Coral Reefs of Cuba (Coral Reefs of the World, 18) 3031367189, 9783031367182

Table of contents :
Foreword
Preface
Contents
Part I: Introduction
1: Insights from Cuban Coral Reefs
1.1 Introduction
1.2 Coral Reef Types and Ecological Zonation of the Scleractinians
1.2.1 General Notes
1.2.2 Fringing Reefs
1.2.3 Barrier Reefs
1.2.4 Reefs on Muddy Bottom
1.3 General Review of the Ecology of the Scleractinians of the Cuban Archipelago
1.4 Contemporary Perspective
References
Part II: History
2: Research History of Corals and Coral Reefs in Cuba
2.1 First Traces of Interest in Corals and Coral Reefs
2.2 The Eighteenth Century: The ``Interesting Stones´´ of Don Antonio Parra
2.3 The Nineteenth Century: Beginnings of Marine Sciences in Cuba
2.4 The Twentieth Century: Birth of the Marine Research Centers, Protected Areas, and First Collaboration Agreements
2.5 The Twenty-First Century: First Steps
References
Part III: Description
3: Physical-Geographic Characteristics of Cuban Reefs
3.1 Introduction
3.2 Data and Methodology
3.2.1 Datasets
3.2.2 Outline of Methods
3.2.3 Definition of Mapping Classes
3.2.4 Estimation of Metrics for Cuban Coral Reefs
3.3 Results and Discussion
3.3.1 Description of the Main Regional Coral Reef Groups in Cuba
3.3.2 Estimating the Metrics of Cuban Reefs
3.3.2.1 Accuracy Assessment
3.3.2.2 Comparison with Existing Coral Reef Maps of Cuba
3.3.2.3 Comparison with Other Countries
3.3.3 Barrier Reefs in Cuba
3.4 Hurricane Frequency by Sectors
3.5 Conclusions
3.6 Recommendations
References
4: Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf and Uplifted Marine Terraces of Cuba: Tec...
4.1 Introduction
4.2 General Geology
4.3 Marine and Coastal Geomorphology
4.3.1 Shelf Seaward Margins
4.3.2 The Shelf Interior
4.3.3 Acoustic Basement of the Shelf
4.3.4 Shorelines and Coastal Areas
4.3.5 Emerged Marine Terraces
4.3.6 Rates of Tectonic Uplift of Pleistocene Coral Limestone
4.4 Late Pleistocene to Holocene Evolution of Coral Reef Communities
4.4.1 Stage Correlated to MIS 7 and 6
4.4.2 Coral Reef Expansion During MIS 5
4.4.3 Coral Reef Demise During MIS 2
4.4.4 Sea-Level Rise and Initiation of Modern Reef Development
4.5 Conclusions
References
5: A Remote Sensing Appraisal of the Extent and Geomorphological Diversity of the Coral Reefs of Cuba
5.1 Introduction: The Millennium Coral Reef Mapping Project of Cuba´s Reefs
5.2 Remote Sensing Image Database
5.3 Key Structural Mapping Choices
5.4 Cuba Geomorphological Representation (Level 3 and Level 5)
5.4.1 Jardines de la Reina, Ana Maria Gulf, and Guacanayabo Gulf
5.4.2 Batabano Gulf and Isla de la Juventud
5.4.3 Continental Barrier Reef in the Moa Region
5.5 Areal Coverage Statistics
5.6 Discussion
5.6.1 Geomorphological Diversity of the Coral Reefs of Cuba
5.6.2 Potential for Coral Reef Management Applications
5.6.3 Future Coral Reef Mapping Efforts
References
Part IV: Biota
6: Macrophytes Associated with Cuban Coral Reefs
6.1 Introduction
6.2 Species Richness and Distribution
6.3 Abundance of Macroalgae on Coral Reefs
6.4 Future Issues for Research
Appendix
References
7: Sponges: Conspicuous Inhabitants of the Cuban Coral Reefs and Their Potential as Bioindicators of Contamination
