both areas the vast majority of bofedales are cushion
the amount and timing of rainfall events, modifying the plant-dominated peatlands with groundwater flowing from
infiltration-runoff patterns, and altering the evapotranspira- hillslopes to bottom valleys (Figure 8) where peat thickness
tion loses. can be more than 10 meters. In Nor Yauyos – Cochas the
dominant species is Distichia muscoides while in Vilacota
OUR RESEARCH PROGRAM Maure D. muscoides is replaced by Oxychloe andina that
Our research has focused on understanding the hydrologi- forms less-compact cushions and spreads more extensively
cal and ecological processes that support wetlands in the reducing the presence of pools.
Andes. We use ground water monitoring wells, piezom- Both in Nor Yauyos – Cochas and Vilacota – Maure the
eters, and geochemical tracers to understand the hydrologic best-preserved sites with the thickest accumulated peat layers
regimes that support different wetland types and the water and highest organic matter content are associated with intact
sources that support them (Cooper et al. 2019). The north- cushion plant communities (Figure 9). Higher diversity of
ern Andes from Colombia to northern Peru are generally plant species occurs usually in places influenced by high
much lower elevation and wetter alpine regions (termed grazing pressure, reduced water provision, a combination
“paramos”) where a higher diversity of peat accumulating of these two pressures, or in artificially created wetlands.
wetland communities exist. For example, in northern Peru Usually, areas with higher plant diversity have lower organic
we identified 20 peat accumulating communities (Cooper et matter content and lower values of hydraulic conductivity
al. 2010). However farther south the elevation of the puna in upper peat layers. Meanwhile, at deeper and permanently
rises to much higher levels, and typically only a couple saturated levels, the peat characteristics as OM content and
of communities occur, all dominated by cushion plants or hydraulic conductivity as well as porosity and structure, re-
bunch grasses. The same is true for the Andes in Colombia, mains relatively homogeneous, indicating possible common
where lower elevation peatlands have higher species and vegetation in the past, that has changed in modern times due
community richness and the highest elevations are occupied to climate changes or human interventions.
primarily by Distichia muscoides dominated cushion plant
communities (Benavides and Vitt 2014). FIGURE 9. Correlation between % cover of Juncaceae (Distichia muscoi-
We have also focused on the processes of peat accumu- des and Oxychloe andina) vs. % organic matter in peatlands in the Vilacota
lation to determine whether in today’s climate and under – Maure Regional Conservation Area.
current land uses the fens are accumulating carbon. We
have used the “moss wire” method (Cooper et al. 2015),
cores with 14C aging (Hribljan et al. 2015), and the use of C
gas fluxes measures to demonstrate that intact cushion com-
munities are peat forming. We are expanding into analyses
of ditched, drained, and heavily grazed peatlands, as well as
restored peatlands, as has been done by Rod Chimner and
his students (Planas-Clarke et al. 2020).
Currently, we are developing a series of research proj-
ects to understand the hydrological behavior and carbon
dynamics of peatlands in Nor Yauyos – Cochas Landscape
Reserve and Vilacota – Maure Regional Conservation Area,
two remarkably different areas of the Peruvian puna (Figure
1, A and B). Nor Yauyos – Cochas is located in the Central

A key research goal for the future is to quantify the Falvey, M. and R.D. Garreaud. 2005. Moisture variability over the South
American Altiplano during the South American low level jet experi-
condition of peatlands, their carbon stock, and the pro- ment (SALLJEX) observing season. Journal of Geophysical Research
cesses that sustain and degrade them. A large proportion Atmospheres 110: 1–12.
of bofedales are highly degraded by land uses, particularly Gallego-Sala, A.V., D.J. Charman, S. Brewer, S.E. Page, I.C. Prentice, P.
heavy and continuous grazing, and their cushion plants Friedlingstein, … and Y. Zhao. 2018. Latitudinal limits to the predicted
have been partly or completely killed. We need to under- increase of the peatland carbon sink with warming. Nature Climate
Change 8: 907–913.
stand the level of grazing that is sustainable for the per-
Hribljan, J.A., D.J. Cooper, J. Sueltenfuss, E.C. Wolf, K.A. Heckman,
sistence of ecosystem services provided by the peatlands E.A. Lilleskov, and R. Chimner. 2015. Carbon storage and long-term rate
and for the persistence of indigenous communities that of accumulation in high-altitude Andean peatlands of Bolivia. Mires and
rely on grazing for their livelihood. For ecosystems that Peat 15: 1–14.
are already highly degraded we need to understand how Olson, D.M., E. Dinerstein, E. Wikramanayake, N.D. Burgess, G.V.N.
to restore their hydrologic regimes, successfully introduce Powell, E.C. Underwood, … and K.R. Kassem. 2001. Terrestrial Ecore-
gions of the World: A new map of life on Earth. BioScience 51: 933–938.
cushion plants, and rebuild plant production and carbon
Planas-Clarke, A., R. Chimner, J. Hribljan, E. Lilleskov, and B. Fuent-
storage. There are many challenges ahead as the people ealba. 2020. The effect of water table levels and short term ditch restora-
of the Andes depend on these bofedales almost completely tion on mountain peatland carbon cycling in the Cordillera Blanca, Peru.
for their livelihoods, but have used them intensively, to the Wetlands Ecology and Management 28: 51-69.
point of severe degradation and collapse in some areas. Rabatel, A., B. Francou, A. Soruco, J. Gomez, B. Cáceres, J.L. Ceballos,
The future will require intensive management and restora- and P. Wagnon. 2013. Current state of glaciers in the tropical Andes: a
multi-century perspective on glacier evolution and climate change. The
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WETLAND RESEARCH
What is the Flora of the Pantanal Wetland?

Arnildo Pott1 and Vali Joana Pott2
ABSTRACT region is not a swamp, but seasonally flooded land that also
he Brazilian Pantanal is a vast Neotropical wetland, in includes slightly higher flood-free ground.
T the upper Paraguay River basin. Rainfall and flooding
are seasonal. The landscape is heterogeneous showing a
The climate is typical of savanna, type Aw (tropical
wet-and-dry), with annual rainfall around 800-1400 mm,
mosaic of vegetation types. Overall, it is predominantly a most of which comes from variable summer rains (Arruda et
savanna, with aquatic plants, riparian and dry forests, for- al. 2016; Figure 2) creating an oscillating pluriannual flood
est islets, woodlands, grasslands and many monodominant cycle (Pott and Pott 2004), as shown in Figure 3. The irregu-
formations. The flora is composed of over 2,200 species of lar flood regime has been associated with oscillations of sea
Angiosperms, and the species-richest families are Fabaceae surface temperature such as ENSO (El Niño South Pacific
(Pea Family) and Poaceae (Grass Family), each with over Oscillation) (Thielen et al. 2020). The general flood pulse is
300 species, followed by Asteraceae (Daisy Family) and monomodal (Junk et al. 2011), although the Paraguay River
Cyperaceae (Sedge Family), both with more than 100 spe- tributaries such as Aquidauana can show more than one
cies each. The species richest genera are Paspalum, Ipo- yearly flood peak. Rainfall is higher in the upper watershed,
moea (morning glories), Mimosa (sensitive plants), Croton, wherefrom whatever surplus runs off to the plain. The asyn-
Eugenia, Ludwigia (primroses) and Arachis (wild peanuts). chrony between local rainfall and delayed river flood creates
Very few are endemic species, as the region is geologi- a prolonged wet period for the riverine vegetation, allowing
cally recent. The flora comes from surrounding domains, some Amazon rain forest species to grow there.
such as Cerrado, Chaco, Amazon, and Atlantic Forest,
FIGURE 1. Map of the Brazilian Pantanal wetland (grey) and the upper
although most species have broad distribution. However, watershed (white) with the tributaries of the Paraguay River. (Source: Arruda
their arrangement and dynamics are unique in the Pantanal. et al. 2016.)
Human population is quite low, except on the edges. Cattle
ranching is the main economy for over 200 years. The
conservation status of the Pantanal is still considered rather
natural and pristine. Tourism is increasing, mainly for the
abundant wildlife.
INTRODUCTION
The Pantanal is the largest continuous freshwater wetland
in South America, located in the middle of the continent,
with 140,000 km2 in Brazil (Figure 1), and additional
15,000 km2 in near Bolivia and 5,000 km2 in Paraguay
(Junk et al. 2011). It is situated in a vast intracontinental
flat lowland, filled with Pliocene-Pleistocene sediments,
forming a somewhat inland delta in the upper Paraguay
River basin (Junk et al. 2011). It is a heterogeneous wetland
for its complex hydrology (Junk et al. 2011), sediments
and vegetation types, and for that reason it has been subdi-
vided into 11 subregions. The term Pantanal, derived from
pântano or swamp, can be misleading since most of the
1 Visiting Professor, INBIO-Instituto de Biociências, UFMS-Universidade Fed-

eral de Mato Grosso do Sul, Campo Grande, MS, Brazil. Corresponding author
contact: arnildo.pott@gmail.com
2 Botanist, Herbarium CGMS, INBIO, UFMS, Campo Grande, MS, Brazil.

FIGURE 2. Mean monthly rainfall and Paraguay River level (Brazilian Navy fluviomet- ceae, with the richest genera being Bacopa (water-
ric gauge at Ladário), showing the asynchrony of rains and flooding. (Source: Arruda hyssops), Cyperus (umbrella sedges), Eleocharis
et al. 2016.) (spike rushes), Ludwigia (water primroses), Rhyn-
chospora (beak rushes), and Utricularia (bladder-
worts). The Pantanal has a range of aquatic habitats
and seasonally wet zones that favor a diversity of
species. Most aquatic species are common to other
vast South American wetlands, such as the Paraná
River floodplain (extending from Brazil to the Es-
teros de Iberá in Argentina; Neiff et al. 2011), Ama-
zonia (including Araguaia, Guaporé and Amapá in
Brazil, and the Beni region in Bolivia; Haase and
Beck 1989), and the Llanos and Orinoco basin in
Colombia and Venezuela (Rial 2009).
Wherefrom does the flora come? The plants
of the Pantanal wetlands are composed of species
from the surrounding phytogeographical provinces:
Cerrado, Chaco, Amazonia and Atlantic Forest, in
addition to a major contingent considered of wide
distribution (Pott and Pott 2004). Cerrado, the Bra-
zilian savanna, nearly surrounds the entire Pantanal. Chaco
Research on the Pantanal in the last 30-40 years has
is a unique vegetation type in Brazil, located the southern-
been made mainly by local institutions such as Embrapa
most subregion of the Pantanal - in Porto Murtinho, on the
Pantanal, IPP (Research Institute for the Pantanal) and sev-
Paraguayan border, belonging to the Humid Chaco (Prado
eral universities (i.e., Federal University of Mato Grosso do
et al. 1992). Amazonian species reach the riparian forests
Sul - UFMS, Federal University of Mato Grosso - UFMT,
because the overflow of the Paraguay River has a three-
Catholic University Don Bosco - UCDB, University of
month delay from the rainy season (i.e., it overflows three
the State of Mato Grosso – Unemat, and University for the
months after the local and upper watershed rainy season;
Development of the State and the Pantanal Region – Uni-
Figure 2). That adds meteorological and edaphic wet
derp), sometimes with collaboration of other Brazilian and
periods with a similar effect of an Amazonian climate. The
international scientists.
Atlantic Forest species include elements from the Season-
PLANT LIFE IN THE PANTANAL ally Dry Forest. The wide-ranging group represents half
The flora of the Pantanal wetlands is represented by over of the species of the Pantanal plain (Pott and Ratter 2011;
2,200 native species of Angiosperms, according to the Pott and Silva 2015) and includes all plant habits, most
updated checklist we compiled from reliably identified of them grassland herbs. However, the plant diversity of
vouchers in herbaria, strictly considering the sedimentary the Pantanal is unique, despite the flora being made up of
floodplain. Nearly 800 species are illustrated in our two associations of species from various phytogeographical
identification guides (Pott and Pott 1994, 2000). The most origins (Pott and Ratter 2011), tolerant of a harsh wet-and-
numerous families are Fabaceae (Pea Family) and Poaceae dry environment.
(Grass Family), each with over 300 species, followed by The number of naturalized species is relatively high
Asteraceae (Daisy Family) and Cyperaceae (Sedge Fam- but mostly concentrated on less flooded ground. We can
ily), both with more than 100 species. Genera exceeding 20 deduce that in a future scenario of a drier Pantanal, they
species are Paspalum, Ipomoea, Mimosa, Croton, Eugenia, will expand their distribution. Indeed, Leucaena leuco-
Ludwigia and Arachis. That follows the pattern of species cephala (leucaena or white lead tree, a fast-growing tree
of these genera being very abundant in Neotropical open from Central America and Mexico) is already spreading
vegetation types. Due to the recent geological age of the on road embankments. So far, there are no massive exotic
floodplain, very few species are endemic to the Pantanal, woody invaders that are common in other tropical wetlands,
and five of them remarkably belong to the genus Arachis although some native shrubs (Mimosa spp. and Vernonan-
(wild peanuts). thura brasiliana) and trees (e.g., Curatella americana) in-
The number of aquatic macrophytes is over 300 spe- crease in dry years. There are many naturalized herbaceous
cies. The species-richest families are Cyperaceae and Poa- plants, mainly on less floodable grasslands. The few exotic

weeds to invade permanently wet habitats are Panicum FIGURE 3. Zonation of monodominant stands: anchored floating mats
repens (torpedo grass) and the worst invader Urochloa ar- of water hyacinths (Eichhornia azurea + E. crassipes) and Pontederia
recta (tanner grass) (Pott and Pott 2004). rotundifolia, on the left, and emergent Oryza latifolia (wild rice), on the
Overall, most of the vegetation of the Pantanal is right, western border of the Pantanal wetland, Brazil. (Photo by A. Pott, June
predominantly a seasonally flooded savanna (Pott and Pott 2, 2009.)
2004), with mosaics of aquatic plants (including floating
mats, floating meadows and swamps) (Figure 3), riparian
forest, dry forest, forest islets, woodlands, grasslands (Fig-
ure 4), and many monodominant formations. Plant species
are arranged over a flooding gradient varying from lakes
and permanent swamps to flood-free ancient levees, and
floodable grassland in between (Figure 5). Soils vary in tex-
ture and fertility from 97% sandy to heavy clays, according
to the sediment type. The largest alluvial fan in the Pantanal
is the Taquari River, encompassing 50,000 km2 of sand.
A striking feature of the Pantanal is the monodominant
vegetation types, mainly floodable savannas or woodlands,
with a single dominant tree, such as Attalea phalerata (uru-
FIGURE 4. Floodable grassland, at drawdown, Pantanal wetland, Brazil.
curi palm or acuri), A. speciosa (babassu palm), Byrsonima (Photo by A. Pott, June 12, 2007.)
cydoniifolia (canjiqueira), Copernicia alba (caranday palm
or carandá) (Figure 6), Erythrina fusca (purple coral tree
or abobreira), Handroanthus heptaphyllus (pink trumpet
tree or piúva) (Figure 7) and its relative Tabebuia aurea
(Caribbean trumpet tree or paratudo) (Figure. 8). Some of
the monodominant species cause encroachment on natural
grasslands, e.g., the trees Curatella americana (sandpaper
tree or lixeira) and Vochysia divergens (cambará) (Figure
9), and shrubs as Combretum spp. (pombeiro). Some her-
baceous species are also monodominant such as Cyperus
giganteus (giant sedge or piri, similar to the Nile papyrus),
Elyonurus muticus (carona), Oryza latifolia (wild rice),
Thalia geniculata (fireflag or caeté) (Figure 10) and Typha
domingensis (cattail or taboa).
A very interesting vegetation type is the floating
meadow (Figure 11). It is a floating islet that starts from a FIGURE 5. Typical landscape of the Pantanal wetland, with pond, floodable
floating mat of free-floating plants such as Salvinia au- grassland and seasonal forest or woodland, over the topographic flooding
gradient, at the end of flood in a wet year, Brazil. (Photo by A. Pott, May 9,
riculata (water fern). On its top the sedge Cyperus blepha-
2009.)
roleptos (formerly Oxycaryum cubense) germinates as an
epiphyte and grows, forming a dense rhizomatous entan-
gled meadow where debris and decaying plants accumulate.
Within a few years a layer of floating organic soil (histosol)
builds up. Caymans and birds nest on it. When that floating
soil reaches 1 m deep, it is possible to walk on, and even
shrubs and a few treelets grow on it. That sudd can clog
old river beds and canals, or drift by winds or float down-
stream. When the lake dries out, the plants die, the organic
soil decomposes or can be consumed by a wildfire, and the
process restarts after the water returns.
Interestingly, alkaline ponds (pH 8-10) called “sali-
nas” also occur; they are surrounded by a flood-free ridge
covered with dry forest and so isolated from other waters.

FIGURE 6. Monodominant stand of Copernicia alba (caranday palm or FIGURE 7. Monodominant floodable savanna of Handroanthus heptaphyllus
carandá), Paraguay River, Pantanal wetland, Brazil. (Photo by A. Pott, July (pink trumpet tree or piúva, or ipê-rosa), in the dry season, Miranda River
9, 2013.) floodplain, Pantanal wetland, Brazil. (Photo by A. Pott, September 2000.)
FIGURE 8. Monodominant floodable savanna of Tabebuia aurea (Carib- FIGURE 9. Monodominant floodable forest of Vochysia divergens
bean trumpet tree or paratudo), at flood, Pantanal wetland, Brazil. (Photo (cambará), at flower, along seasonal streams, in flooded grassland, at
by P.R. Souza, May 22, 2007.) drawdown, Pantanal wetland, Brazil. (Photo by Fábio Edir Costa, July 29,
2008.)
FIGURE 10. Monodominant stand of Thalia geniculata (fireflag or caeté), FIGURE 11. Floating meadow with 1m deep histosol (8 persons walking
Paraguay River floodplain, Pantanal wetland, Brazil. (Photo by A. Pott, on including the photographer): Polygonum acuminatum in the fore-
June 3, 2009.) ground, the red plant is Rhynchanthera novemnervia, and riparian forest
in the back, in the lake Baía Vermelha, near the Paraguay River, Pantanal
wetland, Brazil. (Photo by A. Pott, June 2, 2009.)

Scarce angiosperms grow inside, e.g., Ceratophyllum spp. dichotomus), plus the tapir (Tapirus terrestris). This surplus
(hornworts), besides Characeae (large algae); the water is of forage is quite different from the situation in African
a soup of algae, resulting in ponds of various colors. While savannas that are under high pressure from herds of herbi-
there are no fish, amphibians, myriads of small crustaceans vores. The worst impact in the Pantanal has been from tree
and water insects attract young caymans and migratory clearing of ridges for cultivated pastures. However, lately,
birds such as sandpipers. deforestation has lost financing and lessened this activity in
the region. Instead, many areas of coarse grasses are being
FACTORS AFFECTING VEGETATION replaced by Urochloa humidicola (koronivia grass – an
We have analyzed soil seed banks from seasonal ponds, African tropical forage species, widely cultivated in South
floodable grasslands, Tabebuia aurea (Caribbean trum- America) around the woody patches on hummocks, thereby
pet tree) monodominant savanna, and riparian forest, and maintaining the general landscape pattern and even reduc-
found that annual species predominate even under forest. ing fire incidence. Besides, capybara and deer relish this
The discrepant composition of the standing vegetation and evergreen introduced grass.
the seed bank is a general finding. Under simulated flood- Buffaloes have a heavy impact, mainly in ponds, as we
ing, aquatic plant seedlings emerged from the soil samples, observed, reducing the species richness, in favor of a few
while switching to terrestrial species after alternation to benefiting from their dung, e.g., Pistia stratiotes (water
drained conditions. We called this a “flexible seed bank” lettuce). They are no longer raised in the Pantanal because
– one that is ready to occupy gaps in either the dry or wet they escape and become feral.
season quickly. Furthermore, the aquatic plants can hold the
exotic grasses back (Bao et al. 2020). HAVENS FOR WILDLIFE
As a seasonal savanna, the Pantanal undergoes wild- Like other sedimentary floodplains and alluvial fans, the
fires, often lit by lightning mainly on the surrounding hills, Pantanal contains various types of water bodies, such as
then fed by dry biomass of grasslands and accumulated his- rivers, river branches, old river beds, oxbow lakes, lakes,
tosol. The vegetation evolved under fire. Interestingly, even permanent and temporary ponds, seasonal streams and
riparian forests regenerate after fires and exhibit a striking flooded grasslands. All these water bodies are habitat for
favorable interacting response to fire and flood (Arruda et the abundant wildlife: fish, anaconda, bulldog bat, giant
al. 2016). The most fire-prone habitats are ungrazed deep- otter; birds such as anhinga, cormorant, ducks, herons,
flooded grasslands along the Paraguay and Miranda Riv- ibis, jabiru stork, kingfishers, limpkin, screamer, skimmer,
ers, compared with less flooded savannas where cattle stay spoonbill, terns, wattled jacana, wood stork, and many
year-round. When cattle are excluded, the grasslands either more. However, for the most part, there is no permanent
in swamps or dryland change to tall tussock grasses that are water, only seasonal water bodies including excavated wa-
fuel for inevitable wildfires. In 1988, we excluded the cows ter holes or where underground water is pumped for cattle,
from a 680 ha preserve to see what would happen: within and it is, of course, also utilized by the wild fauna. The
one year the short creeping grasses were replaced by tall harmonic coexistence of domestic and wild animals begins
bunch grasses, and wildfire has swept through many times, by sharing that provided water, as well as the salt.
despite the presence of a 10 m wide firebreak. The same Despite being floodable, the Pantanal is also rich in ter-
has occurred in other areas, e.g., the 108,000 ha private restrial fauna, such as agouti (rodents), armadillos, coati-
preserve SESC Pantanal, created in 1996, frustrating the mundi, crab-eating foxes, giant anteaters, howler monkeys,
expectation that the grasslands would turn into a forest after and peccaries. The big cats – jaguar and puma - are also
cattle exclosure. Prescribed burning can be a management frequent. Many species of birds, especially three species of
tool for conservation. macaws including the hyacinth macaw (elsewhere endan-
Despite over 200 years of cattle ranching on native gered), as well as parrots, parakeets, cardinals, the noisy
grasslands, the impact seems to be low because there is a speckled chachalaca, burrowing owl, flycatchers, hum-
surplus of available forage grass, amounts far above what mingbirds, lapwing, red-legged roadrunner, rhea, toucans
can be consumed by the few species of native herbivo- and woodpeckers.
rous mammals (Pott and Pott 2004). The most numerous
of these herbivores are capybaras (Hydrochoerus hydro- HUMAN USE AND CONSERVATION
chaeris, the world’s largest rodent). Others are naturally The Pantanal people do not hunt for meat, except the non-
more scattered, sometimes with vigorous populations de- native feral pig, as they prefer beef. For over 200 years,
spite being threatened elsewhere; they include the pampas people have raised cattle in the region. The human popu-
deer (Ozotocerus bezoartius), a few related deer species lation is quite low on the plain, except in a few towns on
( i.e., brockets, Mazama spp.and marsh deer, Blastocerus the edges. Crop agriculture is not allowed on the plain,

however it is naturally limited by unfavorable soils. A sub- Junk, W.J., C. Nunes da Cunha, C.J. da Silva, and K.M. Wantzen. 2011.
The Pantanal: a large South American wetland and its position in limno-
sistence shift cultivation used to be practiced by riverside logical theory. In: W.J. Jung, C.J. Silva, C.N. Cunha, and K.M. Wantzen
inhabitants and is now legally restricted under protective (eds.). The Pantanal: Ecology, Biodiversity and Sustainable Manage-
environmental laws. Nonetheless, land mismanagement in ment of a Large Neotropical Seasonal Wetland. Pensoft Publishers,
the 1980s (e.g., land clearing Cerrado woodland of sandy Sofia, Bulgaria. pp. 23-44.
slopes for cattle pastures, creating deep gully erosion and Neiff, J.J., S.L. Casco, A. Cózar, A.S.G. Poi de Neiff, and B. Ubeda.
2011. Vegetation diversity in a large Neotropical wetland during two
damaging small headwater wetlands called veredas) in the different climate scenarios. Biological Conservation 20 (9): 2005-2020.
upper watershed is interfering with water and soil conserva- DOI:10.1007/s10531-011-0071-7
tion, causing significant river silting downstream, mainly Pott, V.J. 1999. Riqueza verde em meio azul. In: E. Scremin-Dias, V.J.
in the Taquari River that has lost its bed and gallery forest Pott, R. Hora, and P.R. Souza (eds.). Nos jardins submersos da Bodo-
and became a swamp. Silt and overflow is spreading over quena: guia para identificação das plantas aquáticas de Bonito e região
(In the underwater gardens of Bodoquena: identification guide for the
a nearly permanently flooded delta. Compared with the aquatic plants of Bonito and region). UFMS, Campo Grande. pp. 59-93.
Pantanal plain, the upper watershed has a higher richness https://ecoa.org.org/wp - content/uploads/2016/12/Nos - Jardins - Sub-
of aquatic plants, as well as of endemic species, mainly in mersos – da – Bodoquena.pdf
vereda wetlands in the Cerrado savanna. These wetlands Pott, V.J., A.C. Cervi, N.C. Bueno, and A. Pott.) 1999. Dinâmica da vegetação
aquática de uma lagoa permanente da Fazenda Nhumirim, Pantanal da Nhe-
are wet year-round, functioning as water storage for the colândia, MS. In: Simpósio sobre Recursos Naturais e Socio-Econômicos do
creeks that feed the rivers. There are also springs on karstic Pantanal; Manejo e Conservação. Corumbá, 1996. Anais. Corumbá, Embrapa.
limestone of the Serra da Bodoquena plateau that support a pp. 227-235.
different aquatic flora (Pott 1999). Pott, A. and V.J. Pott. 1994. Plants of Pantanal. Corumbá, Embrapa.
The official conservation areas are the National Park of Pott, A. and V.J. Pott. 2004. Features and conservation of the Brazilian
Pantanal, the State Park Rio Negro, the State Park Encontro Pantanal wetland. Wetland Ecology and Management 12: 547-522.
das Águas, and the Taiaman Biological Station (Ramsar Pott, V.J. and A. Pott. 2000. Plantas Aquáticas do Pantanal. Brasília,
Embrapa.
site). Fishing is controlled, forbidden during the spawning
period when river waters rise in the rainy season. Ecotour- Pott, A. and J.A. Ratter. 2011. Species diversity of terrestrial plants and
human impact on the vegetation of the Pantanal. In: W.J. Jung, C.J.
ism is a growing economic activity in the region, for its Silva, C.N. Cunha, and K.M. Wantzen (eds.). The Pantanal: Ecology,
abundance of wildlife and scenery, e.g. bird watching. Biodiversity and Sustainable Management of a Large Neotropical Sea-
There is also a potential for scientific tourism. In conclu- sonal Wetland. Pensoft Publishers, Sofia, Bulgaria. pp. 281-300.
sion, as already stated (Heckman 1998; Junk et al. 2011), Pott, A. and J.S.V. Silva. 2015. Terrestrial and aquatic vegetation
diversity of the Pantanal wetland. In: I. Bergier and M.L. Assine (eds.).
the overall conservation status of the Pantanal is still high, Dynamics of the Pantanal Wetland in South America. The Hand-
and it is considered a pristine, natural wetland. n book of Environmental Chemistry 37. Switzerland, Springer. DOI
10.1007/609_2015_352.
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Rojas, J.I. Quintero, and M.I. Marques. 2020. Quo vadis Pantanal?
Haase. R. and S. Beck. 1989. Structure and composition of savanna veg- Expected precipitation extremes and drought dynamics from chang-
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Heckman, C.W. 1998. The Pantanal of Poconé: Biota and Ecology in org/10.1371/journal.pone.0227437
the Northern Section of the World’s Largest Wetland. Kluwer Academic
Publishers, Dordrecht.

WETLAND RESEARCH
Connectivity of River Floodplains - The Case of Ibera Wetlands after 10,000 Years of
Isolation from Parana River
Juan J. Neiff1, Sylvina L. Casco1,2, Andrés Cozar Cabañas3, Alicia S.G. Poi1, Bárbara Úbeda3, Luisa F. Ricaurte4, and
Eduardo M. Mendiondo5
ABSTRACT connectivity may be uneven for populations and commu-

“Esteros del Ibera” is one of the most outstanding wetlands nities (Neiff et al. 2009). Tockner et al. (1998) found that
in South America, by its size (12,300 km2) and biodiver- the connectivity of floodplain habitats to the river course
sity, the largest recorded at this latitude. Owing to this, it showed diverging values for different organisms. While
is recognized as a Ramsar Wetland of International Im- fish diversity was higher in the active Danube floodplain,
portance and as a National Park in Argentina. Esteros del amphibians showed greater diversity in floodplain habitats
Ibera is an “open-air laboratory” since its landscape was isolated from the river. Wiens (1989) pointed out that flood-
created by the lateral migration of the Parana River, leav- plain landscape should be observed from the perspective of
ing a vast paleo-alluvial fan from Argentina to Paraguay. organisms instead of from an anthropocentric viewpoint.
Here, we compare the species richness in the landscapes of Episodes of flooding change the proportion of aquatic and
Ibera with that in the equivalent landscapes of the active terrestrial landscapes, altering physico-chemical properties
course of Parana River in order to understand the causes of and biotic exchanges between water and land (Wiens 2002;
change in diversity patterns over the past 10,000 years. We McClain and Naiman 2008; Almeida and Melo 2009).
found that the loss of connectivity with the pulse regime of Current concepts apply according to the particular
Parana River led to an increase in specific complexity of definition of connectivity adopted. According to the River
Ibera biota. This likely resulted from the combination of a Continuum Concept (Vannote et al. 1980), the idea of
limited change in water quality, the belonging to the vast longitudinal connectivity prevails. The Serial Discontinuity
Amazon biogeographical domain, the natural niche ampli- Concept (Ward and Stanford 1995), used for single-channel
tude of wetland species, and the self-designing capacity of rivers, states that the stability is varyingly influenced by ter-
the Ibera ecosystem. restrial ecosystems. Wiens (2002) argued that connectivity
has three dimensions, namely longitudinal, lateral and verti-
INTRODUCTION cal. The vertical dimension of connectivity seems obvious,
Understanding the spatial and temporal variability of due to the turbulence of the flow. These three dimensions of
wetlands at different time scales requires analyzing the connectivity as well as the pulse regime should be evalu-
underlying biogeochemical and ecological processes (Junk ated at different scales depending on the landscape interac-
1997; Tiner 2003; Neiff 2004; Dawidek and Ferencs 2016). tions and the processes analyzed.
There is consensus on the importance of connectivity To the best of our knowledge, no previous study ad-
between the river’s course and the floodplain (Junk et al. dressing the natural loss on connectivity of fluvial wetlands
1989; Neiff 1990; Junk 1997; Melack and Forsberg 2001) on the scale of thousands of years exists in the literature.
as the variation in water level is responsible for a complex We have an example from South America – for the Ibera
dynamic equilibrium in floodplain landscapes (Amoros wetland in northeastern Argentina, a former floodplain of
and Bornette 2002; Thoms 2003; Bunn et al. 2006; Wiens the Parana River.
2009; Dawidek and Ferencs 2016). However, the effects of
• While numerous studies addressed the limnological
features of Parana River and Ibera wetland since the
1 Centro de Ecología Aplicada del Litoral (CONICET-UNNE). Corrientes, Argen-
tina. Correspondence author contact: jj@neiff.com.ar; guadalupepoi@gmail.com 1980s, basic questions remain:
2 Facultad de Ciencias Exactas y Naturales y Agrimensura (FaCENA-UNNE).
Corrientes, Argentina. sylvina.casco@exa.unne.edu.ar
• Was there a substitution of plant and animal species
3 Departamento de Biología, Instituto Universitario de Investigacion Marina in Ibera as result of the isolation from the river’s
(INMAR), Universidad de Cádiz, Puerto Real, 11510, Spain. andres.cozar@uca. pulses?
es; barbara.ubeda@uca.es
4 Alexander von Humboldt Biological Resources Research Institute, Bogota D.C., • Has the complexity of the assemblages been modified
Colombia. ricaurte.luisa@gmail.com
5 Escola de Engenharia de São Carlos Universidade de São Paulo, Brazil. e.mario.
at the level of species richness, or the spectrum of
mendiondo@gmail.com bioforms in the vegetation?