7.1 Introduction
7.2 Biodiversity
7.3 Ecology
7.3.1 Species Composition
7.3.2 Density
7.4 Sponges as Bioindicators of Contamination
7.5 Final Considerations
7.5.1 Deep Revision of Total Cuban Sponge Species Listed
7.5.2 Assessment of Less and Unexplored Cuban Coral Reef
7.5.3 Incorporation of Sponges as Key Benthos Component in Coral Reef Monitoring
7.5.4 Incorporation of Sponge Biodiversity and Ecology on Marine Protected Areas (MPAs) Management Plan
7.5.5 Continue Biomedical Explorations Based on Cuban Marine Sponges
7.5.6 Increase the Number of Researchers Dedicated to Study of Cuban Marine Sponges
References
8: Species List of Cuban Stony Corals: Class Anthozoa, Order Scleractinia; Class Hydrozoa, Suborders Capitata and Filifera
8.1 Introduction
8.2 Materials and Methods
8.3 Revised List
8.4 References for the Identification of Hermatypic Stony Corals of Cuba
8.5 Discussion
References
9: Octocoral Forests: Distribution, Abundance, and Species Richness in Cuban Coral Reefs
9.1 Introduction
9.2 Reports of Octocorals in Cuba
9.3 Richness and Diversity of Octocorals
9.4 Octocoral Density (Colonies/m2)
9.5 Composition of Octocoral Communities
9.6 Conclusions
References
10: Current State of Knowledge of Reef Mollusks in Cuba
10.1 Introduction
10.2 Cuban Reef Malacofauna
10.3 Reef Mollusks, Between Day and Night
References
11: Herbivory on Cuban Coral Reefs
11.1 Introduction
11.2 What´s Known About Coral Reef Herbivorous Invertebrates in Cuba
11.3 Herbivorous Fishes of Cuban Coral Reefs
11.4 Macroalgal Communities of Cuban Coral Reefs and Herbivory
11.5 Case Studies
11.6 Knowledge Gaps and Future Directions in Research
References
12: Chronology of the Lionfish Invasion in Cuba and Evaluation of Impacts on Native Reef Fishes
12.1 Introduction
12.2 Methods
12.2.1 Visual Censuses and Study Sites
12.2.2 Lionfish Impact Evaluations in Cuba
12.3 Results
12.3.1 Database of Lionfish Sightings in Cuba
12.3.2 Lionfish Impact Evaluation in Cuba
12.4 Discussion
12.4.1 Database of Lionfish Sightings in Cuba
12.4.2 Lionfish Impact Evaluation in Cuba
12.4.3 Other Cuban Locations and Habitats Involved on Lionfish Study and Management
References
13: Sharks and Rays in Cuban Coral Reefs: Ecology, Fisheries, and Conservation
13.1 Introduction
13.2 Materials and Methods
13.2.1 Study Areas
13.2.2 Data Collection
13.2.3 Data Analysis
13.3 Results and Discussion
13.3.1 Studies on Sharks and Rays in Cuba
13.3.2 Species Richness, Relation to Coral Reefs, Economical Importance, and Conservation Status of Cuban Reef Sharks and Rays
13.3.3 Biology of Cuban Reef Sharks and Rays
13.3.3.1 Fisheries-Independent Ecological Information
13.3.3.2 Fisheries-Dependent Ecological Information
13.3.4 Movement Ecology and Critical Habitats for Cuban Reef Sharks
13.3.5 Fisheries That Impact Reef Sharks and Rays in Cuba
13.3.6 MPAs and Cuban Reef Sharks and Rays
13.4 Conclusions and Recommendations for Research and Management of Reef Sharks and Rays in Cuba
References
14: Mesophotic Coral Ecosystems of Cuba
14.1 Research History of Mesophotic Coral Ecosystems in Cuba
14.2 Geomorphology and Distribution of MCEs in Cuba
14.3 Oceanography
14.4 MCE Depth Biozonation
14.5 Biodiversity of Cuba´s MCE
14.6 Ecology