• Has the isolation of the Ibera wetland produced STUDY AREA
changes in resilience? Our study area includes a portion of the Parana River in
• Was the capture and accumulation of carbon in the northeastern Argentina, its active floodpliain and its former
Ibera wetland modified? floodplain - the Ibera wetland (Figure 1). Parana River is
anastomosed in this section and runs with a slope of 0.6-0.8
We believe that studies on a long-term scale (10,000
m/km on a bed of basalts covered by poorly sorted sands and
years) allow a better understanding of the resilience of
clay that form bars and islands originating from the Pleisto-
biota, the functioning and organisation of the natural sys-
cene to the Holocene (Orfeo and Stevaux 2002). The hydro-
tem. Here we provide a first attempt to address these issues
logic regime of Parana River is quite irregular. It includes a
based on available information. This knowledge is expected
period of high waters in summer, with maximum levels in
to prove useful for environmental management, biodiver-
February-March, and a period of low waters, with minimum
sity conservation, and for evaluating carbon sequestration
values between August and the beginning of September.
and current problems of tropical wetlands. We propose that
The mean discharge is around of 18.000 m3/s and peak
the collection of plant and animal species in Ibera, although
flow around 65.000 m3/s. In extraordinary floods, the entire
it evolved in response to the new connectivity conditions
floodplain is covered by a continuous mass of water, expos-
(i.e., isolation from fluvial processes), preserved most of
ing only the treetops of the gallery forest on the floodplain.
the species of the river domain.
Water of Parana River is, generally, neutral (pH
FIGURE 1. Parana River and Esteros del Ibera (adapted from Poi et al. 2017). The blue 6.5-7.3), with low salinity (E.C. 40-90 µS.cm-1),
line delimits the Esteros de Ibera. little calcium, abundant silica, and high turbidity
and color during floods, due to loads of suspended
N
solids reaching 100 mg L-1 (Bonetto 1986a).
PARAGUAY During the Pliocene (5.3 to 2.5 million years
ago), Ibera wetland was part of the extensive
Parana River active floodplain of the Parana paleo-river. River
water in Ibera flowed through braided channels
Corrientes separated by sand bars. Movements within the
Earth’s crust during the Cretaceous Period led
to sweeping changes in the regional topography,
Luna Lake
modifying the slope and direction of surface run-
off. Paleo-river Parana gradually migrated from
Galarza Lake the basin now occupied by Ibera to the current
Itati Lake
channel placed between Paraguay and Argentina
Ibera Lake at the end of Pleistocene roughly 10,000 years
ago (Castellanos 1965; Iriondo, 1991; Popolizio
1977; 2004; Neiff 2004; Orfeo and Neiff 2008;
Corriente River Iwaszkiw et al. 2010). The ancient riverbed of
Parana now known as the Ibera depression is
now a basin of 30,000 km2 fed by rainfall (Herbst
BRAZIL
and Santa Cruz 1999. Currently any excess water
drains into the Parana River through Corriente
River in the south of the Ibera system.
Its biological diversity is similar to that of the
ARGENTINA Parana River, in despite of the absence of a ge-
ographical connection between them. The Iberá
region was a wide fluvial belt from the Pliocene
with low sinuosity braided channels separated by
URUGUAY sandy natural levees. This fluvial system related
with the origin of the Parana River, deposited the
extense Ituzaingó Formation (fine sandy sedi-
ments with iron compounds) with outcrops in the
inner part of the studied area. During the Cre-

taceous Period, the epirogenic movements of the sub-sur- Ibera wetland represents a unique and long-lasting environ-
face basaltic blocks generated local changes of the slope, ment for the collection of Amazon species that settled there
modifying the drainage direction. The Parana paleochannel (Cabrera 1951; 1976; Cabrera and Willink 1973; Carnevali
gradually abandoned the central depression of Corrientes, 2003; Zalocar de Domitrovic 2003).
changing to the present course that shows structural control. It is important to note that the forms of life found in the
The fluvial belt was transformed into a series of intercon- Paleo Iberá date from 3000 to 4500 years BC (Cuadrado
nected waterbodies that receive the influence of local rains. and Neiff, 1994; Morton, 2004; Pacella and Di Pasquo,
The local climate is Humid Subtropical, with rainfall of 2020) and there are no records of plant or animal life
1500 mm/yr, athough rainfall can exceed 2500 mm in years between that date and the time of Iberá’s disconnection
influenced by ENSO events (Poi et al. 2017). During most from the river regime. Paleontologists still do not have an
of the year, temperature typically ranges from 20oC to 30oC, answer to this absence of preserved paleontological ma-
with a maximum absolute temperature of 44oC and mini- terials, although it is known that it was a period of very
mum absolute temperature of 1oC. contrasting climatic changes. There is a significant gap in
Rainfall has driven the geomorphology and hydrology the geological and biological history of Corrientes Prov-
of Ibera for millenia. No differences have been found in the ince from the recent Holocene to the late Pleistocene with
pollen analyses in the whole area of study (Cuadrado and the 40,000-year-old Toropí Formation. Unfortunately, the
Neiff 1993), and 14-Carbon analyses gave an age of 3000 to sequence of the Late Pleistocene and Early Holocene has
3700 years for the current landscape of Ibera and the older not been preserved1.
islands in Parana River. However, geomorphological and
Personal communication from Dr. Alfredo Zurita, Facultad de
sedimentological studies agree that Ibera lost its connec-
1
Ciencias Exactas y Naturales, Universidad Nacional del Nor-
tion with the Parana River 10,000 years ago. Therefore, the deste, Argentina, aezurita74@yahoo.com.ar
TABLE 1. Information published about Ibera and Parana wetlands

Areas Scientific contributions
Geology and Geomorphology Castellanos (1965); Popolizio (1977; 1981); Herbst and Santa Cruz (1999); Iriondo
(2004); Orfeo and Stevaux (2002); Orfeo and Neiff (2008)
Climatic Change effects Neiff et al. (2011); Neiff and Neiff (2013); Úbeda et al. (2013)
Phytoplankton Zalocar de Domitrovic (1990; 1992; 2003); Cózar et al. (2003); Zalocar de Domitro-
vic et al. (2007)
Zooplankton Corrales de Jacobo and Frutos (1982); Frutos (2003; 2008; 2017); Cózar et al.
(2003); Frutos et al. (2009)
Benthic invertebrates Varela and Bechara (1981), Varela et al. (1983); Bechara and Varela (1990)
Invertebrates associated to Poi de Neiff (2003); Poi de Neiff et al. (2006); Poi (2017)
aquatic plants
Ichtyofauna Bonetto et al. (1981); Bonetto (1986a,b); Jacobo (2002); Almirón et al. (2003), Ca-
sciotta et al. (2005); Neiff et al. (2009); Iwaszkiw et al. 2010; Contreras et al. (2017)
Vegetation Cabrera (1976); Neiff (1986; 2003); Arbo and Tressens (2002); Carnevali (2003);
Neiff and Casco (2017)
Wildlife and Biogeography Cabrera and Willink (1973); Alvarez et al. (2003); Giraudo and Arzamendia (2003)
Ecology and Limnology Cuadrado and Neiff (1993); Neiff et al. (1993); Canziani et al. (2003); Gantes and
Torremorel (2005); Poi de Neiff (2003); Neiff (2004); Poi et al. (2017)

FIGURE 2. Satellite image of Parana River paleo-fan indicating the loca- try to show the biotic divergence derived from the isolation
tion of the different wetland groups identified in this study according to of the fluvial dynamics. We think that the indicators used
their connectivity to the Parana River. I: Isolated lakes and marshes; E: here can be easily replicated in other areas as they relate to
Eventually connected wetlands; C: Close connected wetlands. water and nature conservation management projects.
Finally, we will mention what knowledge is needed based
on our analysis and when it is necessary to evaluate environ-
mental impacts of wetland isolation on a long time scale.
WETLAND TYPES AND DISTRIBUTION

For this region, the Parana River ecosystems contains three
types of wetlands based on differences in water supply and
pulse regime (Figures 2-5): isolated, eventually connected,
and close connected.
Isolated wetlands (Group I). They correspond to the Ibera
lakes and are locally called as “esteros”. Ibera lakes are located
in the fluvial paleo flatland very close to the current course of
the river in the Ibera region (27°30–29°00S, 56°25–58°00W).
The lakes are large (8 to 95 km2) and surrounded by extensive
marshes with sudds (floating islands; Figure 3). The water was
relatively transparent, slightly acidic or neutral, the conductiv-
ity ranged between 9 and 52 µS.cm-1 and the dissolved oxygen
concentration varied between 5.3 and 7.5 mg/l (Neiff et al.
2011). The lakes are articulated with each other through chan-
nels of varied development, to finally resolve into a diffuse
drainage system in the headwaters of the Corrientes River.
This river transports water and information from the Ibera to
the Parana River but it is disconnected from the hydrological
pulses of this river. The water fluctuation is exclusively due to
the effect of local rainfall, which is relatively predictable on
annual and interannual bases. Water fluctuations are smoother
APPROACH (less) than in the lakes connected to Parana River. Water flows
Forests and aquatic communities of different individual are predominantly vertical, from and to the atmosphere with
sizes and turnover rates were analyzed in the present study, strong influence from the extensive vegetation (Neiff 2004).
ranging from phytoplankton and zooplankton to ben- There are laminar flows between the marshes and the large
thos and fishes. Using our own information and findings lakes and vice versa that provide a buffer against the drastic
published by others over the last decades (Table 1), we changes in the water table. The wetlands included in the Ibera
compared the two wetland systems: the current floodplain depression occupy 12,000 km2. They are gently concave (0.10
of Parana River and its paleo-floodplain (Ibera) that was to 2.5 m deep) with dense and continuous marsh vegetation
disconnected from the river 10,000 years ago. covering about three quarters of total surface. General NE-SW
To detect changes at ecosystem level and their compo- runoff is very slow and connects with the large lakes (Galarza,
nents, we use common indicators such as total number of Luna, Ibera, Fernández, Trin, Medina, and Itati) to finally
species cited, total abundance, and dominant taxa, especially discharge into the Parana River through the Corriente River
those marking functional differences. On this basis, we aim (Figure 2). These large lakes are 2.5-4.0 m deep and the water
to identify which wetland “compartments” changed and level fluctutates from 0.5 to 0.7 m throughout the year. The
which did not, how they were modified, and then try to ex- western border of Ibera contains low hills of sand deposited by
plain “why”. We analyze whether the causes of change lie in the Parana River and a gentle slope. Thousands of small lakes
habitat variability, the breadth of niches, or other factors. of 1-5 ha (Contreras et al. 2014) with a depth of 1.5-2.5 m are
Based on our previous palynological and paleoecologi- scattered across this area. Since their source of water, physico-
cal research, we confirmed the fluvial origin of the current chemical characteristics, vegetation and fauna are similar to
Ibera wetland system (Cuadrado and Neiff 1993; Morton those in the large lakes of Ibera (Neiff 2004; Poi et al. 2017)
2004; Pacella and Di Pasquo 2020). In the present work, we they are included in Group I.

FIGURE 3. Examples of Group I (Isolated) wetlands: 1) peatland forest FIGURE 4. Examples of Group E (eventually connected) wetlands (from
surrounding Galarza Lake, 2) sudds (floating islands) in Luna Lake, and left to right): 1) marginal riparian forest, 2) bulrush marsh (Schoenoplec-
3) “esteros” landscape in Ibera Lake. tus californicus, Cyperus giganteus, and others) around the lakes, and
3) submerged meadows (Egeria naias, Ceratophyllum demersum and
others) in lakes. Note: Inset E shows the geographic location of these
wetlands. (Source of base image: Google Earth.)
Eventually connected wetlands (Group E). These wet- FIGURE 5. Examples of Group C wetlands: 1) Parana River floodplain with
lands, including island levee lakes, are located on ancient shallow and connected wetlands, 2) oxbow lakes with Pistia stratiotes
riverine islands. The lakes are surrounded by marshes floating meadows, surronded by palm forests (in the distal area of the
floodplain), and 3) meander scroll covered by dense floating meadow of
included in high riverine islands originated by the old Parana water hyacinth (Eichhornia crassipes).
River. These islands are near Ituzaingó city (27°31’19”S,
56°42’55”W, Figure 4). They are situated almost 3 m above
the river course hence they are only connected to the river by
extraordinary floods, that is, once every ten years or more.
These occasional flooding events trigger an exchange of
information (nutrients, organisms, seeds, eggs, etc.) between
the lakes and the Parana River. Most of the time, however,
the lakes are fed by rainfall. The local landscape is very
similar to the Ibera region, at least in the last 3,000 years
(Cuadrado and Neiff 1993). Lake waters show very low con-
centrations of suspended solids and a black-brown color due
to the high concentration of dissolved organic matter.
Close connected wetlands (C). These wetlands include
shallow lakes, oxbow lakes and ponds that occur on recent
lateral riverine islands that emerged in the last few centuries.
They are part of the active Parana floodplain and fed by river
overflows at least once a year. These waterbodies are located
PHYTOPLANKTON
in the tract comprised from the wetlands in Group E to the
Phytoplankton composition shows relevant differences
south, at Itatí city (27°15’34”S, 58°14’ 35”W; Figure 1).
between the Ibera lakes (Group I) and the lakes of the cur-
Silty-sandy sediments and “white waters” (with suspended
rent Parana floodplain, with ten times more species in Ibera
silt, fine sand and clay) predominate. The most frequent veg-
(796 species found by Zalocar de Domitrovic 2003) than in
etation is free floating and reed swamp plants. These lakes
Parana floodplain. In wetlands of Group E and C, density
have a high turnover of plant and animal organisms with dif-
and diversity of phytoplankton decrease during the con-
ferent phases of the river pulses. The water in these wetlands
nection periods in relation to the disturbance and dilution
is similar to that of the Parana River.
produced by the river water entering into the floodplain.

Chlorophyta is the most important group in Ibera lakes (Cózar et al. 2003). Spatial differences in species richness
(Table 2, Group I), while Bacillariophyceae was more im- were less than 38% in the Ibera lakes and the variation
portant in the floodplain lakes of the Parana River (Table 2, between high rainfall and dry periods was only 6%. The
Groups E and C) during the connection period (Zalocar de rotifers Lecane proiecta and Filinia sp. were very abundant
Domitrovic et al. 2007). in severe droughts.
Phytoplankton density in Ibera is highly dependent on In Parana River floodplain (Group C, Table 3), zoo-
the type of environment where the sample is taken, ranging plankton abundance is related to seasonal hydrological fluc-
from 100 to more than 4000 cells/ml. In wetlands of Parana tuations, with higher concentration at the end of the low-
River (Groups E and C), the density can vary between water seasonal period (usually in spring) and lower density
1000-2020 (Zalocar de Domitrovic 1990; 1992) or, 588- during the high-water peak, due to dilution effect. Density
2598 cells/ml (Zalocar de Domitrovic et al. 2007) depend- values are often between 1 and 75 individuals/L in the river
ing on the hydrological phase considered (Table 2). proper (Paggi and José de Paggi 1974; Corrales 1979), al-
though these values can double in floodplain lakes (Bonetto
ZOOPLANKTON 1986a). Rotifers are numerically dominant at all times, and
In Group I, the species richness ranged from 7 to 41 (Frutos cladocerans and copepods alternate as subdominants, both
2003, 2017), with the highest species richness in lakes with in considerably low abundance. Despite the difference in
submerged vegetation. Zooplankton abundance increased diversity, the dominant species are similar to those in Ibera.
in summer (50-450 individuals/L) and decreased in win- For instance, the most abundant rotifers include Keratella
ter (20-350 individuals/L). Rotifers of the genera Kera- cochlearis, Trichocerca similis and Poliarthra trigla. The
tella, Ptygura, and Trichocerca were always numerically main difference with the lakes of Group I is the occurrence
dominant (80-95%). Cladocera and copepods had variable of cladocerans (e.g., Ceriodaphnia cornuta, Diaphanosoma
representation in the samples (Cózar et al. 2003; Frutos bracyurum, and Eubosmina hagmanni) and copepods (e.g.,
2003, 2008). The low density of cladocera and copepods Notodiaptomus incompositus and Mesocyclops longisetus,
in the Ibera lakes is reportedly due to high fish predation among others) in wetlands of Group C
FIGURE 6. Daily water fluctuation at Ibera lakes (red) and Parana River (black) measured at Itatí city from 1970 to 2019. The straight black line shows the
overflow level for Ibera lakes (black), while the dashed lines show overflow levels at Parana River – the lower line indicates overflow into Group C lakes,
while the upper line represent the level at which lakes in Group E are flooded with water from the Parana River. The latter lakes are rarely overflowed.

BENTHOS
The bottom substrate is an important factor influencing the TABLE 2. Phytoplankton in Ibera lakes (grouped as I, isolated) and lakes of
benthic fauna. In Group I lakes, the bottom fauna is mainly current Parana floodplain (grouped as E and C, eventually and closely con-
determined by the presence of organic detritus. Since all nected respectively). Data of the taxonomic group (expressed as relative
lakes have a sandy floor, the limnetic area generally has a abundance (percentage of total abundance), species richness (total number
mobile bottom due to wind effect, so the benthic fauna is of species) and abundance range (cells/ml).
not very abundant. In the littoral zone, or in sites with dense Taxonomic group (%) I (isolation) E and C
submerged vegetation, however, the fauna is more abun- or Other metric (eventually
dant and shows a greater number of species (Varela et al. and close con-
1983; Bechara et al. 1990; Casciota et al. 2005). In Parana nection)
floodplain lakes (Groups E and C in this study), the bottom
is sandy and dynamic due to the current. In the wetlands Chlorophyta 65.7 49.90
of Group C, 75 species were recorded, and in the wetlands Bacillariophyceae 11.93 26.95
of Group I (without connection to the river) 67 species are Euglenophyta 11.80 6.50
mentioned (Table 4). The abundance of fauna is much more
variable in wetlands connected to the river course, which Cyanophyta 6.91 8.51
have a greater amount of Oligochaeta, while in Group I Xanthophyceae 1.63 ----
lakes (Ibera) Chironomidae is the dominant taxa, especially Chrysophyceae 0.75 1.63
in sites with submerged vegetation. Chryptophyta 0.75 6.51
VEGETATION Dinophyta 0.50 ----
Of the 115 macrophytes cited for the floodplain of Parana Relative abundance 100% 100%
River (Neiff 1986), only seven species are not found today Total abundance range 140-4,033 588-2,598
in the wetlands of Ibera and they are members of the Pod- (cells/ml)
ostemaceae - aquatic plants typically growing in the river
Species richness 796 74
rapids (habitats that do not exist in Ibera). These plants are
only found in flowing waters and form a very specialized (Data Sources for Group I: Zalocar de Domitrovic 2003; for Group
community known as “tachyrheophyton” (Neiff 1986). E: Zalocar de Domitrovic 1990 and 1992; and for Group C: Zalo-
In a more recent study (Neiff et al. 2011), 161 species car de Domitrovic 1992; Zalocar de Domitrovic et al. 2007.)
were catalogued for the active Parana River floodplain. They
also account for 40% of the total species reported for float-
ing islands and marshland vegetation in Ibera (400 species)
based on extensive surveys and historical records in herbaria
(Arbo and Tressens 2002). The comparison is especially TABLE 3. Zooplankton in the lakes of Ibera (group I, isolation) and shallow
significant since the main environmental difference between lakes of current Parana floodplain (group C, closely connected). Data of the
Ibera (Group I) and Parana River floodplain (Groups E and taxonomic group (expressed as a percentage of total abundance, %) and
C) is related to the regime of pulses, namely amplitude, fre- abundance (individuals/L).
quency and predictability of the water level fluctuations. Taxonomic
Zooplankton fauna in ecological
Table 5 shows the biological spectrum of the vegeta- group or Other
groups according its connectivity
tion for Groups I and C, for which information of similar metric
amount and quality is available. The total number of spe- C
cies in both groups does not differ by much. The difference I
(Closely
is the distribution of species richness in each plant bioform, (Isolated) connected)
which is due to the different geomorphology of both wet- Rotifers 79.36 63.5
land groups and the variability of the pulse regime. In the
Ibera depression (Group I) the habitat favors the develop- Cladocerans 14.28 17.22
ment of emergent plants (helophytes) that have rhizomes; Copepods 6.36 19.28
they dominate the marsh landscape. In Group C reed bed Abundance 20-450
plants are successful because they are highly resilient to 2-88
(individuals/L)
the irregular hydrological regime of the river (Neiff 1978,
(Data Sources: CECOAL 1981; Frutos 2017.)
1990). The number of free-floating plant species is similar

in Groups I and C, and the species recorded are common The main difference of the vegetation of Ibera in rela-
to both (Table 5). However, this group of plants achieves tion to that of Parana River is the area occupied by her-
greater coverage in the wetlands of active Parana flood- baceous and woody vegetation. On the islands of Parana
plain, due to the periodic input of nutrients during the River, forests cover about 10 to 15%, while the forest area
annual and more frequent floods (Carignan and Neiff 1992; is less than 1% in Ibera where the vegetation is virtually all
Carignan et al. 1994). Although these plants can be found herbaceous. Of the 15 tree species growing in the Parana
in Ibera, they never reach 1% coverage in the lakes. On the gallery forests, only five species are found in small patches
other hand, submerged plants form extensive meadows in on organic (peat) or mineral soils (sand) in Ibera (Figures
the Ibera lakes, while their presence is sporadic and limited 7 and 8). The architecture of these trees is very different
to Group E in wetlands of the Parana floodplain. when they grow in the peaty soils of Ibera: the trees are less
than 8 m high and their roots are distrib-
TABLE 4. Composition of lake bottom fauna in the study area. Data of the taxonomic group (ex-
uted radially in the first 20 cm of the soil
pressed as a percentage of total abundance, %), species richness (total number of species) and to avoid anoxia (Neiff and Casco 2017).
abundance range (individuals/m2). Overall isolation of Ibera has created
Taxonomic group or Benthos fauna in ecological groups a different environment – a lentic one –
Other metric according its connectivity that supports rooted hydrophytes, while
the Parana River favors free-floating
I C plants and floodplain forests (Figures 9
(Isolated) (Closely connected) and 10).
Oligochaeta 52 43
INVERTEBRATES ASSOCIATED WITH
Chironomidae 41 54
AQUATIC VEGETATION
Ostracoda 3 1 In extensive surveys that include several
Amphipoda+Turbelaria + 4 2 species of aquatic and marsh plants, 152
Acari + Mollusca morph species of invertebrates have been
recorded in Parana River floodplain (Poi
100% 100% de Neiff and Neiff 2006) and 98 in Ibera
Abundance (individuals/m2) 5,000-10,000 1,000-100,000 (Poi de Neiff 2003). In both surveys
Species richness 67 75 identical techniques were employed on
seven of the most frequent aquatic plants
(Data Sources: Varela et al. 1983; Bechara et al. 1990; Casciota et al. 2005; in wetlands (Eichhornia crassipes, Eich-
Zilli et al. 2008.) hornia azurea, Pistia stratiotes, Salvinia
biloba, Azolla caroliniana, Lemna gibba,
TABLE 5. Plant bioforms in wetlands with different connectivity (expressed as species richness). and Paspalum repens) and on five species
Emerging plants are always emerging (bulrush, cattails); “Reed bed plants” have life forms in lakes of the large Ibera wetland (Typha
adapted to flooded soil (floating rooted form) and to the emerging soil phase (emerging rooted latifolia, Leersia hexandra, E. azurea,
form). Egeria najas, and Cabomba caroliniana).
Vegetation in ecological groups according its connectivity It is difficult to compare abundance
I C and composition of invertebrates to
(Isolated) (Closely connected) investigate the effects of river connectiv-
ity, because different bioforms of mac-
Emergent (cattail type) 76 38
rophytes were dominant in the Parana
Reed bed plants 18 29 floodpain versus the Ibera lakes. Each
Free-floating plants 9 10 plant bioform (submerged rooted, free
Rooted submerged plants 7 3 floating, or emergent rooted) provides
a different habitat for invertebrates. In
Free submerged plants 6 3
floodplain habitat, both density and
Rooted with floating leaves 6 6 species richness are influenced by hori-
Trees on mineral or peat soils 5 15 zontal flows to and from the river course.
TOTAL (Species richness) 127 104 Surveys of a floodplain lake with high
connectivity to the High Parana (Sirena
(Data Sources: Neiff 1986, 1990, 2003; Neiff and Casco 2017.)

Lake with floating meadows dominated by Paspalum rep- FIGURE 7. Gallery forest of Parana River (Group C) near of Ituzaingó: 1)
ens and Salvina biloba; Poi de Neiff 1981) and two Ibera high, closed, continuous forest, up to 20 m tall, and 2) Trees spaced by 4-6
lakes (Galarza and Trin with dominance of S. biloba and m each, DBH 0.30-0.90 m; shrubs and grasses are scarce or absent as a
Eichhornia azurea; Poi de Neiff 2003) recorded 82 morph result of frequent flooding.
invertebrate species and 61, respectively. The comparison
of similar types of habitats confirms a higher taxon richness
in the lake connected to the river than the isolated lakes of
Ibera. Depending on the site and the mesh size (Table 6)
the overall abundance varied between 18,388 and 72,056
individuals/m2 in Ibera and High Parana, respectively.
Macroinvertebrates (> 500 µm) associated with the
aquatic plants were dominated by oligochaetes (mainly
Naididae), insects and copepods (Table 6) both in High
Parana and in Ibera. When smaller invertebrates (size great-
er than 125 µm) were included, copepods had the highest
relative abundance in Parana and cladocerans in Ibera wet-
lands. Copepod species were also recorded in the plankton
of the more connected lakes (Poi de Neiff 1981; Table 3).
Cladocerans typically associated with vegetated areas, such
as Diaphanosoma, Euryalona, Oxyurella, and Euricercus,
were registered in Ibera (Poi de Neiff 2003). Mollusks and FIGURE 8. Peatland forest in Group I: 1) irregular, low forest, up to 8 m high,
mites were poorly represented, especially in Ibera. At both with species as the Parana riverine forest, although with less diversity, and
2) irregularly shaped trees separated by 4-6 m (DBH 0.15-0.30m) with
sites, the composition of insects was similar. Larvae of two
dense herbaceous vegetation up to 2 m high.
families Ceratopogonidae and non-biting midges (Chiron-
omidae) were the most abundant insects. Air fronds of S.
biloba supported semi-aquatic species such as the grasshop-
per Paulinia acuminata that has a high specificity to this
host plant. There was a high number of genera of Coleop-
tera (Helochares, Enochrus, Derallus, Tropisternus, Para-
cymus, Berosus, Hydrochus, Desmopachria, Laccophylus,
Liodessus, and Hydrochanthus) and Hemiptera (Belostoma,
Pelocoris, Neoplea, and Ranatra) at both sites.
As described above, submerged plants form extensive
meadows in the Ibera lakes. The freshwater prawn Pseudo-
palaemon bouvieri (Decapoda) is adapted to freshwater oligo-
haline waters covered by submerged vegetation. It is restrict-
ed to Ibera (Group I) and other water bodies of the Corrientes
province (Lopretto 1995) fed by rain; this prawn has not been
reported for the Parana River floodplain (Group C).
FISH FAUNA
According to Bonetto (1986b), the fish fauna of this area of Some assemblages of the fish fauna are considered
the Parana River contains about 200 species and does not “sedentary fauna” - smaller fishes that live in ponds and
differ much from that of other large South American rivers. floodplain wetlands on the islands of Groups E and C.
As in other floodplain rivers in South America, characi- Another group of species is the potamodromous (migra-
forms (e.g., toothed fish) comprise almost 40% of the river tory freshwater) fishes that as adults (1-2 m long) make
fish, with many species of Tetragonoptera. Silurids (catfish) extensive migrations upstream in spring and downstream in
make up 20% or more of the total taxa with some being late summer (Bonetto 1986). Available information shows
quite large fish. For example, “surubí” (Pseudoplatystoma that potamodromous species have their immature states
coruscans; a long-whiskered catfish) reaches 2 m in length living in lakes of Group C (Bonetto 1986b; Casciota et al.
and may weigh 120 kg. 2005; Iwaszkiw 2010; Contreras et al. 2017). These species

FIGURE 9. Parana River wetlands: 1) panoramic view of the vegetation on a meander scroll include Prochilodus lineatus, Pterodoras
(Group C) and 2) floating meadow of water hyacinth (Eichhornia crassipes) and other species. granulosus, Oxydoras kneri, Trachydoras
paraguayensis, Serrasalmus spilopleura,
S. marginatus, Pygocentrus nattereri,
Hypostomus latifrons, Loricariichthys
melanocheilus, Schizodon borelli, Lepori-
nus lacustris, Pachyurus bonariensis,
Triportheus paranensis, Odontesthes pe-
rugiae, and Potamotrygon motoro. Some
larger fish (Salminus brasiliensis, Pseu-
doplatistoma coruscans, P. fasciatum,
and Lucypimelodus pati) of the Parana
River are migrants each year during high
flows that occasionally also go upstream
via the Corriente River to the southern
lakes of the Ibera (Itatí, Medina lakes);
they are, however, not found in the
isolated lakes of northern Ibera (Galarza,
Luna, and Ibera lakes).
In Group E lakes, fishes associated
with littoral, emergent or rooted floating
vegetation are very common (Iwaszkiw
et al. 2010). These fishes include Poptella
paraguayensis, Hyphessobrycon eques,
Moenkausia spp., Acesrorhynchus pan-
TABLE 6. Relative abundance of the main taxa (%) and overall abundance expressed as individu-
taneiro, Hypostomus latifrons, Hypopto-
als/m2 of macro- (>500µm) and meso-invertebrates (>125µm) in the Galarza, Sirena and Trin
lakes. (Sources: Reconstructed from Poi de Neiff 1981, 2003 data) poma inexpectata, Cichlasoma dimerus,
Gymnogeophagus balzanii, Apistogram-
Ibera Ibera High Parana High Parana
ma spp., Crenicichla spp., Hoplerythri-
>500µm >125µm floodplain floodplain
nus unitaeniatus, Hoplias malabaricus,
>500µm >125µm
Hoplosternum littorale, and Lepthoplo-
Oligochaeta 25 10.5 33 24 sternum pectoral. Many of these fish are
Cladocera 5 36 1.5 9 also found in Group I where the most
frequent fishes are Hyphessobrycon
Copepoda 15 25 18.5 29 eques, Moenkausia spp., Acesrorhynchus
Amphipoda 1 0.5 4.5 26.5 sp., Cichlasoma dimerus, Gymnogeopha-
gus balzanii and Apistogramma spp. The
Ostracoda 1 + 5 6.5 migratory fish of Parana River (Salminus
Insecta 45 18 31 18 brasiliensis, Pseudoplatistoma corus-
cans, P. fasciatum; Lucypimelodus pati)
Mollusca + + 1.5 0.5 are of occasional presence, as they enter
Hidrachnidia + 0.5 4 2.5 some lakes only during the extraordinary
floods of Parana River.
Other taxa 8 9.3 1 8 As shown in Table 7, fish fauna
100% 100% 100% 100% appears similar across the region regard-
Mean overall 18,388 47,494 38,096 72,056 less of connection to the river. Although
abundance the percentage of characiforms is higher
individuals/m2 than that mentioned by Bonetto (1986b),
the percentage of species included in the
The + sign indicates that the taxa were present but in a very low percentage (<0.5) different taxonomic groups has a similar
proportion in the three connectivity vari-

ants (Groups I, E and C). Of the 200 species reported by exclusive to Group I. Perhaps time (3,000-4,000 years) was
Bonetto (1986a) for the Upper Parana River, there are 111 not sufficient or the differences in environmental conditions
species in Ibera (Casciotta et al. 2005). This total accounts (e.g., climate and water quality), except for flooding and
for those upstream-migrating species found in lakes and soil saturation, were not significantly different between the
streams of the southern Ibera basin (Group I) which are Groups.
still linked to Parana River, at south of Itatí lake (Figure 2). For some components of the system (Group I) such
However, in the lakes of northern Ibera, which are com- as phytoplankton, there was even an increase one order
pletely isolated from the pulses of the Parana River (lakes in magnitude in species richness, due to the lower rate of
Galarza, Ibera and Luna), the same authors reported only water renewal resulting from lower water level fluctuations.
51 species, reflecting the impact of isolation from the river. Despite this, there is no unique species in Group I. All the
The loss of connectivity of the wetlands of Ibera (Group I) species recorded there were found in other environments in
has resulted in a reduction of the number of fish species to the Parana River basin.
a third or a quarter and a loss of species of large migratory In the Group I, more than 80% of the marsh area is
(potamodromous) fish that are restricted to the Parana River occupied by plants with rhizomes (helophytes). This type
and its active floodplain wetlands. of macro-vegetation in the lacustrine scenario is surpris-
ingly not affected by flooding because the entire marsh
COMPARISON OF FLUCTUATION REGIMES UNDER DIFFERENT with its peaty soils simply floats, rising with the increase in
CONNECTIVITY 10K YEAR LATER water level (i.e., floating islands called “embalsados”) and
Daily data on water level fluctuations have been collected falling when water levels decline. The marsh vegetation is
since 1929. The fluctuation of the water level in wetlands sensitive to prolonged dry periods, yet since most of the
of Group I has maintained a pulse regime as a consequence species are helophytes with rhizomes that are able to root in
of the seasonality of the rains (Neiff 2004). The range of the substrate, allowing the plants to persist during the dry
fluctuation between maximum and minimum absolutes was period.
close to 1.5 m (Figure 6). The pulse rate in Parana River is Most of the forests species in Parana River area are
very irregular and the extreme range of fluctuation between adapted to flooding periods. The floodplain trees lack ad-
highs and lows is greater than 8 m historically. When the aptations for survival during periods of prolonged dryness.
water level reaches around 3.5 m, Parana River spills over This fact likely explains both the small area of forest and its
into many of the lakes classified as Group C, receiving new low species richness.
water and information. If the riverwater level exceeds 9 m
line, the lakes of Group E are then flooded;
since 1970, this happened in 1983 and FIGURE 10. Lentic habitats of Ibera wetlands (Group I): 1) view of open water of the Ibera Lake
again in 1992 (Figure 6). and 2) marsh vegetation in littoral zone.
The pulse regime in wetlands indicates
horizontal movements of water from the
river and allows some horizontal circula-
tion of information (nutrients, eggs, seeds,
plankton, etc.). Water turnover produces re-
newal in biotic assemblages through water
circulation but also through change in habi-
tat conditions (e.g., transparency, oxygen,
and nutrients). In Group I (total isolation; at
least for lakes at north of Itatí Lake) there
is no exchange of information with the
river, nor is there a washout effect, and less
variability of the habitat is maintained, fa-
voring the permanence of a greater number
of species, with the taxonomic configura-
tion of Amazonian origin, especially in the
plankton, benthos, aquatic and marsh veg-
etation. Interestingly, the isolation did not
manage to produce any endemic species

ecosystems (Cabrera and Willink 1973)
TABLE 7. Percentage of fish fauna represented by taxonomic group for the three wetland groups.
and, currently, as an ecological corridor
Fish fauna in ecological groups according
Taxonomic group for many species (Bonetto 1986a; 1986b;
its connectivity
Giraudo and Arzamendia 2003), it also
I E
C appears to have functioned as a meridian
(Closely
(Isolated) (Eventually connected) barrier to the dispersal of some birds as
connected) Thamnophilus caerulescens, Cyclarhis
Characiforms 66 46 63 gujanensis, Thraupis sayaca, L. an-
Siluriforms 15 27 23 gustirostris, and Colaptes melanochloros
(Kopuchian et al. 2020). According to
Perciforms 10 16 9
these authors, large rivers as Parana,
Gymnotiforms 4 4 3 function as a barrier to genetic flow in a
Cyprinodontiforms 4 4 - transverse direction between both banks
Pleuronectiforms - 2 1 for some terrestrial birds, leading to
population differentiation and, ultimately,
Beloniforms 1 1 1
allopathic speciation. Isolation changed
Total 100 100 100 the landscape pattern with greater variety
(Data Sources: Casciota et al. 2005; Iwaszkiw 2010; Contreras et al. 2017.) of habitat (large lakes, marshes, peat-
land areas, running waters, and riparian
DISCUSSION forests), and greater species richness in
After 3,000-4,000 years, it appears that the natural system some communities (plankton, benthos, and aquatic plants)
of Ibera has maintained a part of its original configuration and simplification in others (disappearance of migratory
and adapted some elements and processes to a new habi- fish and some tree species from the riparian forests).
tat configuration, according to its capacity of self-design The increased complexity that arose in Ibera from
(Mitsch and Jørgensen 2003; Odum and Odum 2003). isolation can be explained from different perspectives: 1)
Since then Ibera has remained isolated from the Parana the biogeographical context, 2) the extent of the niche of
River with changes in the biota due to dry period climate the resident species, 3) the nature of change in the envi-
changes in the Lower and Middle Holocene, and a progres- ronment, and 4) the ability of the system for self-design.
sively wetter period in the last three or four thousand years From the first point, we consider that the impact of loss of
in the recent Holocene. Previous studies have demonstrated floodplain connectivity to the river becomes particularly
that the Parana River fed Esteros del Ibera produced a significant if the Ibera wetland system constitutes a species-
similar pattern of the landscape that was maintained over at endemic area. For this system, all species belong to the
least three thousand years ago for the entire Group I (Neiff vast Amazon domain, which has remained a biogeographic
2004; Orfeo and Neiff 2008; Pacella and Di Pasquo 2020). dispersal center even during the glacial period. Since the
Although sedimentological and geomorphological evidence Amazon bioprovince is among the most species-rich in the
has shown that the Ibera marshes (Group I) were isolated world, the lack of endemics is not surprising especially
from the Parana River 10,000 years ago (Castellanos 1965; when wetland species generally have very broad niches.
Orfeo 2005; Orfeo and Neiff 2008; Orfeo et al. 2014), paly- Despite this, the ecology of the Ibera has changed com-
nological information shows that the current landscape cor- pletely as a consequence of its isolation, with a greater
responds to a recent humid tropical phase from 3000-3500 extension and variety of lentic environments, although
years old (Cuadrado and Neiff 1994; Pacella and Di Pas- frequent species have disappeared from the river habitat of
quo 2020). The same authors agree that there is no pollen the Parana.
evidence of older landscapes. In this contribution we point The breadth of the ecological niches also plays a key
out that, although Esteros del Ibera was isolated from the role in the pathway for the ecological change. Since the
Parana River for 10,000 years, experiencing long periods Upper Pliocene, the climate of the subtropical zone of
of dry and wet weather, they were able to maintain a fairly South America has gone through very contrasting wet and
similar assemblage of plant and animal species although dry periods (Iriondo 2004), driving a selection of plants and
not all species were able to adapt to the new conditions. animals adapted to the irregular water regimes and drastic
While the Parana River in its north-south direction has changes in the configuration of their habitat. Thus, an im-
served as a vector for the dispersal of genetic information portant number of fluvial species remain in the Ibera lakes
from the Amazon region to Parana River and surrounding (Group I) after 10,000 years of isolation and there is no re-

cord of species unique to Ibera. An example of the ecologi- lands changed from a lotic system dominated by river over-
cal plasticity of the wetlands in the region occurred in 1995, flows to a lentic system of lakes with fluctuations due to
when the Yacyretá Reservoir was constructed by covering local rainfall patterns. It is evident, for example, that Ibera
the marshes of ancient islands in Parana River (Group E) wetlands (in Group I) have a greater number of Cyperaceae
with an eight-meter-thick water layer. After seven months and other helophytes unlike riverine wetlands (Group C)
of being completely submerged, extensive “islands” of that have a greater occurrence of free-floating plants. The
several kilometers of peat rose to the surface. Only 22 days species of trees that live in Ibera are smaller than those on
later, the herbaceous and shrubby vegetation had sprouted the islands of Parana River and have a highly developed
from the buds of the plants that were lying on the surface root system in the shape of a big dish, which allows them to
looking “dead”. It is expected that the floating islands of be supported on organic soils (peat). The process of self-
Ibera formed in a similar manner, thereby maintaining most design seems to have selected species that are extremely
of the plants found on the former Parana floodplain. tolerant of climatic change, which means that the structure
Ibera has experienced strong climate disturbances in of the Ibera landscape and its biotic components are main-
the past, and global climate change is expected to have an tained with a low rate of change in spite of extreme climatic
impact on the region. We studied the possible effects on events of drought and extraordinary waterlogging (Neiff
the Ibera lakes (Group I) under two future climatic sce- et al. 2011; Ubeda et al. 2013). Yet, at the same time, the
narios (A2 and B2) proposed by the International Panel on lower variability of the water sheet and the lower flow rate
Climate Change (IPCC; Neiff and Neiff 2013; Ubeda et has favored the increase of numerous species of plankton
al. 2013). Our results suggest that even though a reduction and benthos that take advantage of the microenvironments
in lakes size could have negative effects on biota, affect- of the waters with different types of vegetation.
ing richness species at local scale (Ubeda et al. 2013), the The P/R ratio is higher in Ibera wetlands than in the
biodiversity will not be significantly affected (Neiff et al. wetlands of Parana River resulting in the accumulation
2011; Neiff and Neiff 2013; Ubeda et al. 2013). The vast- of organic matter in the Ibera marshes (“esteros”) that
ness of Esteros of Ibera wetland complex, with its huge surround the lakes. This organic matter, although slowly
variety of habitat types and ample niches for most species degraded, releases substances that are recycled by the veg-
make it highly resilient from the biodiversity standpoint. etation of the lakes.
We believe that knowledge of the breadth of niches and the At least in the time scale of our analysis, biodiversity does
resilience of the landscape are key aspects for the scientific not seem to be a powerful indicator to evaluate the effect of the
assessment of global climate change risks.
The pulse regime has become more regular, FIGURE 11. Seasonal changes in the water levels for the Ibera wetlands (Group I) and
showing shorter range of water fluctuations the floodplain of Parana River (Group C). The meters represent the vertical variability of
after isolation from the river (Figures 6 and the water sheet in each gauge station. In Group I the seasonal fluctuation is lower than in
11). Water in Ibera comes from rainwater that Group C because it responds to the variation of local rainfall that occurs over a wide area.
The vegetation of the marsh increases the roughness of the surface and decreases the
has been draining through sand for thousands runoff speed.
of years. However, the quality of Ibera waters
is not very different from the waters of Parana
River: low electrical conductivity (EC), slightly
acid to neutral pH, low nutrient content (espe-
cially nitrogen) and the ionic balance is of the
type: HCO3-> Na+> Cl-> Mg+> SO4-> Ca+>
K+. The water exchanges between the marshes
and Ibera lakes determine the contribution of
chemical substances from the organic soils
(Neiff 2004; Ubeda et al. 2013; Poi et al. 2017).
Ibera’s self-designed response to loss of
connectivity is interesting. Nature has selected
those tree species that can persist in water-satu-
rated soils for long periods and go through pro-
longed dry spells - trees that can occupy loose,
acidic soils (pH 4.5), such as peat or sandy soils
with very low nutrient concentration. Ibera wet-

loss of connectivity. We know that all the biota of Parana River out. In the case of Ibera, there was a drastic change in the
comes from the Amazon mega-basin where extensive flood- hydrological regime with a significant attenuation of the vari-
plains have remained since geological-evolutionary times. So ability at interannual and seasonal scales (Figures 6 and 11).
what biotic changes have occurred? The dominance of rotifers This change resulted in increased species richness for most
in the potamoplankton could be a consequence of isolation. biotic assemblages over the millennia time scale (plankton,
Loss of connectivity with Parana River during Upper Pleisto- benthos, and aquatic vegetation), except for forests where the
cene produced wetlands with an increase in different forms of number of species was less than ithe Parana gallery forests.
organic matter, which is the favorable habitat for rotifers.
We think that the analysis of connectivity in terrestrial CONCLUSION
ecosystems based on topological relationships (as the spa- Generally, the connectivity of basin landscapes is analyzed
tial proximity of landscapes or as exchanges between popu- on a current scale, or that of the recent past, without putting
lations or landscape gene banks) is inappropriate in river into a biogeographical context the events and changes that
systems, due to differences in the response mechanisms occur in the support system (physical-chemical environ-
of individual organisms and landscapes to changes to the ment), focusing the analysis on the effects of engineering
pulse regime and variables associated with river connectiv- works on the stability of riverine wetlands (e.g., damming
ity. While in a terrestrial native forest the loss of landscape of rivers or channelization of watercourses). Undoubtedly,
continuity is seen as fragmentation and increased distance any human action on the ecosystems produces disturbances
between patches, in a floodplain of large rivers it is normal that can alter the local, regional or global nature in different
for the natural design of the landscape to include numerous ways. Our challenge is to understand the impact of natural
patches of forest in the form of “galleries” or “patches” in changes in connectivity between landscapes in a basin. We
the landscape matrix due to differences in the topographi- have compared a scenario in which the isolation of riverine
cal position. This determines different eco-hydrological wetlands occurred naturally 10,000 years ago. While the iso-
connectivity and, consequently, different assemblages of lation of Ibera has clearly created a lentic environment on the
species, such as separate cells in the landscape. former floodplain and a decline in fish species, this species
We have a difficult challenge in the study of the niches segregation affected only the northern lakes of Ibera, because
(in Hutchinson`s sense) of plants and animals, in order to large migratory fish need to migrate in order to reproduce.
assess the relationships of river connectivity in different We have not recorded the appearance of unique species typi-
scales of time and space. Although the hydrological regime is cal of the new situation of isolation. Obviously, the absence
one of the main characteristics that condition and define the of rapids in Ibera justifies the disappearance of typically
functioning of aquatic systems, there are other attributes that rheophile species such as those of the Podostemaceae family
determine the character of wetlands, as Tiner (2017) pointed cited for Parana River (Neiff 1986). Nor have we recorded
in Ibera the presence of any invasive species
FIGURE 12. Synopsis of the changes produced by the disconnection of “Esteros de Ibera” that occur in the Parana River in recent decades,
from the water regime of Parana River. such as the golden mussel (Limnoperna fortu-
nei) or the tilapia (Tilapia niloticus). We can
think that the structure and functioning of the
Ibera macro-wetland has retained its biodiversity
and as a result of isolation, it will resist biologi-
cal invasions, at least from riverine species. As
a synthesis we present Figure 12 with the most
notable changes in the long-term scale. n
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558-566.

WETLAND RESEARCH
Urban Wetland Trends in Three Latin American Cities during the Latest Decades
(2002-2019): Concón (Chile), Barranquilla (Colombia), and Lima (Peru)
Carolina Rojas1,2, Juanita Aldana-Domínguez3, Juan Munizaga4, Paola Moschella5, Carolina Martínez6, and Caroline Stamm7
ABSTRACT zones, and 3) the assistance of wetlands when facing

etlands are valuable but threatened natural resources frequent urban disasters has not been considered. It is a
W worldwide. While providing a wealth of environ-
mental benefits, wetlands play a vital role in temporarily
fact that wetlands mitigate the impact of flooding, which
helps make cities more resilient (Ramsar 2019). In rela-
storing flood waters and thereby reducing the risk of dam- tion to this, according to the United Nations (2014), 233 of
aging floods. This is important given the predicted impacts the world’s cities are located in zones that are high at risk
of climate change, especially along the world’s coastline for flooding, affecting approximately 663 million people.
and coastal cities. The continued expansion of urban areas These urban areas require precaution and more resilient
is posing a risk to wetlands in and around metropolitan infrastructure. The importance of wetlands could be even
areas. In this article we examine wetland trends in urban greater when facing extreme events, such as the flooding
areas in three Latin American countries – Chile, Colombia, that occurred in Phoenix, USA in 2014 (Kim et al. 2017). In
and Peru. fact, in the northeastern USA, wetlands helped save $625
million dollars in direct damage from the floods caused by
INTRODUCTION Hurricane Sandy in 2012 (Narayan et al. 2017).
Wetlands, including their associated vegetation and the One of the main ecosystem services of wetlands is
water bodies, cover at least 10% of the planet (Davidson protecting the coast. They are also important for cities, as
et al. 2018). They are disappearing, even though they are they act as carbon sequestering systems, purify water, and
relevant ecosystems for ecological balance and for mitigat- maintain biodiversity and ecological processes. They even
ing the effects of climate change. It is estimated that since provide a place for recreation and relaxation for urban resi-
1900, the world has lost more than 50% of these ecosys- dents (Maltby and Acreman 2011; McInnes 2014). Without
tems, and the increase in urbanization has been identified a doubt, the loss of wetlands is affecting the sustainability
as one of the main causes for this loss (Boyer and Polasky and resilience of our Latin American cities. This is espe-
2004; Faulkner 2004; Bishop et al. 2006; González et al. cially true for coastal cities that face the challenge of being
2014). In fact, more than 55% of the world’s population prepared for the increase in frequency of natural disasters,
lives in cities, and it is expected that by 2050, this figure such as urban flooding. Ramsar (2018) also highlights the
will reach 68% (United Nations 2018). As urban growth role of wetlands in cities by decreasing the impact of urban
increases, wetland area decreases. Unfortunately, Latin flooding caused by strong rainfall, as well as providing a
America leads this worldwide tendency, reporting a loss of buffer from swells and tsunamis. However, Latin America
59% of wetlands over the last decades (1970-2015) (Dar- is dealing with a loss of wetlands, despite being one of the
rah et al. 2019). This loss is combined with the fact that it is regions most exposed to flooding (UN 2011). Evidently,
one of the poorest and most economically unequal regions this tendency also increases the region’s vulnerability to
on the planet (Cepal 2019). climate change (Seto et al. 2011; Hallegatte et al. 2013).
Wetlands are particularly relevant for cities because: 1) We are facing an alarming scenario, where sustained
most large cities are located on the coast, 2) world urban urban growth is the generalized trend in Latin American cit-
growth has been concentrated in low-elevation coastal ies (UN 2018) which in many cases occurs at the expense
of natural spaces (Rojas et al. 2013; Aldana-Domínguez et
1 Institute of Urban Studies, Pontificia Universidad Católica de Chile, Santiago, Chile. al. 2019). Facing pressure to expand, cities have converted
2 Correspondence author contact: carolina.rojas@uc.cl.
3 Department of Chemistry and Biology, Universidad del Norte Colombia, Bar- wetlands to developable land by legal and illegal landfills
ranquilla, Colombia. as is the case in Chile and Argentina (Rojas 2018; Pintos
4 EULA Center, Universidad de Concepción, Concepción, Chile.
5 Department of Humanities - Geography and Environment Section, Pontificia
and Sgroi 2012). Legal clearing is protected by urban laws
Universidad Católica del Perú, Lima, Perú. that do not incorporate factors such as ecological con-
6 Department of Geography, Pontificia Universidad Católica de Chile, Santiago, Chile. nectivity, integration with the coast, or geomorphological
7 Institute of Urban Studies, Pontificia Universidad Católica de Chile, Santiago, Chile.

TABLE 1. Selected images per wetland study area. processes (Rojas et al. 2019). As for the illegal
Study Area Collection Image Source Error clearing, in addition to the environmental im-
Year (RMS) pact, high risk situations are generated, caused
by ground instability. This progressive growth
Aconcagua 2004 – 2019 Google Earth Pro – 0.45 – 0.32
of Latin American cities has altered and frag-
ArcGIS Pro mented wetlands, whether it be for housing or
Ciénaga 2002 – 2019 Google Earth Pro – 0.69 – 0.62 transportation infrastructure, creating coastal
de Mallorquín ArcGIS Pro zones with degraded and devalued ecosystems
Pantanos de Villa 2002 – 2019 Google Earth Pro 0.98 – 0.73 that are marginalized from the territory.
Because of this situation in Latin America,
the study of urban wetlands is highly relevant,
FIGURE 1: Location of the studied wetlands.

as it can contribute to our base of knowledge and support TABLE 2. Land use covers, definitions and reference signatures.
conservation initiatives. Urban wetlands have been little
studied and defined, to the point that the Ramsar Conven-
tion (Secretary of the Ramsar Convention) or global treaty
for the conservation of wetlands recognizes this debt,
admitting that urban wetlands have been forgotten (Het-
tiarachchi et al. 2015). Because of this, these authorities
have highlighted the relevance of urban wetlands, and in
2018, they declared that they were key sites for making
cities healthy and habitable. On the other hand, the lack of
information on the boundaries of urban wetlands has fur-
ther complicated their recognition in the planning of cities.
They are in fact a unique ecosystem. It is very challenging
to physically define them with remote sensing techniques
and geographic information systems, especially when they
coexist in a heterogeneous landscape and are located in
coastal cities (Adam et al. 2014; Gibril et al. 2020). Fur-
thermore, they do not have a uniform plant coverage, are
highly dynamic and their spectral reflectance is easily
disrupted. Additionally, land use has further increased their
variability, causing modifications in their vegetation and
water levels (Gallant 2015).
The objective of this study is to carry out a spatial
and multi-temporal analysis with remote sensors, to deter-
mine wetland surfaces and changes in three cities in Chile,
Colombia and Peru. It seeks to analyze changes in land use
that have led to loss and gain in urban wetlands during the
period (2002-2019) when the greatest loss of these ecosys-
tems has been reported (Darrah et al. 2019). Land changes
are analyzed with high-resolution satellite images, which
offset problems of definition by other sensors such as Land-
sat, as they have only been available for the last two de-
cades in the wetlands situated in the Latin American coastal
cities of Barranquilla, Lima, and Concón. These data will
allow for discovering the main spatial dynamics related to
the reduction and alteration of these ecosystems, thereby
taking a first step towards the recognition of the status of
these ecosystems. The chosen cities are part of the “Urban
Wetlands in Latin America: a solution for sustainable cities
SDG 11” Project (2019 -2021); cities that have experienced
rapid urbanization and are vulnerable to climate change.
OVERVIEW OF URBAN WETLANDS IN LATIN AMERICA

Although Latin American countries have joined the Ramsar
Convention (1981), and represent 11% of the world’s
wetlands, the region leads in wetland loss, reaching 59%
(Darrah et al. 2019). Specifically, the region has observed
losses of wetland surface in cities in Chile, Peru, Colom-
bia, Argentina and Brazil. Decreases are registered on the
Brazilian coast (Sousa et al. 2011; Wittmann et al. 2015),
the Andes and Caribbean regions of Colombia (Patino

areas. The pressure to build has led to the elimination of
and Estupinan-Suarez 2016) and the Luján River basin in
wetlands in cities spanning from north to south, especially
Argentina (Pintos y Sgroi 2012). However, figures of each
in the southern-central zone, which happens to have a high
Latin American country’s contribution and the distribution
recurrence of waterlogging and flooding (Rojas et al. 2014).
of most of the Latin American wetlands are still unknown,
The Aconcagua-Concón wetland is under pressure from in-
which is worrisome (Mitsch and Gosselink 2015).
dustrial activity and from urban growth. There is evidence
Coastal cities in Latin America are vulnerable to the
of an expansion in the real estate market for second homes
effects of climate change, which together with the socio-
(Hidalgo et al. 2016; Martínez et al. 2020), and a concen-
natural risks like flooding, means that more people are
tration of economic activities (ports, tourism and services).
exposed to danger. Chile, Colombia and Peru have been
In addition, since 2015 this coastal zone has been seriously
chosen as example countries with wetlands that are subject
affected by extreme events associated with climate change,
to urbanization and vulnerability to climate change. The
including swells, meteotsunamis, coastal erosion and tsuna-
three countries have experienced significant urban growth
mis like Japan 2011 and Illapel, Chile (Martínez et al. 2011;
and project further increases for 2030. Additionally, 80%
Martínez et al. 2018; Carvajal et al. 2017, Campos-Cava
of their population lives in cities, which are representative
2016); all of which have caused considerable damage to the
of the environmental conflicts caused by urbanization and
infrastructure and connectivity (Winckler et al. 2017). Ad-
conflicts with wetlands located in cities.
ditionally, the areas adjacent to the wetland (the Aconcagua
Chile River estuary) have been categorized as a low-quality land-
In Chile, more than 80% of the population lives in cities. scape because of the high degree of anthropic intervention,
There are more than 4 million ha of wetlands, but only 3% which has led to a further loss of naturalness on the coastal
is protected. None of these protected wetlands are in urban landscape (Rangel-Buitrago et al. 2018).
Colombia
TABLE 3. Land uses changes in wetlands In Colombia, wetlands make up more than 30 million ha,
Wetlands Total Total which is equivalent to 26% of its territory. Wetlands have
Losses Gains historically been associated with the development of human
Aconcagua 33 10 cultures (Jaramillo et al. 2015). The main cause of wetland
transformation has been the change in land use to pastures
La Ciénaga de Mallorquín 224 81
for raising cattle, agriculture and deforestation, and to a
Pantanos de Villa 11 2 lesser extent, urbanization (Patino and Estupinan-Suarez
Total 268 93 2016). These anthropic changes have led to the transforma-
tion of 24% of the country’s
FIGURE 2. Distribution of the land uses and cover types detected in the studied wetlands and surrounding wetlands. It is estimated that
areas for the Aconcagua study area (Concón, Chile) from 2004 to 2019. by 2025, the main factors of
change in wetlands will be, in
first place, the expansion of
ranching, and in second place,
urban and transportation devel-
opment (Ricaurte et al. 2018).
The Magdalena-Cauca basin
(i.e., where the Metropolitan
Area of Barranquilla is located)
is expected to suffer the great-
est changes in land use, with
negative effects on wetlands.
Barranquilla is a coastal city
located on the Magdalena River
estuary in the Caribbean Sea.
This estuary creates a vast area
of wetlands that have been
greatly altered by anthropic

activities throughout history and is highly vulnerable to the TABLE 4. Land cover changes in Aconcagua study area from 2004-2019,
effects of global change (Aldana-Domínguez et al. 2018; the analyzed image included the wetland and its surrounding areas. (Note:
Rodríguez 2015). The transformation of these ecosystems Any difference in net change totals is due to computer round-off.)
has had a negative impact on the ecosystem services, Aconcagua between
mainly on the regulation services (Aldana-Domínguez et 2004 and 2019
al. 2019), which means that new guidelines that allow for Land use Losses Gains Net Change
recovering and conserving the urban ecosystems must be a (ha) (ha) (ha)
priority.
Water bodies 0 9 8
Peru
In Peru, the expansion of cities is one of the main causes Roads 0 0 0
of wetland degradation (MINAM 2015), especially in the Dunes -5 0 -5
coastal and desert strip where wetlands have a special Wetlands -33 10 -23
value as nuclei of biodiversity and freshwater reserves. The Other vegetation -6 4 -2
wetland ecosystems cover a total surface of 6.9 million Plantation forest -9 1 -7
ha, which is equivalent to 5.4% of the territory (MINAM
2019). Among them, the coastal wetlands only occupy Beaches -10 5 -5
0.04%, and are especially vulnerable to growing real estate Bare soils -64 12 -52
pressure and the impact of urbanization on the aquifer. For Urban -1 15 14
example, the expansion of the city of Lima between 1990 Agriculture 0 3 3
and 2013 led to the loss of 203 ha of wetlands (Moschella Grasslands -29 66 37
2018). Studies on the main wetlands in Lima show the deg-
Industrial 0 31 31
radation and loss of ecosystem services caused by anthropic
pressure (Aponte and Cano 2013; Moschella 2012; Pulido
and Bermúdez 2018). Although there is a legal framework
for protecting wetlands, there are weaknesses in the instru- TABLE 5. Land cover changes in Ciénaga de Mallorquín study area from
2002-2019, the analyzed image included the wetland and its surrounding ar-
ments of protection and an adequate regulation for the
eas. (Note: Any difference in net change totals is due to computer round-off.)
application of norms is lacking (Ramírez Aponte 2018).
Further studies are also required to understand the hydrol- Ciénaga de Mallorquín between
ogy of these ecosystems and contribute to their conserva- 2002 and 2019
tion (Rodríguez 2017). In this sense, the National Wetland Land use Losses Gains Net Change
Strategy (MINAM 2015) has identified a lack of studies on (ha) (ha) (ha)
the valuing and management of wetlands as well as a weak Water bodies -14 128 114
participation for the conservation of these ecosystems.
Roads 0 7 6
METHODOLOGY Other vegetation -77 18 -59
Study Areas Beaches -20 36 15
The following urban coastal wetlands were chosen for this Bare soils -82 39 -42
study: 1) Aconcagua (also known as the Concón Wetland) Urban -1 70 70
in the city of Concón and associated with the Andean Acon-
Grasslands 0 17 17
cagua River in the Valparaíso Region (Chile), 2) la Ciénaga
de Mallorquín in Barranquilla (Colombia), and 3) Pantanos Permanent wetland -175 3 -172
de Villa in Lima (Peru). They are all located in coastal areas Semi-permanent
and are subject to the different pressures of changes in use wetland -49 78 29
from urbanization (Figure 1). Industrial -6 28 22
Data Processing
Free satellite images from the last two decades (2002-2019)
were selected from the collection available from the Quick-
bird satellite on Google Earth Pro. Then, a buffer of ap-
proximately 500 meters from the perimeter of each wetland
was defined. The images were georeferenced and classified
by photointerpretation at a scale of 1:2.000 in ArcGIS Pro.