14.7 Coral Connectivity
14.8 Biodiversity of Cuba MCES Versus Other Caribbean MCEs
14.9 Changes over Time
14.10 Conservation/Threats
14.11 Future
References
Part V: Ecology, Conservation and Management
15: Status of Cuban Coral Reefs
15.1 Introduction
15.2 Cuban Coral Reefs: Overview
15.3 Coral Reef Studies in Cuba Regarding Anthropogenic and Environmental Factors and Ecological Relationships
15.3.1 Temporal Trends of Cuban Coral Reefs
15.3.2 Effects of Environmental Factors in Cuban Coral Reefs
15.3.3 Effects of Anthropogenic Factors in Cuban Coral Reefs
15.3.4 Ecological Relationships in Cuban Coral Reefs
15.4 Materials and Methods
15.5 Results and Discussion
15.5.1 Cuban Coral Reefs Trends
15.5.1.1 Temporal Trends of Ecological Indicators
15.5.1.2 Temporal Trends of Environmental Factors
15.5.1.3 Temporal Trends of Anthropogenic Factors
15.5.2 Environmental and Anthropogenic Factors Related to the Status of Cuban Coral Reefs
15.5.2.1 Environmental Factors Related to the Status of Cuban Coral Reefs
15.5.2.2 Anthropogenic Factors Related to the Status of Cuban Coral Reefs
15.5.3 Potential Ecological Relationships on Cuban Coral Reefs
15.5.3.1 Potential Ecological Relationships of D. antillarum
15.5.3.2 Potential Ecological Relationships of Corals
15.5.3.3 Potential Ecological Relationships of Fish
15.6 Integration of Results
15.7 Recommendations for Management and Research
References
16: Population Genetics of Cuba´s Scleractinian Corals
16.1 Introduction
16.2 Genetic Diversity
16.3 Genetic Differentiation and Connectivity
16.4 Information Gaps and Future Avenues of Research
16.5 Conclusion
References
17: Multiple Cumulative Effects on Coral Reefs of the Northwestern Cuban Region
17.1 Introduction
17.2 Coral Reefs in the Northwestern Cuban Region
17.2.1 Environmental Gradient in Northwestern Region
17.2.2 Multiple Stressors in Northwestern Region
17.2.2.1 Anthropogenic
17.2.2.2 Natural
17.3 Coral Reefs Studies from a Multiple Cumulative Effects´ Perspective
17.3.1 Selected Ecological Indicators
17.3.1.1 Community Level
17.3.1.2 Population Level
17.3.1.3 Statistical Analyses
17.4 Responses of Reefs Communities to the Cumulative Effect of Multiple Stressors
17.4.1 Responses of Benthic Communities
17.4.2 Responses of Fish Assemblages
17.4.3 Fish and Coral Interactions
17.4.4 Coral Disease, Bleaching, and Mortality
17.4.5 Impacts of Meteorological Events
17.5 Final Conclusions
17.6 Future
References
18: Guanahacabibes National Park: Research, Monitoring, and Management for the Conservation of Coral Reefs
18.1 Introduction
18.2 Coastline and Fringing Coral Reefs of Guanahacabibes
18.3 Marine Diversity in Guanahacabibes National Park
18.4 Coral Communities
18.5 Bleaching Events
18.6 Coral Transplanting (Acropora cervicornis)
18.7 Fish Communities
18.8 Lionfish and Native Fish Communities
18.9 International Lionfish Fishing Tournaments
18.10 Tools, Strategies, and Regulations Aimed at Tourism Activities Developed in Coral Reefs
18.11 Guanahacabibes National Park Management Effectiveness
18.12 Final Considerations
References
19: Ciénaga de Zapata Biosphere Reserve: Integrating Science with the Management of Coral Reefs
19.1 Introduction