FIGURE 3. Gains and losses in the Aconcagua Wetland (Concón, Chile) Preparation and Selection of Satellite Images for Land
from 2004 to 2019. Classification
Satellite images were selected from Google Earth Pro, ac-
cording to the following criteria:
• Time: Search of the collections that represent chang-
es over the last 20 years.
• Cloudiness: Determining factor for the selection of
an image in its next classification as cloudiness pres-
ents one of the main problems for detection.
• Seasonality: Preference for images with similar sea-
sonal conditions (winter – summer).
These criteria defined the studied time horizon from
2002 to 2019, and the 2.5-meter resolution Quickbird sen-
sor (See Table 1). A priori these images also strengthen
the detection of smaller surfaces like urban wetlands. The
QuickBird sensor images were georeferenced
by ArcGIS Pro by taking control points in
FIGURE 4. Distribution of the land uses and cover types detected in the studied wetland and
KML format on Google Earth Pro. A total of
surrounding area for the Ciénaga de Mallorquín study area (Barranquilla, Colombia) from
2002 to 2019. 18 control points was taken in each image for
each wetland, which were complemented with
points on the ArcGIS Pro basemap and field
work in March 2020. Once georeferencing
was completed, points with an error greater
than 1 meter were eliminated (Table 1).
Classification and Land
Use Covers
Classification was done through a combina-
tion of photointerpretation and image tracing
of the Quickbird images on ArcGIS Pro and
the calculation of the Normalized Difference
Vegetation Index (NDVI) that allowed for
observing the different types of vegetation and
separating the bodies of water and the uncov-
ered land. The recognition of the categories
was based on the definitions and reference im-
FIGURE 5. Gains and losses in the permanent wetland (a) and the semi-permanent ages shown in Table 2 that allowed for distin-
wetland (b) at the Ciénaga de Mallorquín (Barranquilla, Colombia) from 2002 to 2019. guishing natural areas from artificial areas and
The figures also show changes in wetland type (permanent v. semi-permanent). The using the time series of imagery to observe
conversion towards water bodies accounted for most of the permanent wetland loss, with trends from 2002/2004=2019. Surround-
change to semi-permanent wetland also a major factor in loss of this type. ing vegetation that is not part of the wetland
was not classified in detail. For the Ciénaga
de Mallorquín, two types of wetlands were
identified following the Colombian wetland
classification system (Ricaurte el al. 2019):
permanent wetland (permanently flooded),
and semi-permanent wetland (periodically
flooded; mainly covered by mangrove forests
in this area).

Validation of Land Covers the total loss of 33 ha were lost from 2004 to 2019, mainly
The validation of coverages was done through the capturing caused by the conversion to grasslands (17 ha), followed
of 840 points of control, whose verification was optimized by bare soils (8 ha), beaches (3 ha) and finishing with less
in GIS and with field work conducted in March of 2020. pressure of urban zones (2 ha) (Figure 3). The losses were
Then, the veracity of the interpreted classification was somewhat compensated for by a gain of 10 ha from bare
evaluated with the Kappa Statistic Index. soils (9 ha) and beaches (1 ha).
Besides the loss of wetlands, bare soils experienced a
RESULTS loss of 64 ha and gain of 12 ha for a net loss of 52 ha from
Land Covers 2004-2019. This loss accounted for some of the increase
The results of the land cover typing for the three wetland in grasslands surrounding the wetland. Some of the new
areas are shown in Figures 2, 4, and 6; they generated grasslands also came from beaches. A similar situation has
kappa index values of >85% of precision. Therefore, the occurred with the dunes – conversion of dunes to grassland;
maps allow for making a statistically valid interpretation. they experienced a net loss of 5 ha (Table 4).
The distribution and changes in land use are reported in
Tables 3-5, where values from the initial year, gains, losses TABLE 6. Land cover changes in Pantanos de Villa study area from 2002-
and net changes can be observed in the categories identified 2019, the analyzed image included the wetland and its surrounding areas.
for each wetland. (Note: Any difference in net change totals is due to computer round-off.)
A sum of 268 ha of decrease was observed totalizing Pantanos between 2002 and 2019
the three studied wetlands, of which 175 correspond to
loss (Losses – Gains). The main cause was an increase in Land use Losses Gains Net Change
artificial and productive covers such as urbanization and (ha) (ha) (ha)
grasslands, among others. Urbanization increased by a total Water bodies 0 9 9
of 162 ha considering the three studied cases. The land use Roads -1 5 4
covers changes observed over the last two decades them- Wetlands -11 2 -8
selves confirm the general trend of wetland loss in the Latin
Beaches -11 2 -9
American region reported by Darrah et al. (2019).
Of the studied wetlands, La Ciénaga de Mallorquín Bare soils -76 7 -70
(Colombia) has without a doubt suffered the greatest Urban -8 86 78
surface loss, followed by Aconcagua (Chile) and Pantanos Agriculture -1 0 -1
de Villa (Perú). Additionally, among the three wetlands, Green areas -3 0 -3
Pantanos de Villa is under the most
pressure from urbanization in its FIGURE 6. Distribution of the land uses and cover types detected in the studied wetland and sur-
surroundings (i.e., it is virtually surrounding area for the Pantanos de Villa wetland (Lima, Peru) from 2002 to 2019. The larger “Green
rounded by urban development with Area” is a private golf course.
the exception of the “Green Area”).
Aconcagua Wetland Area (Concón,
Chile)
In the case of the Aconcagua wet-
land study area, during the period of
analysis the main interactions and
exchanges were between the wetland,
grassland, bare soils, beaches and
urban and industrial areas (Figure 2).
First, the category experiencing the
greatest gain was grasslands, with 37
ha of net change, closely followed by
industrial area with 31 ha and then by
urbanization with a net gain of 14 ha
(Table 3).
The Aconcagua wetland itself ex-
perienced a net loss of 23 ha, but with

FIGURE 7. Gains and losses in the Pantanos de Villa Wetland (Lima, Perú) Another significant change in the area surrounding the
from 2002 to 2019. There were small changes in wetland area, mainly a wetland was the loss of other vegetation (59 ha; Table 5),
few hectares of losses. which included the last few fragments of tropical dry forest
in the area. The tropical dry forest is one of the most threat-
ened ecosystems and at risk of collapse in Colombia (Etter
et al. 2017), and it is also a key ecosystem for the supply of
ecosystem services in Barranquilla (Aldana-Domínguez et
al. 2019). It continues to lose surface area, mainly because
of urbanization. For the entire study area, as expected,
urbanization has increased (70 ha; Table 5), affecting both
wetlands and tropical dry forest.
Pantanos de Villa Study Area (Lima, Peru)
For the Pantanos de Villa study area, the greatest change
from 2002 to 2019 was the increase of urban areas (86 ha;
Table 6 and Figure 6), which mostly happened on bare
soils, and to a lesser extent on wetlands and beaches. Wet-
land coverage declined by 11 ha: 6 ha due to urbanization,
2 ha lost to bare soils, and the remainder to beaches and
roads. Wetland gained only 2 ha, with the most significant
La Ciénaga de Mallorquín Wetland Area (Barranquilla, corresponding to wetland expansion over an unbeaten path
Colombia) (Figure 7). Although the variation in the wetland’s extent
Both permanent and semi-permanent wetlands at La Cié- seems limited, it is worth mentioning that this wetland has
naga de Mallorquín changed during the study period (Table been converted to urban land for several decades before the
5 and Figures 4 and 5). The main wetland change was the period of this analysis. While most of the new urban areas
loss of permanent wetland surface (net loss of 172 ha). are registered here as conversions from bare soils without
This loss was mainly due to coastal erosion resulting from use, these bare soils are actually cleared wetland (i.e., they
the displacement of the sand bar that separates the wetland were mainly filled to dry the land for future plotting and
from the Caribbean Sea (Figure 5a). This contributed 94 ha building). Currently, the wetland is practically restricted to
to the overall net increase in the water bodies coverage. The the area that is protected and recognized as a Ramsar site.
water bodies are represented by the ocean and the river. Since its borders are mostly developed, it is most likely that
The wetlands have connections to both, but are separated its expanse will not vary.
by a sand bar and a spur. The retraction of the sand bar was Additionally, it is worth mentioning that there are
documented by Rivillas-Ospina and others (2018) who significant differences between the degree of urban plan-
noted an erosion rate of 0.14 m/year from 1973 to 2016. ning and its impact on the wetland’s functioning. To the
The permanent wetland was also reduced by an increase in northeast of the wetland, irregular low-income settlements
the semi-permanent wetland (52 ha), probably due to the and filled wetlands can be found. Here, recent land-use
natural formation of mangrove areas on the western side of changes affect the main springs that supply the wetland and
the Ciénaga de Mallorquín and a recent mangrove refores- the water channels that connect them, putting the quality
tation effort. The formation of beaches and conversion to and quantity of surface flow entering the wetland at risk.
urban development were also responsible for the loss of 22 Specifically, there is an increase of dwellers that use the
ha and 6.7 ha, respectively, of permanent wetland. spring water as they lack piped drinking water. Also, a near
On the other hand, the semi-permanent wetland gained and unstable landfilling might block the channels. For in-
52 ha at the expense of permanent wetland and 25 ha from stance, a public sports field obstructs the water flow. Mean-
bare soils, while it lost some surface area mainly to wa- while, the southwest end presents mostly beach urbaniza-
ter bodies (12 ha), bare soils (11 ha), grasslands (10 ha), tion through construction of high-income condominiums,
and urban zones (8 ha; Figure 5b). Although the Ciénaga which have drained the wetlands and have also reduced the
de Mallorquin is included in the Ramsar Site “Estuarine ecosystem’s connection with the coast.
system of the Magdalena River Ciénaga Grande de Santa
Marta”, anthropic pressures continue to affect it.

CONCLUSION ments, which were built on cleared wetlands and the area
The analysis of Quickbird images from the period of surrounding the protected zone (i.e., designated Ramsar
2002-2019 has allowed for the creation of valid maps that Site). The urbanization of Aconcagua wetland is lesser
show land use cover, including wetlands during two time than grassland growth and, in this period, impacted due to
periods. Consequently, it has permitted spatial and tempo- urbanization process was produced before eighty decades.
ral tracking of wetland trends in selected urban wetlands Overall, the urban wetlands have lost surface, confirm-
in Latin America and provided information on land cover ing the Latin American trend. Urban expansion, mainly for
changes over the past 20 years. housing, is the major impact but urbanization may also be
Our study shows important reduction in surface area of responsible for a change in beaches and grassland. Pres-
most of the studied urban wetlands, mainly caused by three sures surrounding the wetlands that could be interpreted as
factors: coastal erosion/creation of water bodies, expansion an urbanization transformation process were also identified
of grassland and growth of urban areas surrounding them. e.g., kind of settlements in Pantanos de Villa (Perú).
The Ciénaga de Mallorquín wetland has suffered the Latin America faces a great challenge in advancing
greatest loss, caused by coastal dynamics and changes towards urban sustainability according to the 17 Sustain-
in the water body and vegetation. Coastal erosion is a able Development Goals (SDG established by the 2030
complex phenomenon and, in this area, it is related with Agenda (UN, 2015) where the Goal 11 have a target to
oceanographic processes (i.e., currents and waves gener- make cities and human settlements inclusive, safe, resilient
ated by the wind and tides) and the anthropogenic im- and sustainable. The region also needs to improve urban
pacts originated by the construction of infrastructures to planning and management regarding the identification,
maintain the Barranquilla port and the navigable channel definition, and norms for permitted uses of urban wetlands
(Rivillas-Ospina et al. 2018). and their surroundings. Urban wetlands should be “rec-
The spreading of grasslands could be interpreted as ognized spaces” in the city – natural habitats important for
pre-construction activity, where grassland increase over the well-being of the city residents and kept safe, resilient
bare soils and agricultural areas mostly in the case of and sustainable facing the challenges of climate change. n
Aconcagua. The Aconcagua wetland area, although located
ACKNOWLEDGEMENTS
in an area under a great deal of historic pressure from in-
This study is part of the research project “Urban Wetlands
dustrial activity, is currently being altered by an increase of
in Latin America: a solution for sustainable cities SDG
grasslands and bare soils specially when the dunes started
11” (2019 -2020). It was financed by the CODS – Latin
to increase with low vegetation due to natural condi-
American Sustainable Development Goals Center, whose
tions as a precipitation, which is also affecting the losses
general objective is to strengthen interdisciplinary research
in beaches. The Aconcagua wetland is lesser affected by
and cooperation on urban wetlands, recognizing that they
urban growth and agriculture areas.
are key ecosystems fundamental for fostering more sus-
Clearly, urbanization is affecting these three areas.
tainable and resilient cities (i.e., by 2030, as mandated in
The Ciénaga de Mallorquin is the most urbanized wetland
the SDG 11 for Latin American countries), and therefore
studied, losing 15 ha due to urban expansion. Two sectors
strengthening their connection with public policy.
of urban growth are recognized: the eastern side of the
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WETLAND CONSERVATION/EDUCATION/OUTREACH
Wetland Conservation Concerns in Southern Mexico

Tatiana Lobato de-Magalhães, Lucia Guadalupe Cruz, and Everardo Barba1
ABSTRACT Table 1). The region’s wetlands are valuable heritage places
that provide several ecosystem services (Smardon 2006;
A pproximately 16 percent of Southern Mexico’s surface
area is comprised of wetlands which harbor an abun-
dance of plant and animal species, including endangered
Gortari-Ludlow et al. 2015), and substantially contribute
to maintaining biodiversity at local and landscape levels
and endemic species. With two-thirds of the total wetlands (Mora-Olivo et al. 2013; Alcocer and Aguilar-Sierra 2019).
of Mexico and one-third of Mexican Ramsar sites, the This region has some of the highest levels of aquatic plant
Southern Mexico region plays a critical role in wetland species richness and endemism worldwide (Murphy et al.
conservation worldwide. Despite national and international 2019).
efforts, many wetland species and ecosystems are threat- The National Wetland Policy highlights Tabasco State
ened in this region. This review includes information reas having a vast expanse of wetlands (floodplain zones),
lated to seven Southern Mexico states: Campeche, Chiapas, particularly the Pantanos de Centla Biosphere Reserve
Guerrero, Oaxaca, Quintana Roo, Tabasco, and Yucatán. (Figure 1a, Figure 2), with an area of 3,027 km2, covering
From coastal areas to highlands, this region has around 12% of the total state surface. Among these, the Grijalva
2,020 mapped wetlands (64,298 km2) and 41 Ramsar sites. River and the Usumacinta River form an estuarine region
Alarmingly, only 13 of the 41 Ramsar sites have manage- which is considered one of the most important deltas in
ment plans implemented. Regardless of the importance of North and Mesoamerica because of the water flow and the
inland wetlands in terms of their area and economic value, importance for migratory birds and other species (IUCN
issues regarding their conservation and restoration are 2020; SEMARNAT 2020).
generally lacking or neglected. Southern Mexican wetlands
WETLAND TYPES AND CLASSIFICATION
are also severely threatened by changes in natural habitats,
According to the National Wetland Inventory (CONAGUA
particularly those associated with excessive exploitation of
2020), Southern Mexican wetlands are grouped into three
natural resources, tourism, and the oil industry.
major classes: 1) marine and coastal wetlands, including
INTRODUCTION marine and estuarine systems, 2) inland wetlands, including
Southern Mexico is a megadiverse, neotropical region lacustrine, palustrine, and riverine systems, and 3) human-
that harbors several types of wetlands such as mangroves, made wetlands (Table 1; Figure 3). Wetlands are further
riparian forests, floodplains, and cenotes (sinkholes) classified based on their hydrological regime (permanent,
(Figure 1). According to the National Wetland Inventory, intermittent, and temporary wetlands), soil properties
wetlands cover six percent of Mexico (CONAGUA 2020). (texture and composition), and vegetation type, such as
In Southern Mexico, 2,020 wetlands occupy 64,298 km2, the endemic flooded low evergreen forest ecosystem in the
representing two-thirds of the total wetlands in Mexico and Yucatán peninsula (Bala’an K’aax) and the islands of vigor-
16% of the Southern Mexican States territory (the states ous tree vegetation associated with springs and water holes,
of Campeche, Chiapas, Guerrero, Oaxaca, Quintana Roo, which constitute a critical habitat for wildlife, Los Petenes
Tabasco, and Yucatán). in the Campeche State (Figure 1b) (Lot 2004; Ramsar
Mexico is the country with the second highest num- 2020a).
ber of Ramsar sites (142 sites designated as Wetlands of Marine and Coastal Wetlands
International Importance), behind the United Kingdom This wetland type represents around 15% of the total
(175 sites) (Mauerhofer et al. 2015). There are 41 Ramsar mapped wetlands. Marine wetlands (Figure 1c) are most
sites in Southern Mexico, which represents approximately represented by a seagrasses community or ceibadal (e.g.,
29% of the total Mexican Ramsar sites (Ramsar 2020a; Halodule, Syringodium, and Thalassia species) (Creed
et al. 2003) and mangroves (Figure 1d). Some threatened
1 El Colegio de La Frontera Sur (ECOSUR), Mexico; Corresponding author mangrove plant species are Avicennia germinans (black
contact: ebarba@ecosur.mx

mangrove), Conocarpus erectus (but- FIGURE 1. Southern Mexico wetlands: (a) palustrine wetland, El Palmar, Pantanos de Centla
ton mangrove), Laguncularia racemosa Biosphere Reserve, Tabasco State; (b) islands of tree vegetation associated with springs and
(white mangrove), and Rhizophora water holes, Los Petenes, Campeche State; (c) marine wetland with seagrasses vegetation,
mangle (red mangrove) (SEMARNAT Laguna de Términos, Campeche State; (d) mangrove, Tabasco State; (e) riverine rainforest, Rio
2010). Campeche and Quintana Roo Tzendales, Chiapas State; (f) lacustrine wetland, Laguna Catazaja, Chiapas, (g) sinkhole (cenote)
frequented by tourists, Quintana Roo State, and (h) wetland habitats associated with waterfalls,
States have a large portion of mangroves Parque Nacional Cañón del Sumidero, Chiapas State.
on the Atlantic Coast, and Chiapas State
contains large areas of mangrove on the
Pacific Coast.
Inland Wetlands
Most of the Southern Mexico wetlands
are classified as inland wetlands (82%).
They include mostly freshwater palus-
trine wetlands described as swamps,
floodplains, marshes, and forested
wetlands (riparian forests, palm thickets,
and inundated low lands) (Figure 1e).
Riparian forests are comprised of Salix
negra, S. caroliniana, and S. chile (wil-
lows). Lowlands floodable forests are
represented by Annona glabra (swamp
apple) (Campeche State), Dalbergia
brownei (rosewood), and Ficus padofo-
lia (fig tree) (Tabasco State) (Lot 2004).
Lacustrine wetlands (Figure 1f) occur
mostly in highlands and are less abun-
dant than palustrine wetlands (Olmsted
1993). Rooted floating-leaved plants are
numerous in lakes, lagoons, canals, and
open freshwater wetlands (e.g., Nympha-
ea – waterlily, Nuphar – waterlily, Nym-
phoides – floatingheart, Potamogeton –
pondweed, and Sagittaria – arrowhead)
(Lot 2004). Fifty-eight plant species are
associated with calcareous warm-water
rivers of Yucatán peninsula includ-
ing Bacopa monnieri (water hyssop),
Eleocharis geniculata (spikesedge),
Hydrocotyle umbellata (manyflower),
Lemna aequinoctialis (duckweed),
Nymphaea ampla (waterlily), Pas-
palum notatum (bahiagrass), and Typha
domingensis (cattail) (Tapia-Grimaldo
et al. 2017). Sinkholes (cenotes) are a
unique type of inland wetland associ-
ated with a karstic geology. The cenotes
are an important freshwater resource in (Photos a, b, c, e, f courtesy of Everardo Barba; d courtesy of Alejandro Betancourth; g
the Yucatán peninsula region that are and h courtesy of Paula Montoya)
highly impacted by tourism (Figure 1g);
they harbor several rare and threatened
aquatic species (Cervantes-Martínez

FIGURE 2. Panoramic view of Pantanos de Centla Biosphere Reserve, Tabasco State. et al. 2018; Mondragón-Mejía et al.
2019). Highland wetlands are extreme-
ly important for the provision of water
to Southern Mexico cities like San
Cristóbal de las Casas. The dominant
aquatic plants in highland wetlands are
Typha (cattail), Phragmites (common
reed or carrizal), Cyperus (umbrella
sedges), Eleocharis (spike-rushes),
and Schoenoplectus (bulrushes) (Lot
2004; Chediack et al. 2018).
Human-made Wetlands
Human actions have created wetlands
in places, especially through dam con-
struction. They represent about three
percent of Southern Mexico wetlands.
ECOSYSTEM FUNCTIONS AND THREATS

Southern Mexican wetlands provide
(Photo courtesy of the Mexico Mangrove Monitoring System developed by CONABIO/
several ecosystem services and are
SEMARNAT. Photo taken by Joanna Acosta) also impacted by human uses such
as livestock, aquaculture, excessive
exploitation of natural resources, and
industrial expansion (Tables 2 and
FIGURE 3. Wetlands in Southern Mexico States. (Adapted from National Wetland Inventory; CONA-
GUA 2020) 3). For example, Ría Lagartos (Yu-
catán State) is an important estuarine
wetland for flamingo nesting as well
for economic activities such as fish-
ing, agriculture, salt production, and
livestock. Another activity that has
a strong impact on wetlands is the
selective extraction of native palms
such as Pseudophoenix sargentii
(buccaneer palm or kuka’), Thrinax
radiata (thatch palm or chit), and Coc-
cothrinax readii (Mexican silver palm
or nacax), which are used for decora-
tion along avenues and hotels in cities
like Cancun. In Laguna de Terminos
(Figure 1b) (Campeche State), the
exploitation of natural resources has
been crucial for the local economy
during the last three centuries, through
the extraction of dye sticks, precious
woods, and chewing gum. This wet-
land is known for the sustainable use
and management of Crocodylus more-
(Map elaborate by Tatiana Lobato M.)
letti (Mexican crocodile) populations
for commercial purposes based on its
skin (SEMARNAT 2020).

Regarding economic valuation of wetland ecosystem BIODIVERSITY, ENDEMISM, AND THREATENED SPECIES
services, inland wetlands are rated higher than estuarine Southern Mexico wetlands harbor several endemic species
ones in Tabasco State: palustrine ($9,689 USD/ha/year), such as Lithobates brownorum (leopard frog), Bolitoglossa
lacustrine ($6,366), mangrove ($2,653), and coastal lagoon yucatana (Yucatan mushroomtongue salamander), Cyprin-
($1,926) (Camacho-Valdez et al. 2020). Overall, inland odon macularius (desert pupfish), Caretta caretta (logger-
wetlands are threatened by the land-use changes and by head sea turtle), Chelonia mydas (green sea turtle), Eret-
the oil industry, especially the swamps, floodplains, and mochelys imbricada (hawksbill sea turtle), and Sanopus
marshes in Tabasco State. The extraction of hydrocarbon splendidus (splendid toadfish). The following states harbor
has led to major wetland impacts (Domínguez-Domínguez a rich number of aquatic plant species and are considered
et al. 2019; Camacho-Valdez et al. 2020). Furthermore, priority states for the conservation of strictly aquatic plant
in the last two decades the Pantanos de Centla (Tabasco species in Mexico: Chiapas (225 species), Campeche (220),
State) has experienced a notable land-use change – the Oaxaca (210), and Tabasco (186) (Mora-Olivo et al. 2013).
conversion of natural floodplain vegetation to livestock The Pantanos de Centla (Tabasco State) alone harbors
and agricultural areas (De la Rosa-Velázquez et al. 2017). around 569 plant species (76 used by people and 13 rare or
Lowland floodable forests have been drastically reduced by threatened) and a fauna with more than 523 vertebrate spe-
agricultural activity (conversion to pasture and farmland) cies (IUCN 2020; SEMARNAT 2020). The Anillo de Ceno-
and overexploitation of the Haematoxylum campechianum tes are home to endemic species of reptiles (e.g., Terrapene
(campeachy tree or logwood), which was used for a long carolina yucatana – Yucatan box turtle), amphibians (e.g.,
time as a natural source of textiles dye, applied in histology Bolitoglossa yucatana), and birds (e.g., Stelgidopteryx
for staining, and for medicinal uses (Lot 2004). ridgwayi – Yucatan rough-winged swallow, Cyanocorax
Frequent threats to the highland wetlands are urbaniza- yucatanicus – Yucatan jay, and Melanoptila glabirostris –
tion, pollution, mining, and agricultural activities. Several black catbird). These cenotes are also home to a number of
monocotyledons aquatic species historically recorded above endangered or threatened species (Cervantes-Martínez et al.
2,000 m a.s.l. (Chiapas State) were not detected in a recent 2018; IUCN 2020; Ramsar 2020a). The highland wetland
floristic study; the authors consider that it could indicate a Humedales de Montaña La Kisst (Chiapas, 2,120 m a.s.l.)
process of local extinction (Chediack et al. 2018). supports great populations of fish and amphibians, with at
TABLE 1. Extent of wetlands in Southern Mexico based on National Wetland Inventory (NWI) and Ramsar sites.
Wetland Number of Wetlands Total Area (km2) Reference
Total NWI 2,020 64,298 CONAGUA 2020

Marine and Coastal 392 9,602
Inland 1,558 52,610
Artificial 70 2,085
Ramsar sites 41 34,232 Ramsar 2020a
Marine and Coastal 26 29,451
Inland 15 4,782
TABLE 2. Ecosystem services provided by wetlands in Southern Mexico. (Adapted from Smardon 2006; Camacho-Valdez et al. 2020; Ramsar 2020a.)
Ecosystem Services Provided by Wetlands
• Recreation and tourism
• Scientific and educational uses
• Heritage places
• Drinkable water storage
• Hydrological flow regulation
• Biological production (wetland food and non-food products)
• Biogeochemical cycle regulation (erosion protection, pollution control and detoxification, nutrient cycling, and soil
formation)
• Wildlife habitat and biodiversity conservation (genetics, endemism, and rare and threatened species)

FIGURE 4. Ramsar sites in Southern Mexico States. (Adapted from Ramsar 2020a) least 10 species being endemic or under a
protection category (e.g., the endemic fish
Profundulus hildebrandi - Chiapas killi-
fish, and the endemic plant Wolffia colum-
biana – Columbian water-meal) (Chediack
et al. 2018; Ramsar 2020a).
Several wetland species are criti-
cally endangered due to habitat loss. The
International Union for Conservation of
Nature’s Red List of Threatened Species
lists one species of Fungi, 46 plants, and
375 animal species associated with South-
ern Mexican coastal and inland wetlands
(IUCN 2020). Concerning the threatened
categories of IUCN, seven species are
critically endangered, 17 species endan-
gered, 27 species vulnerable, 15 species
near threatened, 276 species least concern,
and 80 species data deficient (Table 4). In
regard to aquatic animal species, there are
four species of mollusks, 125 species of
(Map elaborate by Tatiana Lobato M.) arthropods (Insecta and Malacostraca), 145
TABLE 3. Activities that threaten Southern Mexico wetlands. (Adapted from Gortari-Ludlow et al. 2015; Domínguez-Domínguez et al. 2019; Camacho-
Valdez et al. 2020; Ramsar 2020a.)
Wetland Threats
• Change in natural habitats (agriculture, livestock, aquaculture, and human settlements)
• Excessive exploitation of natural resources (fishing and harvesting aquatic resources, logging and wood harvest-
ing, hunting and collecting terrestrial animals, marine and freshwater aquaculture, and gathering plants)
• Changes in flow regime (drainage and canals construction)
• Wastewater (rural, urban, and industrial)
• Drought (high temperature and high evaporation)
• Infrastructure projects (road construction)
• Oil industry (hydrocarbon extraction and processing)
• Unsustainable use (tourism and navigation)
TABLE 4. Threatened species in Southern Mexico wetlands. (Adapted from IUCN 2020.)
Taxonomic Critically Near Data
Endangered Vulnerable Least Concern Total
Group Endangered Threatened Deficient
Fungi 1 - - - - - 1
Plants - 2 - 1 43 - 46
Mollusks - - - - 3 1 4
Arthropoda 1 7 8 1 72 36 125
Fishes 2 7 12 6 76 42 145
Amphibians 2 - - - 3 - 5
Reptiles 1 - 5 1 22 1 30
Aves - - 1 6 57 - 64
Mammals - 1 1 - - - 2
Total 7 17 27 15 276 80 422

fishes, five amphibians, 30 FIGURE 5. Number of Ramsar sites that satisfy each Ramsar criterion for Southern Mexico. Wetland type
reptiles, 64 species of wa- (criterion 1), biological diversity (criteria 2, 3, and 4), waterbirds (criteria 5 and 6), fishes (criteria 7 and 8), and
terfowl birds, two mammals other taxa (criterion 9). (Adapted from Ramsar 2020a, b)
Rheomys mexicanus (Mexican
water mouse) and Trichechus
manatus (American manatee).
Among the arthropods there
are 152 species of insects that
depend on the aquatic systems
for critical stages in their life
cycles, including dragonflies
and damselflies. Coastal
ecosystems and mangroves are
crucial for the threatened croc-
odiles and caimans (i.e., Cro-
codylus moreletti - Mexican
crocodile and Caiman croco-
dilus - spectacled caiman).
Southern Mexico also shelters
a vast number of waterfowl,
and migratory aquatic birds
that come to its wetlands in
the winter, and several of these
species are threatened (Platt et
al. 2010; Domínguez-Domín-
guez et al. 2019; IUCN 2020).
(Map elaborate by Tatiana Lobato M.)
The aquatic turtle Dermatemys
mawii (white turtle) is the only
critically endangered reptile in the region. Other threatened The first Mexican Ramsar site was designated in
freshwater aquatic turtles are Kinosternon creaseri (creaser’s 1986 and is located in Southern Mexico at Yucatán State,
mud turtle), K. integrum (Mexican mud turtle), K. oaxacae Reserva de la Biosfera Ría Lagartos. The most recent,
(Oaxaca mud turtle), Trachemys ornata (ornate slider), and Humedales de Montaña María Eugenia, was added in 2012
T. scripta (pond slider). and is located in Chiapas state (Ramsar, 2020a). A total
of six sites were also designated as UNESCO Biosphere
RAMSAR SITES Reserves. With regard to geographical distribution, all
Southern Mexico has 41 Ramsar sites covering 34,232 seven Southern Mexican states have at least one Ramsar
km2 (2% of the total country surface). It represents almost site (Ramsar 2020a) (Figure 4). Quintana Roo state has
a third of the 142 total Mexican Ramsar sites. Around the largest number of sites (13), followed by Chiapas (12),
85% (35 wetlands) of Southern Mexico sites occur in low Yucatán (7), Oaxaca (4), and Campeche (3), while Guerrero
elevations (< 300 m a.s.l.), while the highest elevation and Tabasco have only one Ramsar site each. Additionally,
Ramsar sites are attributed to Humedales de Montaña La two Southern Mexico Ramsar sites wetlands extend into
Kisst and Humedales de Montaña María Eugenia, both in the territory of other countries. Parque Nacional Lagunas
Chiapas State (2,120 m a.s.l.). The majority of Ramsar sites de Montebello lies on the Southern border, extending into
are classified as coastal and marine (64%), followed by Guatemala and Parque Nacional Arrecifes de Xcalak into
inland wetlands (36%) (Table 1). Among inland wetlands, Belize border.
13 sites have a permanent water regime and two sites are The Ramsar Convention considers nine criteria to
considered seasonal or intermittent. Mexican Ramsar sites designate wetlands as of international interest for conserva-
have a mean size of 835 km2. The largest site (7,050 km2) is tion (Ramsar 2020a): wetland type (criterion 1), biological
Área de Protección de Flora y Fauna Laguna de Términos diversity (criteria 2, 3, and 4), waterbirds (criteria 5 and
(Campeche State), while the smallest site (0.2 km2) is Playa 6), fishes (criteria 7 and 8), and other taxa (criterion 9)
Barra de la Cruz (Oaxaca State) (Ramsar 2020a). (Ramsar 2020b). Criteria based on wetland type and bio-

diversity are more frequently reported on than ones related 90% of the aquatic plant species richness of Mexico (Mora-
to specific taxa (Figure 5). Among the 41 Southern Mexico Olivo et al. 2013) and having higher economic values than
Ramsar sites none satisfy all criteria and one fills only one estuarine wetlands (Camacho-Valdez et al. 2020), Mexi-
criterion. The latter site is the Parque Nacional Cañón can conservation actions focused on inland wetlands are
del Sumidero (criterion 1) that contains a unique example lacking, particularly for highland wetlands (Alcocer and
of a natural wetland type (humid habitats associated with Aguilar-Sierra 2019).
waterways and waterfalls) (Figure 1h) and harbors threat- Strong synergies among stakeholders that engage the
ened species such as Crax rubra (great curassow), Ateles population, private and governmental sectors, decision-
geoffroyi (black-handed spider monkey), Crocodylus acutus makers, non-governmental organizations, and the acad-
(American crocodile), Leopardus wiedii (margay), and emy are crucial to improving wetland conservation in
Rinodina chrysomelaena (bright yellow crustose lichen) Mexico. Projects that integrate science and practice are
(IUCN 2020). also essential for wetland restoration and conservation
at local and regional levels. Through improved efforts to
CONSERVATION PERSPECTIVE AND CONCLUSIONS increase wetland protection awareness and acquisition
Several actions have been launched since 1986 when Mex- of more detailed data on the degradation and change of
ico became a signatory country of the Ramsar Convention, wetland areas, including risk assessment analysis (Cama-
particularly the creation of the National Wetland Policy and cho-Valdez et al. 2020), Southern Mexican wetlands could
a National Wetlands Committee, the designation of 142 achieve a positive future scenario. n
Ramsar sites, and development of the National Wetland
Inventory (CONAGUA 2020; SEMARNAT 2020). Addi- ACKNOWLEDGMENTS
tionally, since 1936, Mexico has been party to an agreement We thank K. Murphy, D. Jones, and D. Orsini for reviewing
with the United States for the protection of migratory birds, this manuscript. The first author holds a postdoctoral fel-
which has contributed to the implementation of bi-national lowship from ECOSUR (El Colegio de la Frontera Sur) at
initiatives that have improved wetland conservation in CONACyT (National Council of Science and Technology)
Mexico. Despite the international importance of the rein Mexico. We appreciate the valuable comments of the
gion’s Ramsar wetlands, only 71% of the Southern Mexico editor R. Tiner.
Ramsar sites have management plans (25 sites with con-
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Doi:10.1016/j.aquabot.2019.06.006

WETLAND CONSERVATION/EDUCATION/OUTREACH
Propagation of Endangered Aquatic Plants: An Experience that Promotes ex situ Con-

servation and Environmental Education
Sandra Nayeli González Mateos1
ABSTRACT ic times, aquatic plants have been used for various purposes
etlands are among Mexico’s most threatened habitats (Miranda 1980). One of the most important uses was to
W as they have not received the degree of protection and
conservation as they have in North America. To promote
build the “chinampas” - the agricultural production systems
(e.g., “floating gardens”) that helped to shape the city itself
their conservation, the Botanical Garden at the Institute of and served to “gain” space in the water and form new sec-
Biology at the National Autonomous University of Mexico tions of “land” for the very expansion of the great city (for
(UNAM), located south of Mexico City, has established a an example, see https://aztecexplorers.com/2018/07/26/
collection of aquatic plants with the intent to have a repre- travelling-in-time-exploring-the-chinampas-of-tlahuac-
sentative sample of the country’s aquatic plants, with em- mexico-city/). Species such as Lilaeopsis schaffneriana,
phasis on the “Cuenca of Mexico”. The collection is being Schoenoplectus tabernaemontani, Hydrocotyle ranunculoi-
used to propagate aquatic species, conduct research, promote des, Polygonum amphibium, Lemna gibba and Bidens au-
conservation of aquatic plants, and to serve as the founda- rea were used specifically for the construction of “chinam-
tion for environmental education programs to increase public pas” (Lot and Novelo 2004).
awareness of these species and the challenges they face. Despite their historic significance, there are currently
very few studies for these types of aquatic plants in both
WHY CONSERVE AQUATIC PLANTS IN THE BOTANICAL natural populations and ex situ conditions. The Aquatic
GARDEN OF THE INSTITUTE OF BIOLOGY – UNAM? Plant Collection (APC) is a botanical collection of the few
In Mexico, aquatic ecosystems are among the country’s of its kind that exist in Mexico and its goal is to have a rep-
most threatened natural habitats. They have received little resentative sample of the aquatic vascular plants of the flora
attention for their conservation and there are few studies re- of our country, with emphasis on the “Cuenca of Mexico.”
lated to Mexican aquatic species (Mora-Olivo et al. 2013).
Aquatic plants have been linked to man since very ancient WHO ARE WE?
times - the element water has been a very important factor The Botanical Garden of the Institute of Biology - UNAM
for the establishment and development of great civilizations (JB IB-UNAM) is located south of Mexico City and oc-
including the great Tenochtitlan in the “Cuenca de México” cupies an area of 2.75 hectares (Caballero Nieto 2012). The
(the ancient capital of the Aztec empire). Since pre-Hispan- Aquatic Plant Collection (APC) is distributed in 17 ponds lo-
FIGURE 1. Panoramic view of one of the ponds in the Aquatic Plant Collection containing plants with medicinal and/or food use (e.g., Equisetum
hyemale and Nymphaea mexicana). (Photo by Surya Ivonne González Jaramillo.)
1 Botanical Garden of the Institute of Biology –National Autonomous University

of Mexico (UNAM), Mexico City, Mexico; author contact: nayelig@ib.unam.mx

cated in different sections of the botanical garden and covers FIGURE 2. Aquatic plant propagation tank in the greenhouse. (Photo by
an area of approximately 473 m2 (Figure 1). It also contains a Surya Ivonne González Jaramillo.)
greenhouse of 95 m2 for the propagation, growth and devel-
opment of different species of aquatic plants (Figure 2).
Originally, the founders and collaborators of the JB
IB-UNAM made various collections for the production of
species that show the plant diversity of Mexico. Historical
records indicate that in some of the botanical expeditions,
aquatic plants were collected (Equisetum hyemale var. affine,
Equisetum giganteum, Hydrocotyle sp., Sagittaria sagittifolia
and Typha sp. among others) however at that time no spe-
cific collection was consolidated strictly for aquatic plants.
From 2006, on the initiative of Dr. Javier Caballero Nieto
(then Head of the Botanical Garden) all the Garden’s collec-
tions were inventoried, strengthened and redefined into two
different types of collections according to their composition
and function: taxonomic collections and thematic collections.
It was under this concept, that the Aquatic Plant Collec-
tion (APC) was formalized as a thematic collection. At the FIGURE 3. Students working on various projects: Surya González (a)
invitation of Dr. Javier Caballero, Dr. Antonio Lot Helgueras working with Nymphaea odorata and university students - Sara Díaz,
(specialist researcher in aquatic plants of the Institute of Isaac Avalos, and Francisco Mendoz (b) working in the greenhouse. (Pho-
tos by Diana Ferrusca Domínguez – a, and Nayeli González – b.)
Biology UNAM) and landscape architect Pedro Camarena
developed a project for the establishment of this collection.
For this project, a theme was defined for each of the existing
ponds that included a plant palette of aquatic species and an
educational proposal for the use of the collection; this was
accomplished by biologist Teodolinda Balcázar Sol (Coordi-
nator of the Educational Area). From then on biologist José
Luis López was in charge of the maintenance of the ponds
and the asexual propagation counting for it, with the support
of the gardener Jesús Rebollo. Some of the collection sites
of the plant species were the states of Michoacán, Morelos,
Hidalgo and Jalisco.
NEW FLIGHT PLAN: ACTIONS TO BOOST APC

Starting in 2015, I was appointed curator responsible for a
the APC to continue developing the master plan designed
by Dr. Antonio Lot Helgueras. Each pond shows the differ-
ent life forms, endemism, introduced species (naturalized
species are shown, i.e. those that have dispersed to new en-
vironments and have succeeded in these sites and also those
species considered weeds, due to the excessive growth of
their populations that harms other species), as well as the
species used as medicine, food, fibers for the elaboration of
handicrafts and building materials. Species used for orna-
mental (various types and colors of flowers) and ceremonial
purposes are also included.
After analyzing the conditions of the APC and the green-
house, it was determined that the main actions to be taken
would be: 1) boost the sexual spread of species, 2) conduct dif-
ferent studies to learn more about the development of species b

under ex situ conditions, and 3) promote environmental educa- to their ex situ conservation as studies are carried out to
tion activities for the revaluation of Mexican aquatic plants. understand and improve their propagation.
These actions have been made possible through multidisci- N. gracilis or “atzazamolli” is a water lily with white
plinary projects that integrate high school students and college flowers that stand out from the water level and possesses
undergraduates in research topics related to aquatic plants and large green leaves (Figure 4a). Its cultural value is very
even fauna associated with the ponds in the collection. important as it is represented in the “Florentino Codex” in
Currently the APC includes 35 species from 19 botani- the section that mentions edible wild plants. This species is
cal families with the ultimate goal to increase the number endemic to Mexico and, unfortunately, is highly vulnerable
of species and individuals per species. Since 2016, 34 to the loss and contamination of water bodies.
students of high school or university levels have been Nymphaea mexicana called “atlacuetzon” or
performing volunteer activities, summer stays, profes- “paskurinda” inhabited the Mexico Basin and unfortunately
sional internships, social service, or thesis research. Basic its natural populations have declined drastically. This species
horticulture activities are carried out in addition to the with bright yellow flowers (Figure 4b) was a very impor-
design and execution of research that promotes the study tant element in the consolidation of the “chinampas.” It is
of aquatic plant species. This work expands our knowledge even represented in pre-Hispanic murals that demonstrate
about the biology of species, while encouraging students to its cultural value. In addition, it had an ornamental use for
develop technical skills for hydrophytic plant management the Day of the Dead offerings and its leaves were used in the
and the revaluation of wetlands (Figure 3). preparation of a special food that was made with the mixture
of other elements of the “chinampas” (López Martínez 2018)
SEXUAL REPRODUCTION RESEARCH N. odorata or “apapatla” is a very elegant and showy
Studies are carried out with some of the most attractive species whose white flowers float on the surface of the water
species of the Family Nymphaceae that are subject to more and offer a very pleasant sweet aroma (Figure 4c). This species
environmental pressures. The APC includes five species was a very important element for the formation of “chinam-
from this family, of which Nymphaea gracilis, N. odorata pas” however it has not been seen in the Basin of Mexico
and N. mexicana (Figure 4) are those for which germination since the 1940s (Note: It is the most common water lily in
and growth tests have been initiated. These three species North America). Its leaves are large, which favors the coloni-
were abundant in the “Cuenca of Mexico” and have been zation by aquatic fauna, both below water (i.e., for the deposit
elements of the landscape with a great cultural significance of snail eggs) and above their leaves (e.g., some small birds
(López Martínez 2018). Since their populations have de- perch on them in order to feed on various insects).
clined or disappeared completely they are now considered Since the summer of 2016, individuals of Nymphaea
as threatened species of extinction, according to current odorata, N. gracilis and N. mexicana have been monitored
environmental regulations. The JB IB-UNAM contributes in the different exhibition ponds and/or in the propagation
FIGURE 4. Three water lilies being studied: Nymphaea gracilis (a), N. mexicana (b), and N. odorata (c). (Photos by APC – a, and Nayeli González – b and c.)
a b c

TABLE 1. Summary of studies on propagation of aquatic plants.
N. mexicana N. odorata N. gracilis
Vegetative Successful Successful Successful
propagation
Fruits Not enough have been Bee pollination and cross pollination. Successful cross pollination.
obtained. Abundant fruits in older specimens.
Seeds Scarce Abundant Abundant

Germination Viable seeds have not been Low germination. Successful germination.
obtained. Stimulation with constant Stimulation with constant
temperature and dark conditions. temperature conditions.
Next actions Fertilization tests for flower Pre-germination treatments in seeds. Germination in different light and
and fruit production. temperature conditions.
FIGURE 5. From left to right: the first two images (with the yellow margin) shows the formation of stoloniferous rhizomes in N. mexicana for vegetative
propagation; the central pair of images (with the orange margin) shows the horizontal rhizome of N. odorata that can be cut for production of new individu-
als and N. odorata observed in fruit that is collected to obtain seeds; the images on the right (with the blue margin) show N. gracilis - its rhizome with
small leaves initiating growth and a student performing artificial pollination. (Photos by APC.)
FIGURE 6. Diana Ferrusca separating rhizomes from N. mexicana. (Photo by Surya Ivonne FIGURE 7. One of the new species added to the APC col-
González Jaramillo.) lection: Anemopsis californica. (Photo by Nayeli González.)

greenhouse. Sites were periodically reviewed to monitor lieve that ponds are a shelter option for them, but they are not.
the formation of flowers and fruits. Fruits were collected The ponds do not have a warm and constant temperature for
and seeds obtained. Species leaflets or rhizomes were also them and in many of them there is no food suitable for them.
collected to determine favorable characteristics for their Also many of the turtles arrive sick and fight each other as
spread. In addition, the greenhouse pollination of Nym- there are many (more than 25 individuals) for a small pond (60
phaea odorata and N. gracilis produced fruit, viable seeds, m2 approx.). Their presence poses a threat to the development
and seedlings of both species. Although some seeds of N. of water lilies and the proper functioning of the system. The
mexicana were produced, no seedlings have been estab- turtles consume leaves and buttons preferably of water lilies
lished. However N. mexicana did achieve favorable vegeta- and climb the leaves and damage them. Since 2017, monitor-
tive reproduction by means of rhizomes. There are different ing of ponds has been carried out for the detection of exotic
factors (e.g., substrate, light, and water level) that influence aquatic organisms. In 2019, a study was conducted to assess
both germination and the establishment of seedlings so ob- the effect of the overpopulation of the Japanese turtle on one
taining good-sized individuals from seeds involves a slower of the APC ponds. The turtle population was assessed, individ-
process but with greater advantages promoting genetic di- ual health was determined, then the turtles were removed from
versity (Bornette and Puijalon 2011). Table 1 highlights the the pond, treated, and when they were in good condition they
main results and the next actions with each of the species, were put up for adoption so that they could be removed from
while Figure 5 shows the propagation process. the pond to evaluate the response of the plants by removing
an element from the system. Four workshops have been held
PROPAGATING SPECIES UNDER EX SITU CONDITIONS
to inform visitors about the problem of abandonment of these
When plants are grown in greenhouses and gardens, situ-
turtles in the ponds.
ations arise that must be dealt with and often quickly to
Incorporation of new species into exhibition ponds.
maintain a healthy and productive environment. Plant
Through links with other botanical gardens of the Mexican
cultivation at APC requires an investment to time to ensure
Association of Botanical Gardens A.C. specimens or seeds of
the well-being of aquatic species in the collection.
aquatic plants have been received in donation for the strength-
Phenological monitoring of species and their horticultural
ening of APC’s collection. The main donors have been the
management. Fortnightly tours are carried out to identify the
Botanical Garden of Fundación Xochitla, A.C. in the State of
stages of development of the different species in the collection.
Mexico and the Botanical Garden of Culiacán, Sinaloa. Also
The formation of flowers, fruits and/or seeds is mainly identi-
some academics of the JB IB-UNAM, like biologist Ivonne
fied and recorded. These observations also help determine
Olalde, have collected aquatic plants that are currently propa-
whether the specimens require pruning or transplantation.
gated in the greenhouse for later incorporation into the ponds.
Comprehensive pest and disease management. Fort-
In the last three years seven new species have been added to
nightly observations also detect the presence of an insect or
the collection: Anemopsis californica (Figure 7), Ludwigia sp.,
any other organism whose presence may cause some damage
Thalia geniculata, Hydrocotyle ranunculoides, Limnobium
to aquatic species. Upon detection, Dr. Bonifacio Don Juan
laevigata, Marsilea mollis, and Lilaeopsis schaffneriana.
(responsible for the phytosanitary management of the collec-
Environmental education activities: the revaluation of
tions) is notified and the recommended treatment is applied.
Mexican aquatic plants. The general purpose of educational
Some problems caused by aphids, molluscs, and fungi have
activities is to promote the public’s interest in aquatic plants
been detected. Their treatment has been timely and health
and to recognize the importance of aquatic environments and
problems have been kept under control (Figure 6).
their plant diversity. For four years (2015-2018) we participat-
Wildlife program: native versus exotic species. The
ed in environmental education events: National Day of the Bo-
ponds provide habitat for wildlife. In some of the ponds
tanical Gardens and summer courses with the development of
and according to the season, you can observe various birds
different activities related to the APC. More than 300 people
such as the Mexican duck (Anas platyrhynchos ssp. diazi),
have participated in the programs. Some activities were guided
blue heron (Nycticorax nycticorax), frogs (e.g., Lithobathes
tours, children’s workshops or didactic games (Figure 8). The
montezumae), and dragonflies (e.g., Rhionaeschna multi-
activities were based on the “Environmental Education Action
color). Other organisms flock to the ponds to get their prey
Plan for the Botanical Gardens of Mexico” which promotes
(e.g., some insects become part of the diet of the collar
meaningful learning by involving different audiences and
lizard - Sceloporus torquatus) or to drink water and cool
diverse strategies considering that it is not only important to
down like “tlacuache” (Didelphis virginiana).
know the biology of plants but to have a comprehensive vision
Unfortunately, however, there are also exotic wildlife such
of plant resources (Martínez-González et al. 2012). We have
as the Japanese turtle (Traechemys scripta elegans) abandoned
observed after the implementation of the activities that 90% of
by visitors. Turtles are abandoned because their owners be-
the participants were unaware that there was so much diver- FIGURE 8. Examples of environmental education activities to revalue
sity of aquatic plants as they only knew the common water aquatic resources: guided tours (a) and children’s workshops (b). (Photos by
hyacinth (Eichornnia crassipes). They were most surprised to APC – a, and Surya Ivonne González Jaramillo – b.)
learn that some aquatic plants are so small (Lemna and Wolffia)
while others are insectivorous (Utricularia). Participants also
learned about the uses of aquatic plants since pre-Hispanic
times and their relationship with aquatic fauna. Some sugges-
tions for future activities include: 1) perform new activities
considering different educational levels and age ranges, 2)
implement workshops involving aquatic plant-animal relation-
ships and 3) develop horticultural workshops for propagation
of particular species such as horsetail (Equisetum hyemale).
CHALLENGES: THE IMMEDIATE FUTURE

APC will continue to develop the three basic lines of action:
1) sexual propagation of species, particularly for species of
other botanical families or others with a different conserva-
tion status, 2) optimal development of species under ex situ
conditions (e.g., assessing their growth and reducing the
incidence of health problems), and 3) conducting environ-
mental education activities and planning new strategies for
educational intervention in locations (field sites) that still
have natural populations of native aquatic vegetation.
The Aquatic Plant Collection provides an experience
that allows not only the study of Mexican aquatic plants,
but is also like a blue and green island in an urban environ-
ment that allows the development of an artificial wetland
with the presence of wildlife. In addition, it contributes to
the development of new professionals (Figure 9) and offers
Botanical Garden visitors an introduction to aquatic plants -
learning more about them (e.g. biology, ecology or ethno-
botany) and the need for their conservation. n
REFERENCES FIGURE 9. Students in the greenhouse (Surya González, Diana Ferrusca,

Caballero Nieto, J. 2012. El Jardìn Botánico del Instituto de Biología y
Nayeli González, Luis Silva, and Rogelio Yañez). (Photo by Surya Ivonne
la Estrategia Global para la Conservación Vegetal. In: J. Caballero Nieto
(ed.). Jardines Botánicos: contribución a la conservación vegetal de González Jaramillo.)
México. Ciudad de México, México: Comisión Nacional para el Cono-
cimiento y Uso de la Biodiversidad. pp. 75-86.
Bornette, G. and S. Puijalon. 2011. Response of aquatic plants to abiotic
factors: a review. Aquat. Sci. 73: 1-14.
López Martínez, C. 2018. Atlacuetzonan, planta acuática en el paisaje de
la Cuenca de México a través de sus usos. Ciudad de México: Facultad
de Arquitectura, UNAM.
Lot, A. and A. Novelo. 2004. Iconografía y estudio de plantas acuáti-
cas de la Ciudad de México. Ciudad de México. Universidad Nacional
Autónoma de México.
Martínez-González, L., and V. Franco, ad T. Balcázar. 2012. Plan de
Educación Ambiental para los Jardines Botánicos de México. Ciudad de
México. Asociación Mexicana de Jardines Botánicos.
Miranda Arce, M.G. 1980. Plantas Acuáticas útiles del Valle de México.
México, D.F.: Facultad de Ciencias, UNAM.
Mora-Olivo, A., J.L. Villaseñor, and M. Martínez. 2013. Las plantas vas-
culares acuáticas estrictas y su conservación en México. Acta Botánica
Mexicana 103: 27-63.

NOTES - WETLAND RESEARCH
Bark Traits: A Predictor for Recognition of Successional Groups in Riparian Forest

Species
Jane Rodrigues da Silva1,2, Augusto Cesar de Aquino Ribas3, Diogo da Silva Matos4, Edna Scremin-Dias2, and Rosani do
Carmo de Oliveira Arruda2
F rom the anatomical point of view, “bark” is the term used

to designate all tissues exterior to the vascular cambium,
therefore including peridermis, secondary phloem, and if they
Ocotea diospyrifolia a thick rhytidome was observed. In pio-
neer species Triplaris gardneriana (Figure 1a) and Inga vera
and, in late successional species Handroanthus heptaphyllus
remain, the primary phloem and the cortex (Angyalossy et and Vitex cymosa the secondary phloem is formed by tangen-
al. 2016). Bark traits, such as thickness and density, provide tial layers of axial parenchyma and conducting cells alternat-
a remarkable data source that can be exploited to understand ing with tangential layers of sclerenchyma from vascular
the plant’s life strategies in a community. Here, we evaluated cambium towards periderm. The sclerenchyma is composed of
whether the bark functional traits and anatomical features large sclereids in Vitex cymosa, and fibers with thickened walls
could be used as indicators of successional groups in woody in other species. In late successional Ocotea diospyrifolia
species from the riparian forest of the Paraguay River, Mato small aggregates of sclereids and large, oval sclereids clusters
Grosso do Sul state, Brazil. We hypothesized that pioneer spe- with thickened wall disperse were observed in the entire bark
cies have thicker bark, with lower density and low percentage (Figure 1f). In Vochysia divergens, large sclereids clusters are
of sclerenchyma tissues than the late successional species. restricted to the collapsed secondary phloem (Figure 1e). We
We collected bark samples from the main stem at 1.30 m detected starch stored in the parenchyma cells in all examined
above ground from three specimens in each group: pioneer species and phenolic compounds in the inner bark of pioneer
species (Inga vera, Fabaceae; Triplaris gardneriana, Polygo- species Inga vera and Triplaris gardneriana and late succes-
naceae; Vochysia divergens, Vochysiaceae) and late successional species, Ocotea diospyrifolia. We observed that the bark
sional species (Handroanthus heptaphyllus, Bignoniaceae; traits differ between pioneer and late successional species (n =
Ocotea diospyrifolia, Lauraceae; Vitex cymosa, Lamiaceae; 18; t-value 2.33; p-value = 0.03), where RBT seems to be the
Damasceno-Junior et al. 2005). Histological slides of the bark major bark trait to determine the successional groups than the
were prepared following standard techniques. We estimated bark density (pioneer = 0.39 g/cm3; late successional = 0.40 g/
the relative bark thickness (RBT) as the total bark thickness di- cm3) and percentage of sclerenchyma (pioneer = 30.8%; late
vided by bole diameter multiplied by 100 (Midgley and Lawes successional = 32.6%) (Figure 1f).
2016). Bark density was calculated as dry mass per fresh vol- Contrary to our expectation, late successional species
ume (Borchert 1994). We estimated the area of sclerenchyma (4.61%) have thicker bark than pioneer species (2.99%).
(sclereids and fibres cells; %) in cross-sections of the inner bark Thicker bark in late successional species can act in stem
(secondary phloem and cortex, if any). We performed multidi- protection from decay and against pathogens and herbivores
mensional scaling (MDS) analysis based on a Bray-Curtis dis- (Loehle 1988) which these species are prone to during periods
similarity matrix and used mixed models to test the difference of flooding in riparian forests. At the same time, thick bark
between species groups using species as a random factor. consisting of rhytidome or phellem with numerous layers of
A single periderm was observed in pioneer species (Figure thick-walled cells can play a key role in cambium protec-
1a) and Handroanthus heptaphyllus, a late successional spe- tion from lethal temperatures during fires. In the face of the
cies. In this species, the phellem is formed by numerous layers alarming increase of fire events in the riparian forest of the
of thick-walled, sclerified cells interspersed with few layers Pantanal, thick bark will be an important strategy and adaptive
of thin-walled cell layers (Figure 1b). In Vochysia divergens functional trait for woody species in these communities. Bark
(a pioneer species) the formation of aerenchymatous phellem adaptation in pioneer species involves the development of aer-
(Figure 1d) was observed. In Vitex cymosa (Figure 1c) and enchymatous phellem in Vochysia divergens, a feature that can
increase aeration in the bark tissues caused by anoxia during
1 Correspondence author contact: janersbio@gmail.com
2 Laboratório de Anatomia Vegetal, Instituto de Biociências (INBIO), Universi- flooding (Yáñez-Espinosa and Terrazas 2001) and the accu-
dade Federal de Mato Grosso do Sul, Campo Grande, Mato Grosso do Sul, Brazil. mulation of phenolic compounds in Inga vera and Triplaris
3 Agência de Tecnologia da Informação e Comunicação, Universidade Federal de
Mato Grosso do Sul, Campo Grande, Mato Grosso do Sul, Brazil.
gardneriana, which provides chemical defense in the bark
4 Programa de Pós-Graduação em Biologia Vegetal, Universidade Federal de (Roth 1981). Production of these compounds is stimulated by
Mato Grosso do Sul, Campo Grande, Mato Grosso do Sul, Brazil. flooding in several species (Yule et al. 2018). n
REFERENCES FIGURE 1. Examples of bark traits and ordination results: (a) General aspect of stem bark in transverse
Angyalossy, V., M. Pace, R.F. Evert, C.R. sections of the pioneer species Triplaris gardneriana. Note the layers of axial parenchyma and conducting
Marcati, A.A. Oskolski, T. Terrazas, E. cells alternating with layers of sclerenchyma (arrows) in the inner bark. (b) Phellem (Ph) with thick-
Kotina, F. Lens, S.C. Mazzoni-Viveiros, walled, sclerified cells in Handroanthus heptaphyllus, a late successional species. (c) Rhytidome (Rh)
G. Angeles, S.R. Machado, A. Crivellaro,
K.S. Rao, L. Junikka, N. Nicolaeva, and P.
with successive development of periderms (arrowhead) in a late successional species Vitex cymosa. (d)
Baas. 2016. IAWA list of microscopic bark Aerenchymatous phellem in the pioneer species Vochysia divergens. (e) Secondary phloem in Vochysia
features. International Association of Wood divergens with large sclereids clusters (*). (f) Secondary phloem in a late successional species Ocotea
Anatomists (IAWA) Journal 37: 517–615. diospyrifolia with small (arrows) and large (*) sclereids groups. (g) Multidimensional scaling (NMDS)
doi:10.1163/22941932-20160151. ordination based on a Bray-Curtis matrix of dissimilarities showing the distribution of the pioneer and late
Borchert, R. 1994. Soil and stem water successional species in the riparian forest of the Paraguay River. Scale bars: (a) = 500 µm; (b, c, d and e)
storage determine phenology and distribu- = 100 µm; (f) = 100 µm.
tion of tropical dry forest trees. Ecology
75(5): 1437–1449. doi:10.2307/1937467.
Damasceno-Junior, G.A., J. Semir, F.A.
Maës Dos Santos, and H. De Freitas
Leitão-Filho. 2005. Structure, distribution
of species and inundation in a riparian
forest of Rio Paraguai, Pantanal, Brazil.
Flora 200(2): 119–135. doi:10.1016/j.
flora.2004.09.002.
Midgley, J.J., and M.J. Lawes. 2016. Rela-
tive bark thickness: towards standardised
measurement and analysis. Plant Ecology
217(6): 677–681. Springer Netherlands.
doi:10.1007/s11258-016-0587-8.
Roth, I. 1981. Structural Patterns of Tropi-
cal Barks. Gebrüder Borntraeger, Berlin.
Yáñez-Espinosa, L. and T. Terrazas. 2001.
Wood and bark anatomy variation of An-
nona glabra L. under flooding. Agrocien-
cia 35(1): 51–63.
Yule, C.M., Y.Y. Lim, and T.Y. Lim. 2018.
Recycling of phenolic compounds in Bor-
neo’s tropical peat swamp forests. Carbon
Balance Management 13: 1–14.

Mangrove Ecological Restoration Inside Pantanos de Centla Biosphere Reserve
(Tabasco, México)
Raúl Alejandro Betancourth1,2, Pilar A. Gómez-Ruiz3, and Paulo Carbajal-Borges2
antanos de Centla Biosphere Reserve (PCBR) is located the Universidad Autónoma del Carmen and Biomasa A.C
P in southern México within Tabasco state (Figure 1).
It was declared as a Natural Protected Area by Federal
NGO. The main actions executed during the project were:
• Evaluation of socio-ecological conditions to deter-
Government in 1992 and in 1995 it was designated as a mine the technical viability and necessity of man-
Wetland of International Importance (RAMSAR Conven- grove ecological restoration.
tion Site 0733). PCBR is considered the most extensive and
important wetland in Mesoamerica, being a strategic area • Implementation of ecological restoration actions on
due to the presence of migratory species and high biodiver- 50 hectares through reforestation and hydrologic
sity richness. Main ecosystems in PCBR are hydrophytic rehabilitation of mangroves.
communities like popal-tular vegetation and mangroves • Conducted training workshops to strengthen local ca-
represented by associations of Rhizophora mangle, Lagun- pacities for conserving and managing mangroves, with
cularia racemosa and Avicennia germinans distributed in gender inclusion as a strategy for climate change adap-
fringe and riverine mangrove types. tation and extreme weather phenomena risk prevention.
Since PCBR is a protected area, a series of measures The project was implemented at El Palmar and Tem-
for conserving and protecting ecosystems biodiversity have bladeras ejidos, both located on northern PCBR (Figure
been developed including wildlife monitoring, mangrove 1). El Palmar community has 87 people and Tembladeras
restoration, firewall breaches construction, forest watch has 122 people. These are communities that have commu-
and environmental education. However, natural protected nal property control over land, and thereby make collec-
areas are not exempted from impacts associated with global tive decisions via an assembly of communal landowners
climate change, and for that reason PCBR managers recently (ejidatarios). Their main economic activity is fishing for
started to implement a plan for mitigation and adaptation to self-consumption and selling in local markets.
climate change, to increase ecosystem resilience, to protect
biodiversity, and to minimize vulnerabilities of local commu- PARTICIPATORY ECOLOGICAL RESTORATION PROCESS
nities that inhabit this important region of southern México. Project development was done through a participatory
Because of these needs, the “Resiliencia Project” has been framework, where local knowledge and experience play a
developed to address this environmental problem in different fundamental role. Every decision related to actions, design,
protected areas of Mexico. Its main objective is to decrease techniques, and work seasons was made after dialogue and
direct and adverse impacts of climate change on biodiversity coordination among communities, local authorities and the
and human communities through strengthening the effective- PCBR managers.
ness of management and spatial configuration of México´s A diagnostic evaluation was carried out with socio-
Natural Protected Areas. This project has been implemented ecosystemic focus to evaluate the viability of different sites
in 17 Natural Protected Areas in México, including PCBR. for restoration, both in El Palmar and Tembladeras. To do
The initiative was funded by the Global Environmental Fund this, we used three sources of information: 1) cartographic
(GEF) and was implemented by United Nations Develop- analysis, 2) gathering of local knowledge, and 3) ecologi-
ment Program (UNDP) and National Commission of Natural cal evaluation. Through cartographic analysis, we found
Protected Areas (CONANP in Spanish). vegetation distribution patterns that were helpful in locat-
Project implementation in PCBR was conducted between ing biological corridors. We were also able to correlate
2019 and 2020 by the Mexican NGO - Foro para El Desarrollo geography with historical data on wildfires and obtain local
Sustentable A.C., with academic and technical support from knowledge about fire-prone areas that helped us detect and
avoid sites that are under constant threat from fire.
1 Corresponding author contact: rabetancourthb@gmail.com.
2 Foro para el Desarrollo Sustentable A.C., San Cristóbal de Las Casas, Chiapas, According to the analysis of all this information, the
México. most suitable restoration action for ejido El Palmar was
3 CONACYT-Universidad Autónoma del Carmen, Campeche, México.

mangrove reforestation with Rhizophora mangle propa- fires. This participatory ecological restoration process aims,
gules (Figure 2). On the other hand, for ejido Tembladeras in the short- and long-terms, to recover mangrove ecosys-
the needed action was hydrologic rehabilitation through tem at landscape level and to increase local resilience and
cleaning natural channels in a previously reforested area improve climate change adaptation by local communities
where mangroves grew up without control and caused hy- inside PCBR. n
drologic connectivity problems at
the landscape level. A few months FIGURE 1. Pantanos de Centla Biosphere Reserve (PCBR) and case study area localization. (A) General
later, during the first monitoring overview of Mexico with study area indicated, (B) state of Tabasco, indicating PCBR location, (C) PCBR area
evaluation of restoration activi- with case study locations “ejidos” El Palmar and Tembladeras and (D) Amplification of both ejidos surface.
ties, we recorded a global survival Elaborated by Juan Paulo Carbajal Borges based on Instituto Nacional de Estadística y Geografía (INEGI).
of 75% of R. mangle propagules 2010. Red Carretera de México, Comisión Nacional de Áreas Naturales Protegidas (CONANP) 2017. Áreas
Naturales Protegidas Federales, Registro Agrario Nacional (RAN) 2017. Perimetrales de Núcleos Agrarios.
planted in ejido El Palmar. In the
case of Tembladeras, fish spe-
cies of interest were captured in
rehabilitated channels, providing
evidence that hydrologic connec-
tivity is recovering.
Training workshops were
planned to cover these topics:
mangrove ecosystem services,
methodological phases of man-
grove ecological restoration
projects, and restoration monitor-
ing techniques. In addition, a basic
course on forest fires was given,
with the approval of the National
Forest Commission (CONAFOR
in Spanish). A community-expe-
riences exchange was done with
a neighborhood community that
had wide experience in mangrove
sustainable use and restoration
practices in Tabasco state. In this FIGURE 2. Hydrophytic vegetation in channels or “picadas” where reforestation action was implemented in
activity, people from El Palmar ejido El Palmar, PCBR. (Photo by Alejandro Betancourth.)
and Tembladeras communities
visited a mangrove nursery, re-
stored sites with L. racemosa, and
a charcoal production operation.
Currently, Foro para el De-
sarrollo Sustentable A.C. together
with its academic and technical
allies, are continuing the initia-
tive through a variety of activities
including additional restoration
actions, developing a protection
strategy for restored sites through
establishment of firewall breach-
es, designing local climate adap-
tation and disaster risk reduction
plans, and implementing early
warning systems for mangrove

Diversity of Temporary Ponds from the Guajira, Colombia
Cesar E. Tamaris-Turizo1, Pedro Eslava, Juan M. Fuentes-Reinés, Daniel Serna-Macías, Diana Tamaris, and Luis Castro
emporary ponds are ecosystems of great importance for Biological samples were taken during wet period of
T the biodiversity of the arid region of northern Colom-
bia. These temporal systems support resident and migratory
2018. The plankton samples were collected using a 25-l
bucket at both littoral and limnetic habitats. Samples were
animals including birds, frogs, snakes, plankton, macroin- filtered with a zooplankton net (mesh size = 45 μm) and
vertebrates, and fishes. These organisms display strategies, phytoplankton net (mesh size = 23 μm) and preserved in
for example, some invertebrates burrow their larvae in the 70% ethanol. Macroinvertebrates associated with macro-
sediment or produce resistant eggs in diapause state, that phytes were taken with a hand net (mesh size = 300 μm)
permit them to resist the drought periods that could last and macroinvertebrates associate with sediment with an
more than six months. Despite their importance to local Ekman dredge. The fishes were taken by tie and hand net
and migratory species, temporary ponds in Colombia have (both, mesh size = 500 μm), some fishes were transported
received little attention from researchers. to laboratory to confirm the species. Birds were identified
The aim of our study was to characterize the biodiver- though free tours around the ponds from 5:00 am to 9:00
sity of five temporary ponds located on the middle Guajira am and from 4:00pm to 6:00pm.
(Maicao, Manaure, Uribia, Mayapo and El Ebanal) of The fauna found in the systems of temporary ponds of
northern Colombia and compare the communities between La Guajira contain typical tropical species. More than 12,000
sites (Figure 1). Examples of these ponds are shown in organisms were collected and observed, belonging to five
Figure 2. large groups: macroinvertebrates, microphytes, fishes, birds,
and zooplankton). A total of 10,465 macroinvertebratres
were collected, representing 30 genera
FIGURE 1. General locations of the temporary ponds (The base image was taken from https://www. and 20 families. Among this group,
laguajira.gov.co/web/la-guajira/division-politica-administrativa.html). Triops (tadpole shrimp), Dendrocepha-
lus (fairy shrimp), and Thamnocephalus
(fairy shrimp) were the most abundant
genera with 3,207, 2,269 and 3,169
individuals tabulated, respectively.
Triops and Thamnocephalus are first
records from La Guajira. A total of 204
taxa of microphytes were identified,
grouped in 66 genera and 40 families.
Euglenophyceae was the family with
greatest richness (25% of all families)
whereas Trachelomonas and Lepocin-
clis were the most abundant genera of
euglenoids. Only seven species of fishes
were found Austrofundulus guajira (kil-
lifish), Rachovia hummelincki (killifish),
Astyanax magdalenae (tolomba), Mugil
incilis (mullet), Dormitator maculatus
(whitebait), Eleotris amblyopsis, and
Ctenolucius hujeta (hujeta gar). The two
1 Grupo de Investigación en Ecología y Biodiversidad, Universidad del Magda-
lena, Santa Marta, Colombia; corresponding author contact: ctamaris@unimagda-
lena.edu.co.

former species occurred in isolated ponds of fluvial systems FIGURE 2. Sampled temporary ponds in the northern Colombia: a)
while the others were found in ponds with influence of lotic Maicao, b) Uribia, c) Manaure, d) Mayapo, and e) El Ebanal. (Photos by
ecosystems. Fifty-two bird species were observed, with the P. Eslava). Note: The first four ponds (a, b, c and d) are temporary ponds
Carib grackle (Quiscalus lugubris) being most frequent. The isolated from fluvial systems, while the last pond (e) is connected with
zooplanktonic community were represented by eight taxa the Ranchería River.
of Rotifera (Lecane sp, Platyas brevicornis, Asplanchna sp,
Brachionus caudatus, B. quadridentatus quadridentatus, B.
mirabilis, Filinia sp., and Dicranophorus sp), 18 of Cladoc-
era (Diaphanosoma breverime, Diaphanosoma spinulosum,
Diaphanosoma dentatum, Sarsilotona serricauda, Pseudosi-
da sp., Ceriodaphnia cornuta, Moinodaphnia macleayi, Moi-
na micrura micrura, Moina reticulata, Macrothrix elegans,
Macrothrix spinosa, Grimaldina freyi, Kurzia polyspina,
Leydigia cf striata, Ovalona cf glabra, Chydorus nitidulus,
Dunhevedia odontoplax, and Dunhevedia crassa) and four
taxa of Copepoda (Prionodiaptomus colombiensis, Microcy-
clops cf ceibaensis. Mesocyclops brasilianus, and Thermo-
cyclops tenuis). Of these Platyias quadricornis quadricornis,
Brachionus caudatus, B. quadridentatus quadridentatus,
B. mirabilis, Prionodiaptumus colombiensis, Mesocyclops
brasilianus, and Thermocyclops tenuis are new records to the
Department of La Guajira. The most abundant zooplankton
species was a copepod - Prionodiaptomus colombiensis (860
individuals) whereas the least abundant was a rotifer - As-
planchna sp. with only two individuals collected. Lecane sp.,
Thermocyclops tenuis, Microcyclops cf ceibaensis, Moina
micrura, Moinodaphnia macleayi, Diaphanosoma spinolu-
sum, and Leydiga cf striata were the most frequent taxa. The
temporary pond with the greatest richness of zooplankton
was Maicao pond with 20 species, while El Ebanal pond had
the least richness with only nine species.
Overall, Maicao ponds had the highest biodiversity,
probably due to the habitat heterogeneity marked by the oc-
currence of different patches of macrophytes (i.e., Eleocha-
ris elegans and Paspalum sp.) and scrubs (i.e., Caesalpinia
coriaria and Parkinsonia praecox). Research about the
community structure in temporal ponds in Colombia is
scarce and it is expected that the results of our study will
help stimulate similar investigations in order to generate
local listings that reveal new records of species occurrence
and, consequently, to better understand the diversity of the
fauna of Colombia’s temporary ponds. n

Treatment Wetlands - Experiencies in Mexico
Armando Rivas1
installed in several communities. Each system is composed

I n Mexico treatment wetlands are mostly utilized for waste-
water treatment, mainly due to their low operating costs,
simple operation, aesthetics and successful experiences.
of: pretreatment (screening and grit removal), septic tank,
vertical subsurface flow wetland (sludge wetland planted
Approximately 40% of the treatment infrastructure with Arundo donax), subsurface flow wetland (planted
(mainly systems requiring electricity) are out of operation, with Typha sp.), and maturation pond and subsurface flow
due to, among other causes, insufficient economic re- wetland (with Typha sp.). Treatment wetlands were created
sources and slightly trained personnel. There is therefore a in five communities (2004 in Cucuchucho, 2005 in Santa
need for new technological alternatives to help solve this Fe de la Laguna, 2006 Erongaricuaro, 2007 San Francisco
serious problem. Uricho and two more, 2007, at San Jeronimo Purenchecu-
In the Patzcuaro Lake basin, the installed treatment aro. So by 2007, a total of six wetland treatment systems
capacity (activated sludge and oxidation ditches) is 190 L/s, had been built in the Patzcuaro Lake basin.
of which 52 L/s are treated (72% of the flow rate was used Prior to the construction of the systems, studies of
for agricultural activities before getting to treatment plant). wastewater characteristics, topography and soils mechan-
There is also the problem of discharging residual sludge. ics were carried out. After one year of operation, the first
The problem is compounded by the direct discharge, into system (Cucuchucho locality, for 1.0 L/s) was evaluated
the lake, of wastewater generated in agricultural and indus- during the dry season (December to May) by four sam-
trial activities, which cause eutrophication into the lake. An pling campaigns. Average influent and effluent values
additional problem, perhaps the most important is con- (and removal efficiencies –%-) were: BOD 454 and 12
cerned to social considerations, since there is an obvious mg/L (97%); TN 79 and 10 mg/L (87%); TP 17 and 6
rejection of conventional systems for the above mentioned mg/L (65%); FC 3.42E 06 and 5.72E 02 MPN/100 mL
reasons and because people don’t like the bad odors gener- (99.9834%), and helminths 12 and 0 (100%). The Mexican
ated from wastewater treatment facilities. regulation (NOM-001-SEMARNAT-1996, discharge into
As an alternative solution and through the imple- lakes and protection of aquatic life) are: BOD 30 mg/L,
mentation of an intensive social participation program TN 15 mg/L; TP 5 mg/L, Helminths 1/L and FC 1000
(workshops, videos, brochures, visits to wetlands in opera- MPN/100 mL. So for this treatment all parameters were
tion, legal aspects of land, etc.) treatment wetlands were met except for the TP standard. Figure 1 shows the BOD
FIGURE 1. Results for Cucuchucho wetland treatment system during first year of operation.
1,000 BOD
TN
TP
Constituents, mg/L
100
10
1
Inlet Septic Wetland Pond Wetland
Tank 1 2
1 Instituto Mexicano de Tecnología del Agua. Av. Paseo Cuauhnáhuac Nº 8532

Col. Progreso. Jiutepec. Morelos. México. C.P. 62550; rivas.hz@gmail.com

results (graph on left) during the four sampling campaigns maize), and 4) areas to protect wildlife (lake birds). The
(dry season) and the average results of BOD, TN, and TP successful application of wetland systems for wastewater
for each component of the system (graph on right). treatment now serves as a model for other communities
One year later, a second system (3.0 L/s) was built in (total of five built systems) and have been used for educa-
the community of Santa Fe de la Laguna. The obtained tion on the value of natural treatment systems in workshops
efficiencies were similar to those achieved in the Cucu- of water culture for several communities. In addition, the
chucho system. wetlands are frequently visited by schoolchildren, academ-
The impact of treated water on lake water quality was ics, and people interested in the use of this technology for
subsequently assessed by a study of bio indicators (macro- wastewater treatment.
invertebrates), used in their concept of Family Level Biotic After installation of the five systems, water quality
Index (FBI), in four sampled points (locations with or studies were carried out at the sites where the treated water
without treatment wetlands). After two years of the start-up is discharged, either through wetlands or with electrome-
of Cucuchucho system (first evaluated point), the FBI was chanical systems.
12; at the second point (Santa Fe de la Laguna, a system The results, after ten years of operation, indicate
operating after one year of the start-up), the FBI was 10; there is a lower concentration of nutrients (TN and TP) in
the third point, near Opongio community (blank site, inside the lake where wetlands and lagoons are functioning, in
the lake, with no treatment plant or wastewater discharge) comparison with where the electromechanical systems are
the FBI was 9; and the fourth place (Tzintzuntzan commu- employed. It was also observed that the efficiency of TP
nity, without treatment plant, therefore direct discharge of removal by conventional treatment plants is decreasing
waste water into the lake) the FBI was 0. after the start-up; a cause, which is likely to contribute to
The BOD, inside the lake, near the discharge of treated this result, could be due to insufficient activities of opera-
water, before the installation of the treatment wetland, tion and maintenance.
was 414 mg/L. After three years of water treatment by the In conclusion, the use of treatment wetlands has
wetland system BOD dropped to 88.9 mg/L, indicating a produced successful results by improving the quality of
significant improvement in water quality within the lake lake’s water, protecting the lake’s biodiversity and health
from wastewater treatment by wetlands. Other benefits of local people and livestock, providing additional in-
by using wetlands were: 1) sale of ornate flowers (Zant- come through the sale of by-products, and creating areas
edeschia aethiopica, placed on the periphery to improve for wild life protection. Perhaps best of all, the wetland
aesthetics), 2) sale of Typha angustifolia as raw material for treatment systems keep on working with a minimum of at-
handicrafts manufacturing, 3) treated water that complies tention and have been widely accepted in social terms by
with the regulations for crops irrigation (vegetables and local communities. n

NOTES - WETLAND CONSERVATION/EDUCATION/OUTREACH
An Ecosystem-based Approach to Managing Fish, Cattle and Forests on the Amazon

Floodplain
David G. McGrath1,2, Antonia Socorro Pena da Gama2,5, Laura L. Hess3, Bruce Forsberg4, Antonio José Mota Bentes5,
Poliane Batista5
he Lower Amazon floodplain is the vast fluvial wet-

T land associated with the Amazon River upstream of its
internal delta and estuary. It extends from above the mouth
introduced by Japanese colonists in the 1930s and after
WWII, it expanded throughout the Amazon floodplain,
reaching a peak in the 1970s before declining through the
of the Xingu River in the East to the border between the mid-1980s when production ended. Since the colonial
states of Pará and Amazonas in the West and has a total period, the natural grasslands of the floodplain have been
area of 16,000 km2, of which roughly 11,000 km2 is season- used to graze cattle. This intensified in the 1980s with the
ally inundated (Figure 1). It is characterized by large open seasonal movement of cattle between floodplain and up-
water bodies surrounded by seasonally flooded grasslands lands. Today, cattle ranching and fishing are the dominant
and forests that vary in extent and inundation state during resource use activities on the floodplain. Between the mid-
the annual flood cycle. Forests occupy the levees border- 1970s and 2008 more than 50% of forest cover was cleared
ing river channels while grasslands occupy the transition and today there is no primary forest on the Lower Amazon
zone between the levees and the open water bodies of the floodplain (Renó et al. 2016).
floodplain interior (Figure 2). Cattle ranching is practiced on a range of scales from
The river level peaks in early June, falls to its minimum artisanal fishers who raise a few head of cattle to large-
level in early November and then rises gradually from De- scale ranchers with herds of several hundred. Deforestation
cember to June. Flood amplitude is about 6 meters, with the and degradation of forests and grasslands is, in part, the
highest floods exceeding 8 meters. At peak high-water level result of the economic strategies of small-scale fishers who
approximately 95% of the floodplain is inundated, with in addition to fishing in floodplain lakes, practice shifting
open water, flooded grasses and flooded forests occupying cultivation for annual crops and raise cattle on community
59, 21 and 20% of this area, respectively (Hess et al. 2003). grasslands (Figure 2). Within household economic strate-
At low water, only 67% of the floodplain is flooded, with gies, fishing provides subsistence and cash income, while
open water, flooded grasses and flooded forests occupying cattle serve as a source of savings. While small-scale fish-
69, 18 and 13% of this flooded area, respectively (Figures ers understand the close relationship between floodplain
3 and 4). Many trees have adapted to this flood regime by fisheries and forests, many continue to raise cattle, under-
producing fruits and seeds that mature as floodwaters reach mining management efforts to increase the productivity of
their peak, providing an important food source for fish, local fisheries.
which in turn act as a key dispersal agent for the plants The tendency for smallholders to invest in cattle as a
(Goulding et al. 2019; Schöngart et al. 2002). Many spe- savings strategy, despite the consequences for the local
cies of fish, including important commercial species such fisheries on which they depend for their cash income, is
as Characins, use the floodplain as nursery habitat and – related to the perceived security of investments in cattle
once mature – as seasonal feeding habitat. As adults these versus those in managing the fishery. Insecurity regard-
species move out of the floodplain to migrate upstream to ing fisheries is exacerbated by the lack of government
spawn during rising water levels and then move back onto enforcement of management agreements. Property rights
the floodplain to feed during the high-water period. to fish in a lake are diffuse and follow the law of capture,
The Lower Amazon is the most deforested section of while property rights to cattle grazing on collectively
the Amazon floodplain. During the 18th and 19th Centuries, owned grasslands are clear and enforced by the state.
levee forests were cleared for cacao plantations. Jute was Consequently, smallholders invest in fishing for immedi-
ate needs (subsistence and cash), while for long-term sav-
1 Earth Innovation Institute, San Francisco, CA, USA; ings, they invest in cattle. Until these conditions change,
2 Federal University of Western Pará, Santarém, PA, Brazil; it is unlikely that smallholders will agree to reduce their
3 Earth Research Institute, University of California , Santa Barbara, CA, USA;
4 Vermont Department of Environmental Conservation, Montpelier, VT, USA; investment in cattle to conserve the habitat that improves
5 Sapopema (Sociedade para a Pesquisa e Proteção do Meio Ambiente), San- fisheries productivity.
tarém, PA, Brazil.

To address the problems of FIGURE 1. Location map showing the Lower Amazon region centered on the city of Santarém, at the
overfishing in community lakes confluence of the Tapajós and Amazon Rivers. The box shows the area covered in Figure 3.
and overgrazing of floodplain
grasslands, communities have
implemented inter-community
fishing agreements and formal
contracts between individual com-
munities and local cattle owners.
These informal fishing agreements FIGURE 2. Transect of the Lower Amazon floodplain showing main habitat types and resource use by
led to the development of a co- floodplain smallholders.
management policy that provides
a pathway for their legal recogni-
tion. Communities then adapted
this model to negotiate agreements
between communities and local
cattle owners to regulate cattle
grazing on floodplain grasslands.
These were legalized through the
Public Ministry via a contract
called a Term of Adjustment of
Conduct (TAC). Some 50 TACs
FIGURE 3. Vegetation cover of Lower Amazon floodplain at low-water stage (top) and high-water stage
were negotiated by communities
(bottom).
in the region. Between 2006 and
2008, community territories were
incorporated into 41 Agro-extrac-
tive Settlement Projects (PAE),
a type of extractive reserve for
traditional populations, with a total
area of 740,000 hectares. Both
agreements were incorporated into
the Utilization Plans of the PAE,
creating the potential for the inte-
grated management of settlement
resources (Figure 5). However,
lack of enforcement continues to
2018). These studies make it possible to calculate the cost
be a problem.
to fishers of forest and grassland degradation caused by
Floodplain residents understand the importance of
cattle and shifting agriculture. These economic estimates
floodplain forests and grasslands to their livelihoods, not
can then be used to generate scenarios so fishers can clearly
only as feeding and nursery habitat for fish, but also for
see the costs of investing in cattle when designing lake
controlling bank erosion and as wave barriers to protect
management plans that regulate fishing and cattle grazing
houses during flood season thunderstorms. There have been
and reforestation of floodplain lake margins. We plan to
numerous initiatives to reforest levees and replant aquatic
use this approach in working with floodplain communi-
macrophytes, but these are often frustrated by uncontrolled
ties to negotiate more effective agreements for regulating
cattle grazing.
cattle and to justify community investments in large-scale
In recent years, several research initiatives have docu-
reforestation projects that improve the productivity of lake
mented the close relationship between the extent of forest
fisheries and increase aquatic biodiversity and the resilience
cover surrounding lakes and the productivity of fishing
of floodplain fisheries and household subsistence strategies.
effort at both the regional scale and through comparison
Our strategy going forward builds on these community
of fishing productivity, fish diversity and fish commu-
initiatives for reforesting lake margins, planting floating
nity structure in individual floodplain lakes with varying
grass storm barriers and other local approaches to conserv-
amounts of forest cover (Castello et al. 2018; Arantes et al.
ing habitat. This partnership harnesses the potential of
universities in Santarém through collaborations with local Hess, L. L., J. M. Melack, E. M.L.M. Novo, C. C.F. Barbosa, and M.
Gastil. 2003. Dual-season mapping of wetland inundation and vegeta-
NGOs, community schools and community management tion for the central Amazon basin. Remote Sensing of Environment 87:
organizations, combining science, local knowledge and 404–428.
local organizational capacity to sustainably manage flood- McGrath, D. and M. Crossa. 1998. Restoration of floodplain lake habitat:
plain fisheries, grasslands and forests. n a PLEC demonstration project. PLEC News and Views 11: 10-17.
McGrath, D., A. Cardoso, O.T. Almeida, and J. Pezzuti. 2008. Con-
REFERENCES: structing a policy and institutional framework for an ecosystem-based
Arantes, C. C., K.O. Winemiller, M. Petrere, L. Castello, L.L. Hess, approach to managing the lower Amazon floodplain. Environment,
and C.E.C. Freitas. 2018. Relationships between forest cover and fish Development and Sustainability 10: 677-695.
diversity in the Amazon River floodplain. Journal of Applied Ecology
55: 386–395. Renó, V., E. Novo, and M. Escada. 2016. Forest fragmentation in the
Lower Amazon Floodplain: Implications for biodiversity and ecosystem
Castello, L., L. L. Hess, R. Thapa, D. G. McGrath, C. C. Arantes, V. F. service provision to riverine populations. Remote Sensing 8: 26.
Renó, and V. J. Isaac. 2018. Fishery yields vary with land cover on the
Amazon River floodplain. Fish and Fisheries 19(3): 431-440. Schöngart, J., M. T. F. Piedade, S. Ludwigshausen, V. Horna, and M.
Worbes. 2002. Phenology and stem-growth periodicity of tree spe-
Goulding, M., E. Venticinque, M. L. de B. Ribeiro, R. B. Barthem,R. G. cies in Amazonian floodplain forests. Journal of Tropical Ecology 18:
Leite, B. Forsberg, P. Petry, U. Lopes da Silva-Júnior, P. Santos Ferraz, 581–597.
and C. Cañas. 2019. Ecosystem-based management of Amazon fisheries
and wetlands. Fish and Fisheries 20(1): 138-158.
FIGURE 4. Contrasting low-water (left) and high-water (right) conditions, Curuai Lake, Amazon
floodplain. PlanetScope imagery for 2 November 2017 (low) and 29 July 2017 (high). Images
courtesy of Planet Labs.
FIGURE 5. Distribution of Agro-extractive Settlement Projects (PAE) of the Lower Amazon floodplain.

Strengthening Governance in the Monterrico Multiple Use Natural Reserve: Planning
for Conservation with a Bottom-Up Approach
Ana Silvia Morales1
he Monterrico Multiple Use Natural Reserve to improve the current biodiversity and ecosystem services
T (MMUNR) is an important protected wetland with
high conservation values located on the Pacific Coast of
management by creating new and pertinent models for
protected areas management.
Guatemala (Figure 1). It is comprised of estuarine and In this regard, after an assessment of the current
coastal-marine ecosystems that provide goods and services wetland management approach, it was necessary to rede-
to the local people whose livelihoods depend on them, sign the way CECON was interacting with the local com-
such as fisheries, tourism, coastal protection, among others. munities to achieve better conservation outcomes (Figure
Since the MMUNR was created, like most of the protected 2). Taking into account that the MMUNR is a wetland of
areas back in the 1970s, there was no consultation pro- considerable relevance for local people, CECON is work-
cess with local communities to obtain prior and informed ing on a governance platform through updating its master
consent to be part of conservation effort. The reserve is plan with a bottom–up approach to include inputs from
managed by the Center of Conservation Studies (CECON) local communities regarding their perceptions and concerns
of the University of San Carlos, a research center that also about conservation of ecosystem goods and services for
manages six other protected areas. Their objectives are to reducing threats to the MMUNR wetland, and also to get
conduct permanent biodiversity research programs aimed local people to interact with public and private sectors. This
updating process is also consider-
FIGURE 1. Location of the MMUNR in the Pacific Coastal of Guatemala, Central America (Miguel Ávila, ing actions beyond the borders of
CECON). the protected area, by focusing
on activities related to watershed
management taking place upstream
that are altering the natural hydro-
logical cycle of the wetland.
The main goals of this partici-
patory process are: 1) to strengthen
CECON as the protected area
manager with the continued sup-
port of the local communities and
the other stakeholders for decision
making, and 2) establishing a com-
mittee to create an environment for
sustainable use of the wetland eco-
systems through a mechanism of
effective and equitable governance
that includes all sectors involved in
the area. This process also contem-
plates an expansion of the role of
CECON as the MMUNR Manager
to include serving as both techni-
cal–scientific advisor and mediator
for conflicts related to the use of
1 Coordinator of the Monterrico Multiple Use Natural Reserve, Center of Conser-

vation Studies, University of San Carlos of Guatemala; ansilmo@gmail.com

biodiversity and natural resources. Moreover, recent inter- resources. CECON is recognized as a management author-
national cooperation has been directed to strengthen con- ity not only regarding the protected area but in general for
servation of coastal marine protected areas. In this regard, environmental concerns in other local communities outside
CECON is helping local communities implement produc- the borders. As a result of this ongoing shared work, a
tive sustainable practices in agriculture, strengthening local mutual collaboration agreement has been signed by both
community organization, and directing conservation incen- CECON and the local communities. The main challenge is
tives towards those stakeholders involved in conservation to consolidate this process with the active participation and
and restoration of the protected area. consent of the local communities together with the public
The governance process carried out so far has achieved and private sectors. In time, this can be achieved through
great success in terms of community involvement as local the equitable sharing of benefits from the wetland conserva-
people have become more aware of the relevance of work- tion for all stakeholders in the area. n
ing together for the conservation of biodiversity and natural
FIGURE 2. Guatemala’s Monterrico Multiple Use Natural Reserve is important to both people and wildlife. (Photos by Ana Silvia Morales and
Homero Escobar.)

Amphibious Colombia: A Country of Wetlands
Ronald Ayazo Toscano1, Wilson Ramirez, Ana Carolina Santos, Olga Lucía Hernández-Manrique, Angélica Batista, and
Margarita Roa
he Biological Research FIGURE 1. Map of Colombia and its amphibious nature prepared by César Aponte adapted from Amphibian
T Institute Alexander von
Humboldt is a civil non-
Colombia. A country of wetlands (Jaramillo Villa et al. 2015). This map shows the general distribution of five
wetland categories in the five hydrographic areas of Colombia: Amazon, Caribbean, Magdalena-Cauca, Orinoco,
profit organization associated and Pacific. (Source: The Biological Research Institute Alexander von Humboldt, Bogotá.)
with Colombia’s Ministry of
Environment and Sustainable
Development (MinAmbi-
ente). It is part of the Colom-
bian National Environmental
System (SINA), which aims
to store and share data to
generate knowledge about
Colombian environment and
ecosystems. The von Hum-
boldt Institute is recognized
internationally as an authority
on the status and trends of
biodiversity, invasive species,
biological collection, open
data, citizen science, ecologi-
cal restoration, ecohydrology,
and supportive knowledge
networks. The Institute con-
ducts scientific research on
biodiversity in the continental
territory of Colombia, includ-
ing freshwater resources. The
von Humboldt Institute has been an active participant in ous Colombia: A country of wetlands, the most complete
supporting MinAmbiente in the construction of Colombia’s and extensive scientific work about Colombian’s wetlands
National Wetland Policy (Naranjo et al. 1999). (Jaramillo Villa et al. 2015). This project created figures and
Colombia has a heterogeneous topography and geogra- images to guide the natural resource management and risk
phy that requires continuous study and up-to-date cartog- management of wetlands in the country, beginning with a
raphy to inform environmental policy-making. In contrast carefully developed set of maps (Figure 1).
to other countries of the Andes region, Colombia has three One of those maps, using official information later pub-
distinct mountain ranges separated by rivers, creating a lished by Flórez-Ayala et al. (2016), showed more than 30
diverse hydrographic network supporting a vast wetland million hectares of wetlands, encompassing 26% of the na-
ecosystems. To support its work, the von Humboldt Insti- tional inland area (see figure above). Wetlands were classi-
tute has several institutional associations, such as the Na- fied into four categories: open permanent, permanent under
tional Adaptation Fund who financed the project Amphibi- canopy, temporary, and potential. Open permanent repre-
sents wetlands where water presence is constant and there
is no tree cover (e.g., lakes, lagoons, ciénagas, rivers, and
1 Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, glaciers). Permanent under canopy identifies forested wet-
Bogotá, Colombia; Corresponding author: rayazo@humboldt.org.co.

lands that are always flooded (e.g., Atrato River swamps REFERENCES
or mangroves). Temporary wetlands do not have water Flórez-Ayala, C., L.M. Estupiñán-Suárez, S. Rojas, C. Aponte, M.
year-round, but are periodically wet. The last category Quiñones, O. Acevedo, S.P. Vilardy Quiroga, and Ú. Jaramillo Villa.
2016. Identification and mapping of Colombian inland wetlands. Biota
- Potential - identifies areas where soil and/or landform Colombiana 17(1):179–207. https://doi.org/10.21068/c2016s01a03
characteristics indicate the likely presence of a wetland, Jaramillo Villa, Ú., J. Cortés-Duque, and C. Flórez-Ayala (Editors).
although no flooding was detected during the analyzed pe- 2015. Amphibian Colombia. A country of wetlands. Alexander von
riod (2007-2011); this category must be examined at scales Humboldt Research Institute of Biological Resources. Bogotá D.C.,
Colombia. http://hdl.handle.net/20.500.11761/34868
with greater detail. Another map of interest identified and
classified 89 macro-habitats across marine-coastal, inland, Mesa-S., L.M., G. Corzo, O. L. Hernández-Manrique, C.A. Lasso, and
G. Galvis. 2017. Freshwater ecoregions from Colombia: a proposal for
and anthropogenic systems using the Brazilian wetland territorial planning of the Trasandean region and part of the Orinoco and
classification system with geomorphological adjustments Amazon basins. In: L.A. Moreno, G.I. Andrade, and L.F. Ruíz-Contre-
for Colombia (Ricaute et al. 2019). Lastly, in association ras (Editors). 2016. Biodiversity 2016. Status and Trends of Colombian
Continental Biodiversity. Research Institute of Biological Resources
with Ecopetrol (the Colombian State Petroleum Com- Alexander von Humboldt. Bogotá D.C., Colombia. http://reporte.hum-
pany), the von Humboldt Institute identified 28 freshwater boldt.org.co/biodiversidad/en/2016/cap4/406/index.html
ecoregions for Colombia, based on hydro-geographic and Naranjo, L., G. Andrade, and E. Ponce de León. 1999. Humedales
biological criteria like fish species composition, the inter- interiores de Colombia: Bases técnicas para su conservación y uso
sostenible.: Instituto de Investigación de Recursos Biológicos Alex-
pretation of their drainage network, and geomorphological ander von Humboldt y Ministerio del Medio Ambiente, Bogotá D.C.,
characteristics (Mesa-S et al. 2016). All these findings are Colombia.
provided to help improve and facilitate the creation of laws Ricaurte, L.F., J.E. Patiño, D.F.R. Zambrano, J.C. Arias-G, O. Ace-
and public policies that include the protection, restoration vedo, C. Aponte, R. Medina, M. González, S. Rojas, C. Flórez, L.M
and sustainable use of freshwater ecosystems necessary for Estupinan-Suarez, U. Jaramillo, A.C. Santos, C.A. Lasso, A.A. Duque,
S.R. Calle, J.I. Vélez, J.H. Caballero, S.R. Duque, M. Núñez-Avella-
the territorial planning. n neda, I.D. Correa, J.A. Rodríguez-Rodríguez, S. Vilardy, A Prieto-C,
A. Rudas-Ll, A.M. Cleef, C.M. Finlayson, and W.J. Junk. 2019. A clas-
sification system for Colombian wetlands: an essential step forward in
open environmental policy-making. Wetlands 39: 971–990. https://doi.
org/10.1007/s13157-019-01149-8

Amphibian Territories in Transition: Socio-ecological Rehabilitation of Wetlands
Ronald Ayazo Toscano1, Wilson Ramírez1, Klaudia Cárdenas1, Olga Lucía Hernandez-Manrique1, Natalia Gómez López2,
William Vargas2, Paola Isaacs-Cubides1, Mauricio Aguilar1, Yenifer Herrera1, Henry Huertas1, Juan C. Linares3, Wendy
López4, and Jorge Bedoya5
T he successful rehabilitation of a wetland

landscape requires more than a single res-
toration method, especially when productive
FIGURE 1. Map showing location of The Mojana and its wetlands prepared by Paola Isaacs-
Cubides. The inset map on the right shows the location of The Mojana in Colombia and the
location of Colombia in the Western Hemisphere. The enlarged map on the left shows rivers
activities are carried out there. In Colombia, (Drenaje Doble), marshes (Cienagas - open permanent wetlands), and freshwater swamp
the La Niña phenomenon from 2010-2011 forests (Zapales) along with the municipality boundary and the limits of The Mojana. (Source:
The Biological Research Institute Alexander von Humboldt, Bogotá.)
was particularly destructive, causing signifi-
cant damage to infrastructure, economy, and
human lives. For this reason, in The Mojana,
a process of rehabilitating wetlands through
socio-ecological services and ecosystem res-
toration was initiated to increase the adaptive
capacity of natural ecosystems and the liveli-
hoods of people dependent on it by reducing
the risk of flooding and drought associated
with climate change and its variability.
The Mojana Region is an internal delta
located at the south center of the Colombian
Caribbean region (Figure 1). It is a flood-
plain landscape with marshes, swamps,
lagoons, streams, rivers, and zapales (fresh-
water swamp forest of The Mojana). It is
part of the region known as the “Momposina
Depression” that is characterized by the
flood pulses of the rivers Magdalena, Cauca
and San Jorge. In addition, this region is a
candidate for World Heritage site (UNESCO
2020) and includes the marsh of Ayapel
a Wetland of International Importance
(Minambiente 2018). According to the latest
municipal measurement to the Multidimen-
sional Poverty Index, this region is also one
of the poorest in Colombia (DANE 2015)
and its alluvial valleys represent one of the
country’s highest levels of anthropic impact
according to the first spatiotemporal evalua-
tion of human pressure in Colombia (Correa
Ayram et al. 2020).
1 Instituto de Investigación de Recursos Biológicos Alexander von Humboldt.

Corresponding author contact: rayazo@humboldt.org.co.
2 Corporación Paisajes Rurales (Colombia)
3 Universidad de Córdoba (Colombia)
4 Ministry of Environment and Sustainable Development (Colombia)
5 UNDP - United Nations Development Programme (Colombia)

FIGURE 2. Fisherman in the marsh of Palo (Pescador en Ciénaga de Palo) by Klaudia Cárdenas. rescuing the cultural heritage of being a
This photo received the Best Wetland Photo awarded by Society of Wetland Scientists at the I mojanero (an amphibious culture with
Congreso Internacional y II Congreso Nacional de Ríos y Humedales. Colombia, 2017. behaviors, beliefs, and practices related
to the management of land and water
resources in The Mojana), and monitoring
with citizen science (Ayazo Toscano et al.
2020).
As a result, between 2016 and 2019,
it was possible to implement a socio-eco-
logical rehabilitation through a participa-
tory process that facilitated the recovery
of ecological connectivity for 4,822
hectares of this floodplain. To date, these
processes of social transformation and
ecological rehabilitation are expanding
the most successful interventions piloted
through the project to an additional elev-
en municipalities in The Mojana Region
(UNDP Colombia 2020). n
REFERENCES
Ayazo Toscano, R., W. Ramírez, and Ú. Jaramillo
Villa (Editors.). In Press. Territorios Anfibios en
Transición. Rehabilitación Socioecológica de
Colombia’s Ministry of Environment and Sustainable Humedales. Instituto de Investigación de Recursos Biológicos Alexander
Development, through a project funded by the Kyoto Pro- von Humboldt. Bogotá, D.C., Colombia.
tocol’s Adaptation Fund and in partnership with the United Correa Ayram, C.A., A. Etter, J. Díaz-Timoté, S. Rodríguez Buriticá, W.
Nations Development Programme Colombia, developed a Ramírez, and G. Corzo. 2020. Spatiotemporal evaluation of the hu-
holistic view for climate change and disaster risk manage- man footprint in Colombia: Four decades of anthropic impact in highly
biodiverse ecosystems. Ecological Indicators 117: 106630. https://doi.
ment for the region. This included a plan for the rehabilita- org/10.1016/j.ecolind.2020.106630
tion of degraded wetland ecosystems and ecosystems-based DANE. 2015. Pobreza Monetaria y Multidimensional: Principales Resul-
livelihoods as integral factors of the system necessary for tados 2014, Colombia.
both its operation and the well-being of the locals (Figure Jaramillo Villa, Ú., C. Cárdenas, R. Ayazo Toscano, W. Vargas, N.
2). In this context, the most feasible strategy to guarantee Gómez, J.C. Linares, M. Carillo, A. Martínez, and W. Ramírez. 2018.
the success and sustainability of this rehabilitation work Recuperar modos de vida, para rehabilitar ecosistemas: rehabilitación
del socio-ecosistema anfibio en la Mojana. In: L.A. Moreno, G.I.
was to design a plan based on understanding and recogniz- Andrade, and M.F. Gómez (Editors). 2019. Biodiversidad 2018. Estado
ing the interdependence of the region’s inhabitants and y tendencias de la biodiversidad continental de Colombia. Instituto de
natural ecosystems (Jaramillo Villa et al. 2018). Investigación de Recursos Biológicos Alexander von Humboldt. Bogotá,
D.C., Colombia. http://reporte.humboldt.org.co/biodiversidad/2018/
Through this initiative, the von Humboldt Institute in cap4/404/#seccion13
association with the University of Córdoba and the Rural Minambiente. 2018. Decreto N° 356. Diario Oficial de la República de
Landscapes Corporation worked with landowners, com- Colombia, Bogotá, Colombia, 22 de febrero de 2018
munities, local and regional governments to implement and United Nations Development Programme Colombia (UNDP Colombia).
monitor different social-ecological rehabilitation strategies 2020. Mojana: Climate and Life. Retrieved from United Nations Devel-
to enhance connectivity of wetland ecosystems. To account opment Programme. https://tinyurl.com/y9unbsut
for the extremes of droughts and floods in the region, activ- UNESCO. 2020. Pre-Hispanic Hydraulic System of the San Jorge River.
https://whc.unesco.org/en/tentativelists/5764/
ities focused on the household level by diversifying crops
in home gardens, rehabilitating the floodplain landscape,

Guardians of Wetlands (Los Guardianes de los Humedales): Young Peruvians
Committed to Wetlands
Héctor Aponte1
etland conservation is an arduous task; that is why After an arduous and challenging selection process,
W young people need to get closely involve in its sustain- 30 guardians were selected for the 2020 GDH program.
ability. The Universidad Científica del Sur, one of the Peruvian Between January and March 2020, the guardians received
universities that have promoted the study of wetlands in recent training on pre-Inca cultures and coastal wetlands, its func-
years, proposed an alternative that involves young university tioning and characteristics. Also, they had the opportunity
students. The Guardianes de los Humedales (Guardians of to do some handicrafts from marshes plants, using tradi-
Wetlands, GDH) program seeks to introduce young people to tional patterns. Likewise, they participated in field work-
environmental research and education related to the protection shops on the identification of native birds and plants as well
of coastal wetlands. The study of wetlands is a cross-cutting as they received training to identify the threats and impacts
area of interest; it is multi-disciplinary involving lines of to the wetlands visited.
research from several university departments such as environ- Once trained, the guardians participated in two diffu-
mental engineering, agribusiness, tourism, and biology. This sion activities, situated in the center and south of Lima.
program allows us to promote environmental research and These activities were focused on children, where we
education from different approaches such as tourism, biodi- explained the importance of wetland conservation through
versity management, business studies, evaluation of medicinal different play and learning strategies. Nowadays, the guard-
plants and land management in Peruvian coastal wetlands. ians are preparing virtual youth awareness activities for
The GDH project provides different activities to the the coming months of 2020, as well as meetings with other
students. For example, students receive courses and carry young people committed to wetlands (such as the Youth
out fieldwork, which allows them to learn about biodiver- Engaged in Wetlands initiative). We hoped that as a result
sity, culture and research in wetlands. The workshops in- of the GDH program, these students would be empowered,
clude training in dissemination techniques and awareness- and the network of professionals supporting the protection
raising on the protection of coastal wetlands. Therefore, we of wetlands will be strengthened. n
can reach a genuinely environmental awareness:
knowledge and motivation. FIGURE 1. Guardian of Wetland working with local youth, stimulating an interest in
wetlands and their conservation. (Photo by Héctor Aponte.)
The GDH program is committed to the training
of young people who can play a role as dissemi-
nating intermediaries of knowledge, and research
and conservation agents of the Peruvian coastal
wetlands. It is not possible to achieve environmen-
tal awareness without knowledge that sustains the
motivation for conservation and that is why we
consider the training part as the basis for this proj-
ect. This program is led by three wetland scientists
-Gustavo Lértora, Dámaso W. Ramírez, and Héctor
Aponte - who have the support of the Universidad
Científica del Sur (Perú) for the execution of the
program. It is the second time that this program
has been carried out and we are pleased to have the
support of the Humedales Costeros (humedales-
costeror.org) initiative this time.
1 Professional Wetland Scientist. Associate Researcher, Universidad Científica del

Sur, Lima-Perú; haponte@cientifica.edu.pe.

We Are Wetlands (@Somos_humedales)
Karina Paz Arteaga1
ince childhood I’ve been concerned about wetlands and promoting and facilitating the work of others learning about
S their protection. I began my journey when I discov-
ered the Porvenir bay wetland in my hometown (Porvenir
their experiences and thereby expanding one’s knowledge,
and 3) communicate in simple terms technical knowledge
Tierra del Fuego in the southernmost region of Chile) about the environment and legal issues so that citizens can
where migratory flamingos resided in winte. At the age decide how to act and what to do to contribute to the care
of 14 I began to study and disseminate scientific informa- of nature, especially wetlands.
tion about the Laguna de Los Cisnes wetland located in For the communication objective, a series of initiatives
the same region. By then I knew that I wanted to become have been developed that promote the care of wetlands
a biologist and dedicate my life to wetland conservation. I from different organizations, with a citizen vision. The
stopped worrying about wetlands and to take care of them, empowerment and identification of the community with
working on wetland conservation initiatives, highlight- wetlands and their biodiversity are fundamental to lo-
ing their key role in the conservation of species and their cal wetland protection and make it to highlight projects
importance to people. that have been developed in the region of Los Ríos Chile,
Somos_humedales is a personal project reflected especially in cities such as Futrono and Los Lagos with a
through social networks (currently via Facebook and Insta- powerful educational focus.
gram) designedto provide general knowledge and technical “Somos_humedales” also actively participates in the
information on wetlands in open access forum (free access) National Network of Wetlands of Chile. Although this
for everyone in a simple language (easy to understand). As initiative is born from a personal concern and professional
such communication and education about wetlands are the interest, it would not be possible without cooperation from
main goals of my initiative. The three objectives of this people who love wetlands and who have the spirit and
project are: 1) bring the community closer (make aware) to conviction support wetland conservation. Collectively, we
projects that are being developed for the care and protection are part of nature “We are wetlands” (“Somos humedales”).
of wetlands, as well as provide news from reliable sources My hope is that, in the near future, “Somos humedales”
about conservation, 2) build a connection space (story) will become an organization to promote wetland conserva-
of experiences from different territories, with the aim of tion in Chile. n
1 Author contact: karinapaz.medioambiente@gmail.com

Urban Wetlands Interactive Platform
Camila Teutsch Barros1
any countries in South America and throughout the frame settings. Users can also download the map as a png
M world still lack the financial and human resources to image, and export data as kml or GeoJSON files for further
deliver sound baselines and implement monitoring programs processing through GIS software.
to support wetland conservation and management. This ex- Another key feature of the platform is the ‘Theme Map’
plains why many wetlands are not even listed within official section. Theme Maps work on the same logic as the Gen-
information sources around the world, which makes them eral Map, but also include the option of drawing lines and
particularly vulnerable, especially in contexts of rapid urban polygons, and are entirely administered by a particular user
growth. This reality is what inspired the development of the or group of users to serve a particular purpose.
Urban Wetlands Interactive Platform (Figure 1). The Urban Last but not least, the Platform includes a ‘Campaigns’
Wetlands Interactive Platform is an online participatory map- section, which are based on Theme Maps that are made
ping tool that helps gather local information about wetlands public for a certain period of time so as to involve the com-
and other natural assets world-wide, in order to support munity in participatory processes, citizen science projects
water-sensitive urban planning and decision-making. or such. Active Campaigns are showcased in the Platform’s
The Platform is not only a cost-effective tool to identify homepage, and Campaign leaders get a Communications
wetlands and gather information about them, but also to Kit which include graphics to help them spread the word
dynamically build on the local knowledge about them and through social networks.
to share ideas for their protection, restoration and sustain- The Platform was developed by Patagua in collabora-
able management. In addition to this, the Platform helps to tion with independent researcher Francisco Vasquez. So far,
connect people and organizations who share an interest on it is being successfully used in Chile and Colombia. It is
wetlands. It fosters collaboration among them and engages currently available in Spanish only, but there should be an
local communities in environmental protection. English version coming out in the short term. Learn more
The main feature of the Platform is a ‘General Map’ to by visiting www.humedalesurbanos.com or contact the
which registered users can add data points on five different developers at hola@humedalesurbanos.com. n
themes (each of which is repre-
sented by a color): 1) Initiatives, FIGURE 1. Themes that can be displayed on the interactive platform.
2) Problems, 3) Public Use, 4)
Historical Data, and 5) Location.
Each point allows users to add
a short description as well as a
picture. All the information in
the General Map is public and
free for everyone to explore and
comment on.
The Platform was developed
as an intuitive tool that allows
everyone to contribute, no ad-
vanced technical skills required.
Both street map and satellite
view are available as basemaps,
and registered users can also
save their preferred zoom and
1 Executive Director, Patagua; camila@patagua.cl.

WETLANDS IN THE NEWS
L isted below are some links to some random news articles that may be of interest. Links from past issues can be ac-
cessed on the SWS website: http://sws.org/Sample-Content/wetlands-in-the-news.html. 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. n
Huntington Beach wetlands continue to expand, following decades of Brazil revokes mangrove protections, weakening another ecosystem key
degradation to curbing climate change
https://www.ocregister.com/2020/10/16/huntington-beach-wetlands- https://www.cnn.com/2020/09/29/americas/brazil-revokes-mangroves-
continue-to-expand-following-decades-of-degradation/ protection-climate-intl/index.html
The World’s Largest Tropical Wetland Has Become an Inferno Invasive sea lampreys in Great Lakes, and the lake trout they prey on,
https://www.nytimes.com/interactive/2020/10/13/climate/pantanal- puzzle scientists
brazil-fires.html https://phys.org/news/2020-09-invasive-sea-lampreys-great-lakes.html
Why Scientists Made Venus Flytraps That Glow USDA Awards $5 Million to Support Wetland Mitigation Banking
https://www.nytimes.com/2020/10/12/science/venus-flytraps-close.html https://www.nrcs.usda.gov/wps/portal/nrcs/detail/national/newsroom/
releases/?cid=NRCSEPRD1660022
To protect nature, bring down the walls of fortress conservation
https://www.aljazeera.com/opinions/2020/10/12/to-protect-nature-bring- Blue Origin to fill wetlands for rocket test site
down-the-walls-of-fortress-conservation/ https://www.floridatoday.com/story/news/local/environment/2020/09/29/
blue-origin-fill-wetlands-rocket-test-site/3559508001/
Florida seeks to take over federal wetland permits
https://www.naplesnews.com/story/news/environment/2020/10/12/ Why Nova Scotia wants to poison a lake to kill off invasive species
florida-seeks-take-over-federal-wetland-permits/3627492001/ https://www.cbc.ca/news/canada/nova-scotia/nova-scotia-lake-poison-
invasive-species-1.5739446
Why trout need wetlands
https://www.wisconsinwetlands.org/updates/why-trout-need-wetlands/ A Watershed Study for Wetland Restoration
https://www.newswise.com/articles/a-watershed-study-for-wetland-
What’s Green, Soggy and Fights Climate Change?
restoration
https://www.nytimes.com/2020/10/09/climate/peat-climate-change.html
Study finds spreading ghost forests on NC coast may contribute to
Droughts are threatening global wetlands
climate change
https://phys.org/news/2020-10-droughts-threatening-global-wetlands.html
https://phys.org/news/2020-09-ghost-forests-nc-coast-contribute.html
How to reverse global wildlife declines by 2050
The World's Marvellously Freaky Carnivorous Plants Are in More
https://theconversation.com/how-to-reverse-global-wildlife-declines-
Trouble Than We Knew
by-2050-146041
https://www.sciencealert.com/the-world-s-marvellously-freaky-carnivo-
Elusive eastern black rail threatened by rising sea levels rous-plants-are-in-dire-need-of-our-help
https://apnews.com/article/habitat-destruction-wildlife-climate-change-
Two new species of wetland plant discovered from Western Ghats
rising-sea-levels-climate-5a8ea861445582c2625d93ba82069c70
https://indianexpress.com/article/india/two-new-species-of-wetland-
Mercury on the Rise plant-discovered-from-western-ghats-6619464/
https://wrri.ncsu.edu/blog/2019/11/mercury-on-the-rise/
Wetland Silviculture & Water Tables
40 Percent of the Amazon Is on the Brink of Becoming Savanna https://www.srs.fs.usda.gov/compass/2020/01/30/wetland-silviculture-
https://earther.gizmodo.com/40-percent-of-the-amazon-is-on-the-brink- water-tables/
of-transitioni-1845274803
Florida Gulf Coast University unveils new ultrasonic technology to fight
USDA Seeks New Partnerships to Safeguard, Restore Wetland Ecosys- algae blooms
tems https://www.fox4now.com/news/protecting-paradise/florida-gulf-coast-
https://www.nrcs.usda.gov/wps/portal/nrcs/detail/national/newsroom/ university-unveils-new-ultrasonic-technology-to-fight-algae-blooms
releases/?cid=NRCSEPRD1664617
Extraordinary Fires Char the Pantanal, a Vast Floodplain in South America
How to Revolutionize Biodiversity Conservation in the U.S. https://scitechdaily.com/extraordinary-fires-char-the-pantanal-a-vast-
https://www.scientificamerican.com/article/how-to-revolutionize-biodi- floodplain-in-south-america/
versity-conservation-in-the-u-s/
Brazilian wetlands fires started by humans and worsened by drought
MR-GO closure improving environment, but more wetlands, swamp https://www.theguardian.com/world/2020/sep/18/brazilian-wetlands-
restoration needed fires-started-by-humans-and-worsened-by-drought
https://www.nola.com/news/environment/article_06d67378-0423-11eb-
Army Corps launches Everglades, Biscayne Bay restoration plan
bcbf-5f2aad29d796.html
https://www.miamiherald.com/news/local/environment/arti-
40 Percent of World's Plants at Risk of Extinction cle245828505.html
https://www.ecowatch.com/plants-biodiversity-extinction-2647868027.html
New wetland recharge park opens in Ocala
New Study Shows a Vicious Circle of Climate Change Building on https://www.wcjb.com/2020/09/17/new-wetland-recharge-park-opens-
Thickening Layers of Warm Ocean Water in-ocala/
https://insideclimatenews.org/news/28092020/ocean-stratification-
‘The warning lights are flashing.’ Report finds nations failing to protect
climate-change
biodiversity
The world’s largest wetland is on fire: how can we save the Pantanal? https://www.sciencemag.org/news/2020/09/warning-lights-are-flashing-
https://blog.ecosia.org/what-is-the-pantanal-and-why-is-it-burning/ report-finds-nations-failing-protect-biodiversity
'The Blob': Low-oxygen water killing lobsters, fish in Cape Cod Bay Handing federal wetlands permitting to FL DEP is an idea that’s all wet
https://www.capecodtimes.com/news/20200928/the-blob-low-oxygen- https://www.floridaphoenix.com/2020/09/17/handing-federal-wetlands-
water-killing-lobsters-fish-in-cape-cod-bay permitting-to-fl-dep-is-an-idea-thats-all-wet/

Fires in Pantanal, world's largest tropical wetlands, 'triple' in 2020 Pitkin County launches project to restore ancient wetland at North Star
https://www.bbc.com/news/world-latin-america-53500288 Preserve near Aspen
https://www.9news.com/article/life/style/colorado-guide/pitkin-
Portugal's airport plans threaten wetlands
county-project-restore-ancient-wetland-north-star-preserve-
https://science.sciencemag.org/content/369/6510/1440.1
aspen/73-78622b09-815a-48ff-938f-999d67771d77
Australian stinging trees release spider-like venom that can hurt for weeks
New floating wetlands in Winton Woods pond hold aquatic plants
https://www.cnn.com/2020/09/17/asia/australia-venomous-stinging-
https://www.cincinnati.com/story/news/2020/09/05/new-floating-wet-
trees-scn-scli-intl/index.html
lands-winton-woods-pond-hold-aquatic-plants/5730368002/
Soggy coastal soils? Here’s why ecologists love them
Invasive green crab species spotted in Padilla Bay, Anacortes
https://www.sciencenewsforstudents.org/article/wetlands-coastal-soils-
https://mynorthwest.com/2139258/invasive-green-crabs-padilla-bay/
ecology-flooding-storm-surge-climate-change
Invasive aquatic plant found in 4 Michigan inland lakes
Long-term change in habitat and vegetation in an ungrazed, estuarine salt
https://www.wnem.com/news/invasive-aquatic-plant-found-in-4-michi-
marsh: Man-made foreland compared to young marsh development
gan-inland-lakes/article_9aa160ae-ef97-11ea-978a-3381cd594084.html
https://www.sciencedirect.com/science/article/abs/pii/
S0272771419302537 Brazil Pantanal Scorched by Fires
https://www.nytimes.com/2020/09/04/world/americas/brazil-wetlands-
Conservationists Split Over Poseidon Desal Project’s Potential to Help
fires-pantanal.html
Bolsa Chica Wetlands Voice of OC
https://voiceofoc.org/2020/09/conservationists-split-over-poseidon- Court Overturns Administration Efforts to Weaken the Migratory Bird
desal-projects-potential-to-help-bolsa-chica-wetlands/ Treaty Act
https://abcbirds.org/article/court-upholds-mbta-2020/
Birds are mysteriously dying in New Mexico in 'frightening' numbers
https://www.lcsun-news.com/story/news/2020/09/12/mass-deaths-migra- Quakertown swamp gets Wetland of Distinction title | Southeastern
tory-birds-new-mexico-environment/5780282002/ Pennsylvania
https://www.wfmz.com/news/area/southeastern-pa/quakertown-swamp-
Battle on to save Brazil's tropical wetlands from flames
gets-wetland-of-distinction-title/article_f6c60f74-e7ea-11ea-838d-
https://apnews.com/35368749c3619220d79a2ecfa9ab1e3b
4707c0188c10.html
Trump Administration Announces more than $130 Million in Public-
NASA Research Reveals the True Causes of Sea Level Rise Since 1900
Private Funding for Wetland Conservation Projects
https://scitechdaily.com/nasa-research-reveals-the-true-causes-of-sea-
https://www.mychesco.com/a/news/national/trump-administration-an-
level-rise-since-1900/
nounces-more-than-130-million-in-public-private-funding-for-wetland-
conservation-projects/ Human-Nature Connectivity: Wetlands Within Sustainable Futures
https://link.springer.com/chapter/10.1007/978-3-030-41306-4_6
Buzzy’s Ranch wetlands restoration seeks grant to buy water rights
https://sierranevadaally.org/2020/09/09/buzzys-ranch-wetlands-restora- Hamoun wetlands restoration project started
tion-looks-for-grant-to-buy-water-rights/ https://www.tehrantimes.com/news/451584/Hamoun-wetlands-restora-
tion-project-started
World's wildlife populations in devastating decline warns WWF report
https://www.cnn.com/2020/09/09/world/wwf-report-species-decline- Inverleith Park Pond - Water Gems
climate-scn-intl-scli/index.html https://www.watergems.co.uk/portfolio_page/inverleith-park-pond/
More than half of government environmental scientists say their work Massachusetts grants $798,500 for river, wetland restoration projects
has been suppressed https://www.masslive.com/news/2020/08/massachusetts-grants-
https://www.abc.net.au/news/science/2020-09-09/environment-scientists- 798500-in-river-wetland-restoration-projects.html
censored-suppressed-data/12643824 BCCC & Sound Rivers to start construction on new wetlands
Wildlife in 'catastrophic decline' due to human destruction, scientists warn https://www.witn.com/2020/08/22/bccc-sound-rivers-to-start-construc-
https://www.bbc.com/news/science-environment-54091048 tion-on-new-wetlands/
Bending the Curve of Biodiversity Loss: Ambitious Conservation and Potsdam professor turns to beetles in fight against invasive purple loosetrife
Restoration Efforts Required https://www.newyorkupstate.com/adirondacks/2020/08/potsdam-profes-
https://scitechdaily.com/bending-the-curve-of-biodiversity-loss-ambi- sor-turns-to-beetles-in-fight-against-invasive-purple-loosetrife.html
tious-conservation-and-restoration-efforts-required/ VEWH - Improving plants in Lower Barwon wetlands
As the Arctic thaws, Indigenous Alaskans demand a voice in climate https://www.vewh.vic.gov.au/news-and-publications/stories/improving-
change research plants-in-lower-barwon-wetlands
https://www.sciencemag.org/news/2020/09/arctic-thaws-indigenous- Brazil fires threaten world's largest wetland
alaskans-demand-voice-climate-change-research https://www.reuters.com/article/us-brazil-environment-fires-idUSKC-
How artificial salt marshes can help in the fight against rising seas N25E2FC
https://www.theguardian.com/environment/2020/sep/09/how-artificial- Watch hummingbirds ‘dance’ through waterfalls
salt-marshes-can-help-in-the-fight-against-rising-seas-aoe https://www.sciencemag.org/news/2020/08/watch-hummingbirds-dance-
With Baylands under flood threat, Palo Alto explores projects to address through-waterfalls
sea level rise The sea otter rescue plan that worked too well
https://www.paloaltoonline.com/news/2020/09/07/with-baylands-under- https://www.bbc.com/future/article/20200818-the-canadian-sea-otter-
flood-threat-palo-alto-explores-projects-to-address-sea-level-rise rescue-plan-that-worked-too-well
Why plants in wetlands are highly productive Carnivorous Plants of Delaware's Wetlands: sundew, bladderworts & more
https://phys.org/news/2020-09-wetlands-highly-productive.html https://wmap.blogs.delaware.gov/2020/03/05/wetland-carnivorous-
Wetland area does not get Greene County stamp of approval, land trust plants-nothing-to-be-afraid-of/
makes plans for another proposal Under the radar rollback of stream and wetland protections
https://www.daytondailynews.com/news/wetland-area-does-not-get- https://thehill.com/opinion/energy-environment/512499-under-the-radar-
greene-county-stamp-of-approval-land-trust-makes-plans-for-another- rollback-of-stream-and-wetland-protections
proposal/HV5ON7B34ZCOBN5GVCXRTCPUMI/

Greenland's ice sheet has melted to a point of no return, study finds Fridays On The Farm: Restoring Wetlands and Creating Habitat
https://www.cnn.com/2020/08/14/weather/greenland-ice-sheet/index.html https://www.farmers.gov/connect/blog/conservation/fridays-farm-restor-
ing-wetlands-and-creating-habitat
Concrete to be removed to create wetlands in East Bay
https://www.eastbaytimes.com/old-aircraft-taxiway-in-alameda-to-be- Climate change: UK peat emissions could cancel forest benefits
converted-to-wetlands-park https://www.bbc.com/news/science-environment-53684047
'It overtakes everything:' Invasive aquatic plant threatens to overrun Wind farms built on carbon-rich peat bogs lose their ability to fight
Potomac watershed climate change
https://wjla.com/news/local/invasive-plant-species-potomac-watershed https://theconversation.com/wind-farms-built-on-carbon-rich-peat-bogs-
lose-their-ability-to-fight-climate-change-143551
Scientists decry federal rule that removes protection from 'unconnected'
streams and wetlands Articles on international activities in wetland and nature conservation
https://phys.org/news/2020-08-scientists-decry-federal-unconnected- https://news.mongabay.com/author/genevieve/
streams.html New wetlands open to the public
High-latitude Climate Change - Alaska Nature and Science (U.S. Na- https://www.dailystandard.com/archive/2020-08-03/stories/41000/new-
tional Park Service) wetlands-open-to-the-public
https://www.nps.gov/subjects/aknatureandscience/hi-latclimatechange.htm Peat takes millennia to generate, and bogs store 10 times more carbon
Dead sturgeon found on Lake Michigan beaches than forests — using it in gardening is madness
https://www.michiganradio.org/post/dead-sturgeon-found-lake-michi- https://www.countrylife.co.uk/nature/peat-takes-millennia-to-generate-
gan-beaches and-bogs-store-10-times-more-carbon-than-forests-using-it-in-garden-
ing-is-madness-214851
We mapped the world's frozen peatlands – what we found was very
worrying Ancient bones in disturbed peat bogs are rotting away, alarming
https://theconversation.com/we-mapped-the-worlds-frozen-peatlands- archaeologists
what-we-found-was-very-worrying-144235 https://www.sciencemag.org/news/2020/07/ancient-bones-disturbed-
peat-bogs-are-rotting-away-alarming-archaeologists
Half Of Wisconsin’s Wetlands To Lose Federal Protection, Nature Con-
servancy Estimates Smaller habitats worse than expected for biodiversity
https://www.wxpr.org/post/half-wisconsin-s-wetlands-lose-federal-pro- https://phys.org/news/2020-07-smaller-habitats-worse-biodiversity.html
tection-nature-conservancy-estimates#stream/0 Kabartal Wetland
A century ago, it was a Florida landscape shrub. Now, it's a pest plant https://definearth.com/2020/04/05/kabartal-wetland/
that keeps spreading. 'A win for the Everglades': 5,000 pythons removed in state-sponsored
https://phys.org/news/2020-08-century-florida-landscape-shrub-pest.html capture program
UNEP supports project to restore peatlands in Indonesia https://www.msn.com/en-us/news/us/a-win-for-the-everglades-
https://www.unenvironment.org/news-and-stories/story/unep-supports- 5-000-pythons-removed-in-state-sponsored-capture-program/ar-
project-restore-peatlands-indonesia BB17lBQM?li=BBnbcA1
Wetland revival Wetland Wildlife of Arkansas
https://www.iowafarmbureau.com/Article/Wetland-revival http://www.naturalheritage.com/blog/wetland-wildlife
Market St. development on wetland overcomes opposition, heads to Ireland's Peatlands - The National Trust for Ireland
Wilmington council https://www.antaisce.org/issues/irelands-peatlands
https://portcitydaily.com/local-news/2020/08/10/market-st-development- New wetland installed in Putnam County
on-wetland-overcomes-opposition-heads-to-wilmington-council/ https://www.limaohio.com/news/419468/new-wetland-installed-in-
Citrus Flavoring Is Weaponized Against Insect-Borne Diseases putnam-county
https://www.nytimes.com/2020/08/10/health/tick-mosquito-repellant- Two new bird species discovered nesting in restored Arkansas wetlands
nootkatone.html https://www.kark.com/news/local-news/two-new-bird-species-discov-
Abandoned Fiberglass Boats are Polluting the Marine Environment ered-nesting-in-restored-arkansas-wetlands/
https://www.maritime-executive.com/editorials/abandoned-fiberglass- Plan to dissolve Peatland Restoration Agency raised concerns
boats-are-polluting-the-marine-environment https://www.thejakartapost.com/news/2020/07/16/plan-to-dissolve-peat-
Toxic Chemicals From Fossil Fuels Are Poisoning East Coast Dolphins land-restoration-agency-raised-concerns.html
and Whales, Study Finds Global methane emissions soar to record high
https://www.ecowatch.com/toxic-chemicals-dolphins-whales- https://phys.org/news/2020-07-global-methane-emissions-soar-high.html
us-2646906589.html
Plants cropping up in lost Michigan lakes where dams failed
https://news.yahoo.com/plants-cropping-lost-michigan-
lakes-130604416.html

WETLAND BOOKSHELF
oug Wilcox, former editor of Wetlands, has edited and Although the list of participating scientists is huge (71
D produced a book that provides a review of the history
of wetland science from the experiences of today’s wetland
contributors) and contains the names of many familiar wet-
land scientists, it is not exhaustive as others were unable to
scientists. The book entitled “History of Wetland Science: contribute for various reasons. Nonetheless, the book of-
A Perspective from Wetland Leaders” was published in fers a unique frame of reference for presenting the history
July 2020 and is available from Amazon (https://www. of wetland science by having today’s wetland scientists
amazon.com/History-Wetland-Science-Perspectives-Lead- provide short stories of their introduction to wetlands and a
ers/dp/B08DC6GXDM). In the Preface, Doug reveals reflection on their careers.
that his inspiration for the book came from the fact that For the latest news on wetlands and related topics,
many of his old wetland friends were retiring, retired and readers are referred to the Association of State Wetland
a few had passed and he wanted to record some of their Managers website: https://www.aswm.org/. Their “Wetland
experiences. He felt it would be a meaningful exercise to News Digest” section include links to newspaper articles
have wetland scientists who have made significant contri- that should be of interest: https://www.aswm.org/news/
butions to wetland science, management, or conservation wetland-breaking-news.
provide short autobiographies that would offer presentday Please help us add new books to this listing. If your
and future students of wetland science some perspective on agency, organization, or institution has a website where wet-
how early wetland scientists became involved in wetlands land information can be accessed, please send the informa-
and the challenges they faced in their careers. Students as tion to the Editor of Wetland Science & Practice at ralphtin-
well as today’s wetland scientists should find the chapters er83@gmail. com. Your cooperation is appreciated. n
interesting, inspiring, and sometimes humorous as contrib-
utors offer a personal glimpse of their life with wetlands.
BOOKS
• History of Wetland Science: A Perspective from Wetland • Wetland Soils: Genesis, Hydrology, Landscapes, and Clas-
Leaders https://www.amazon.com/History-Wetland-Sci- sification https://www.crcpress.com/Wetland-Soils-Gene-
ence-Perspectives-Leaders/dp/B08DC6GXDM sis-Hydrology-Landscapes-and-Classification/Vepraskas-
• An Introduction to the Aquatic Insects of North America Richardson-Vepraskas-Craft/9781566704847
(5th Edition) https://he.kendallhunt.com/product/introduc- • Creating and Restoring Wetlands: From Theory to Practice
tion-aquatic-insects-north-america http://store.elsevier.com/Creating-and-Restoring-Wetlands/
• Wading Right In: Discovering the Nature of Wetlands
Christopher-Craft/isbn-9780124072329/
https://press.uchicago.edu/ucp/books/book/chicago/W/
• Salt Marsh Secrets. Who uncovered them and how?
bo28183520.html
http://trnerr.org/SaltMarshSecrets/
• Sedges of Maine
• Remote Sensing of Wetlands: Applications and Advances.
https://umaine.edu/umpress/books-in-print/
https://www.crcpress.com/product/isbn/9781482237351
• Sedges and Rushes of Minnesota https://www.upress.umn.
• Wetlands (5th Edition). http://www.wiley.com/WileyCDA/
edu/book-division/books/sedges-and-rushes-of-minnesota
WileyTitle/productCd-1118676823.html
• Wetland & Stream Rapid Assessments: Development,
• Black Swan Lake – Life of a Wetland http://press.uchicago.
Validation, and Application https://www.elsevier.com/
edu/ucp/books/book/distributed/B/bo15564698.html
books/wetland-and-stream-rapid-assessments/dor-
• Coastal Wetlands of the World: Geology, Ecology, Dis-
ney/978-0-12-805091-0
tribution and Applications http://www.cambridge.org/
• Eager: The Surprising Secret Life of Beavers and Why
us/academic/subjects/earth-and-environmental-science/
They Matter https://www.chelseagreen.com/product/eager/
environmental-science/coastal-wetlands-world-geology-
• Wetland Indicators – A Guide to Wetland Formation, Iden-
ecology-distribution-and-applications
tification, Delineation, Classification, and Mapping
• Florida’s Wetlands https://www.amazon.com/Floridas-
https://www.crcpress.com/Wetland-Indicators-A-Guide-to-
Wetlands-Natural-Ecosystems-Species/dp/1561646873/
Wetland-Identification-Delineation-Classification/Tiner/p/
ref=sr_1_4?ie=UTF8&qid=1518650552&sr=8-
book/9781439853696
4&keywords=wetland+books

ONLINE SOURCES OF WETLAND INFORMATION
• Mid-Atlantic Freshwater Wetlands: Science, Management, The following is a listing of some government agencies and
Policy, and Practice http://www.springer.com/environment/ environmental organizations that provide online informa-
aquatic+sciences/book/978-1-4614-5595-0 tion on wetlands and where their publications on wetlands
• The Atchafalaya River Basin: History and Ecology of an may be accessed.
American Wetland http://www.tamupress.com/product/ • U.S. Army Corps of Engineers, Waterways Experiment
Atchafalaya-River-Basin,7733.aspx Station, Environmental Laboratory https://www.erdc.
• Tidal Wetlands Primer: An Introduction to their Ecology, usace.army.mil/Locations/EL.aspx
Natural History, Status and Conservation https://www. • U.S. Army Corps of Engineers, National Wetland Plants
umass.edu/umpress/title/tidal-wetlands-primer Database http://wetland-plants.usace.army.mil/nwpl_stat-
• Wetland Landscape Characterization: Practical Tools, ic/v34/home/home.html
Methods, and Approaches for Landscape Ecology http:// • U.S. Environmental Protection Agency https://www.epa.
www.crcpress.com/product/isbn/9781466503762 gov/wetlands
• Wetland Techniques (3 volumes) http://www.springer.com/ • U.S. Fish and Wildlife Service, National Wetlands Inventory
life+sciences/ecology/book/978-94-007-6859-8 https://fws.gov/wetlands/
• Wildflowers and Other Plants of Iowa Wetlands • U.S. Geological Survey, Wetland and Aquatic Research
https://www.uipress.uiowa.edu/books/2015-spring/wild- Center https://www.usgs.gov/centers/wetland-and-aquatic-
flowers-and-other-plants-iowa-wetlands.htm research-center-warc
• Wetland Restoration: A Handbook for New Zealand • U.S. Geological Survey, Northern Prairie Wildlife Re-
Freshwater Systems https://www.landcareresearch.co.nz/ search Center https://www.usgs.gov/centers/npwrc
publications/books/wetlands-handbook • U.S. Geological Survey, Patuxent Wildlife Research Center
• Wetland Ecosystems https://www.wiley.com/en-us/ https://www.usgs.gov/centers/pwrc
Wetland+Ecosystems-p-9780470286302 • National Oceanic and Atmospheric Administration, Office
• Constructed Wetlands and Sustainable Development of Coastal Management https://coast.noaa.gov/
https://www.routledge.com/Constructed-Wet- • U.S.D.A. Natural Resources Conservation Service, Hydric
lands-and-Sustainable-Development/Austin-Yu/p/ Soils https://www.nrcs.usda.gov/wps/portal/nrcs/main/
book/9781138908994 soils/use/hydric/
• Association of State Wetland Managers https://www.
aswm.org/

WETLANDS JOURNAL
What’s New in the SWS Journal - Wetlands?

The following articles appear in Volume 40, Issue 4 of Wetlands, Journal of the Society of Wetland Scientists.
• The Twin Limit Marsh Model: A Non-equilibrium Approach to Predicting Marsh Vegetation on Shorelines and in
Floodplains
• Denitrification Capacity of Hill Country Wet and Dry Area Soils as Influenced by Dissolved Organic Carbon Con-
centration and Chemistry
• Dissolved Organic Matter Export from Surface Sediments of a New England Salt Marsh
• Nutrient Inputs and Hydrology Interact with Plant Functional Type in Affecting Plant Production and Nutrient Con-
tents in a Wet Grassland
• Patterns of Denitrification and Methanogenesis Rates from Vernal Pools in a Temperate Forest Driven by Seasonal,
Microbial Functional Gene Abundances, and Soil Chemistry
• The Difference in Light use Efficiency between an Abandoned Peatland Pasture and an Adjacent Boreal Bog in
Western Newfoundland, Canada
• A Suitable Method for Assessing Invasibility of Habitats in the Ramsar Sites - an Example of the Southern Part of
the Pannonian Plain
• Multiple Stressors Influence Salt Marsh Recovery after a Spring Fire at Mugu Lagoon, CA
• Impact of Barrier Breaching on Wetland Ecosystems under the Influence of Storm Surge, Sea-Level Rise and Fresh-
water Discharge
• Feedbacks of Alpine Wetlands on the Tibetan Plateau to the Atmosphere
• Coastal Marsh Bird Habitat Selection and Responses to Hurricane Sandy
• Do Breeding Bird Communities or Conservation Value Differ Among Forested Wetland Types or Ecoregions in
Nova Scotia?
• The Missing Metric: An Evaluation of Fungal Importance in Wetland Assessments
• Foundation Species Loss Affects Leaf Breakdown and Aquatic Invertebrate Resource Use in Black Ash Wetlands
• Habitat Suitability Modelling of Benthic Macroinvertebrate Community in Wetlands of Lake Tana Watershed,
Northwest Ethiopia
• Impact of Peatland Restoration on Soil Microbial Activity and Nematode Communities
• Plant Community Establishment in a Coastal Marsh Restored Using Sediment Additions
• A Framework for Considering Climate Change Impacts in Project Selection for Deepwater Horizon Restoration Ef-
forts
• In-Situ CO2 Partitioning Measurements in a Phragmites australis Wetland: Under-
standing Carbon Loss through Ecosystem Respiration
• Correction to: Morphology of Drained Upland Depressions
on the Des Moines Lobe of Iowa
• Correction to: “Relationships Between Salinity and Short-
Term Soil Carbon Accumulation Rates from Marsh Types
Across a Landscape in the Mississippi River Delta"

WSP SUBMISSION GUIDELINES
About Wetland Science & Practice (WSP)
etland Science and Practice (WSP) is the

W SWS quarterly publication aimed at provid-
ing information on select SWS activities (technical
WSP Manuscript – General Guidelines
LENGTH:
committee summaries, chapter workshop overview/ Approximately 5,000 words; can be longer if necessary.
abstracts, and SWS-funded student activities), brief
summary articles on ongoing or recently completed STYLE:
wetland research, restoration, or management See existing articles from 2014 to more recent years available
projects or on the general ecology and natural his- online at:
tory of wetlands, and highlights of current events. http://www.sws.org/Publications/wsp-contents.html
WSP also includes sections listing new publications
and research at various institutions, and links to TEXT:
major wetland research facilities, federal agencies, Word document, 12 font, Times New Roman, single-spaced; keep
wetland restoration/monitoring sites and wetland tables and figures separate, although captions can be included in
mapping sites. The publication also serves as an text. For reference citations in text use this format: (Smith 2016;
outlet for commentaries, perspectives and opinions Jones and Whithead 2014; Peterson et al. 2010).
on important developments in wetland science,
FIGURES:
theory, management and policy.
Please include color images and photos of subject wetland(s) as
Both invited and unsolicited manuscripts are
WSP is a full-color e-publication.
reviewed by the WSP editor for suitability for pub-
Image size should be less than 1MB – 500KB may work best for
lication. Student papers are welcomed. Please see
this e-publication.
publication guidelines at the end of this issue.
Electronic access to Wetland Science and REFERENCE CITATION EXAMPLES:
Practice is included in your SWS membership. • Claus, S., S. Imgraben, K. Brennan, A. Carthey, B. Daly, R.
All issues published, except the the current issue, Blakey, E. Turak, and N. Saintilan. 2011. Assessing the ex-
are available via the internet to the general public. tent and condition of wetlands in NSW: Supporting report
At the San Juan meeting, the SWS Board of A – Conceptual framework, Monitoring, evaluation and re-
Directors voted to approve release of past issues porting program, Technical report series, Office of Environ-
of WSP when a new issue is available to SWS ment and Heritage, Sydney, Australia. OEH 2011/0727.
members only. This means that a WSP issue will • Clements, F.E. 1916. Plant Succession: An Analysis of the
be available to the public four months after it has Development of Vegetation. Carnegie Institution of Wash-
been read by SWS members (e.g., the June 2017 ington. Washington D.C. Publication 242.
issue will be an open access issue in September • Clewell, A.F., C. Raymond, C.L. Coultas, W.M. Dennis,
2017). Such availability will hopefully stimulate and J.P. Kelly. 2009. Spatially narrow wet prairies. Casta-
more interest in contributing to the journal. And, nea 74: 146-159.
we are excited about this opportunity to promote
• Colburn, E.A. 2004. Vernal Pools: Natural History and
the good work done by our members.
Conservation. McDonald & Woodward Publishing Com-
HOW YOU CAN HELP pany, Blacksburg, VA.
If you read something you like in WSP, or that you • Cole, C.A. and R.P. Brooks. 2000. Patterns of wetland
think someone else would find interesting, be sure hydrology in the Ridge and Valley Province, Pennsylvania,
to share. Share links to your Facebook, Twitter, USA. Wetlands 20: 438-447.
Instagram and LinkedIn accounts. • Cook, E.R., R. Seager, M.A. Cane, and D.W. Stahle. 2007.
Make sure that all your SWS colleagues are North American drought: reconstructions, causes, and con-
checking out our recent issues, and help spread the sequences. Earth-Science Reviews 81: 93-134.
word about SWS to non-members! • Cooper, D.J. and D.M. Merritt. 2012. Assessing the water
Questions? Contact editor Ralph Tiner, PWS needs of riparian and wetland vegetation in the western United
Emeritus (ralphtiner83@gmail.com). n States. U.S.D.A., Forest Service, Rocky Mountain Research
Station, Ft. Collins, CO. Gen. Tech. Rep. RMRS-GTR-282.

WEB TIP
Resources
at your fingertips!
For your convenience, SWS has compiled a hefty list
of wetland science websites, books, newsletters,
government agencies, research centers and more,
and saved them to sws.org.
Find them on the Related Links page sws.org.
Coming in 2021!
ADVERTISING OPPORTUNITIES IN WSP
Stay tuned for more information to be released soon!
Wetland Science
&
WSP is the formal voice of the Society of Wetland Scientists. It is a quarterly publication focusing
on the news of the SWS and providing important announcements for members and opportunities
for wetland scientists, managers, and graduate students to publish brief summaries of their works
Practice and conservation initiatives. Topics for articles may include descriptions of threatened wetlands
around the globe or the establishment of wetland conservation areas, and summary findings from
research or restoration projects. All manuscripts should follow guidelines for authors listed above. All papers published in WSP will be reviewed by the
editor for suitability and may be subject to peer review as necessary. Most articles will be published within 3 months of receipt. Letters to the editor are
also encouraged, but must be relevant to broad wetland-related topics. All material should be sent electronically to the current editor of WSP. Com-
plaints about SWS policy or personnel should be sent directly to the elected officers of SWS and will not be considered for publication in WSP. n
SOCIETY OF WETLAND SCIENTISTS

1818 Parmenter St., Ste 300,
Middleton, WI 53562
(608) 310-7855
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October 2020 Wetland Science &amp; Practice

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Wetland Science

SWS - Celebrating Our 40th Anniversary in 2020

SOCIETY OF WETLAND SCIENTISTS

REFLECTIONS, CONNECTIONS AND THE CHANGING SEASONS

I’ve been watching intently and enjoying the seasons change

216 Wetland Science & Practice October 2020

WORLD’S LEADING AQUATIC SCIENTIFIC SOCIETIES PUBLIC POLICY UPDATES

218 Wetland Science & Practice October 2020

Introducing the Society of Wetland Scientists 40th Anniversary Award

Student Section Virtual Conference SWS Grants International Travel Awards

Wetland Science & Practice October 2020 219

2020 Virtual Meeting

220 Wetland Science & Practice October 2020

STUDENTS - APPLY FOR SWAMMP PARTICIPATION

More info: http://sws.org/Awards-and-Grants/sws-undergraduate-mentoring-program-swammp.html n

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Passing of a Leader in Wetland Policy Development

From the Editor’s Desk, continued from page 214

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This Issue: Emphasis on Latin America Wetlands

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Ephemeral wetlands, Lençóis Maranhenses, Brazil by Tatiana Lobato de Magalhães.

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Lacustrine marsh, Panguipulli, Los Rios Region, Chile by Karina Arteaga.

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Fire in Paradise Declaration: A Call for Your Participation

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TABLE 2. Principal research areas of the Andean States’ organizations.

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Wetlands of the Coast of Lima: Patterns of Plant Diversity and Challenges

1 Professional Wetland Scientist. Associate Researcher, Universidad Científica del

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Andes, Bofedales, and the Communities of Huascarán National Park, Peru

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October 2020 Wetland Science & Practice

October 2020 Wetland Science & Practice