19.2 Coral Reefs Under Different Management Categories
19.3 Main Uses, Impacts and Management Measures on Coral Reefs in the RBCZ
19.3.1 Fishing
19.3.2 Diving
19.3.3 Coral Bleaching Events
19.3.4 Climate Impacts
19.3.5 Water Dynamics and Fringing Reefs in the RBCZ
19.4 Local Context for Integrated Management of Coral Reefs in the RBCZ
19.5 General Considerations
References
20: Coral Reefs in Cuban Marine-Protected Areas
20.1 Introduction
20.2 Representativeness of Coral Reefs in Cuban MPAs
20.3 Plans, Programs and Projects That Have Contributed to the Conservation of Coral Reefs in Cuban MPAs
20.4 Coral Reef Monitoring Efforts in the Cuban MPAs
20.5 Main Uses, Impacts and Challenges Faced by Coral Reefs in Cuban MPAs
20.6 Conclusions
References
Part VI: Economic Valuation
21: Economic Valuation of the Coral Reefs of Jardines de la Reina and Punta Francés National Parks, Cuba
21.1 Introduction
21.1.1 Ecosystem Services of Coral Reefs and Their Benefits to Coastal Communities
21.1.2 Economic Valuation of Environmental Goods and Services of Coral Reefs
21.1.3 Valuation Methods
21.2 Total Economic Value of Coral Reefs
21.2.1 Study Area
21.3 Total Economic Value of the Coral Reefs of Punta Francés and Jardines de la Reina National Parks
21.3.1 Direct Use Value of Jardines de la Reina National Park´s Coral Reefs
21.3.2 Indirect Use Value of Jardines de la Reina National Park´s Coral Reefs
21.3.3 Option Value of the Jardines de la Reina National Park´s Coral Reefs
21.3.4 Existence Value of the Jardines de la Reina National Park´s Coral Reefs
21.3.5 Integration of Environmental Goods and Services of the Jardines de la Reina National Park´s Coral Reefs
21.3.6 Direct Use Value of Punta Francés National Park´s Coral Reef
21.3.7 Indirect Use Value of Punta Francés National Park´s Coral Reef
21.3.8 Option Value of Punta Francés National Park´s Coral Reef
21.3.9 Integration of Environmental Goods and Services of Punta Frances National Park´s Coral Reefs
21.4 Discussion
21.4.1 Limitations on Case Studies
21.4.2 Value of Cuban Coral Reefs
21.5 Implications for Conservation and Management
References
22: The Economic Value of Coral Reefs in the Context of Marine-Protected Areas: Experiences of the South Cuban Archipelago Pro...
22.1 Introduction
22.2 Description of Economic Evaluation Case Studies
22.2.1 Criteria for the Selection of Study Sites and Methodology
22.2.2 Site Description
22.2.3 Conservation Status, Uses and Threats of Coral Reefs in Guanahacabibes NP and Desembarco del Granma NP
22.2.4 Economic Valuation Framework
22.2.5 Data Collection
22.3 Outcomes of Economic Evaluation Case Studies
22.3.1 Economic Valuation of Ecosystem Goods and Services
22.3.2 Guanahacabibes NP
22.3.2.1 Tourism and Recreation
22.3.2.2 Habitat of Key Species
22.3.2.3 Scientific Educational Value
22.3.3 Desembarco del Granma NP
22.3.3.1 Coastal Protection
22.3.3.2 Habitat for Species
22.3.3.3 Nature Tourism
22.3.3.4 Proposal of Alternative Sustainable Economic Activities for Coastal Communities
22.4 Evaluation of Studies and Lessons Learned
22.5 Conclusions
References
23: Fish Can Be more Valuable Alive than Dead
23.1 Introduction
23.2 Recreational Fisheries for Bonefish, Tarpon and Permit: International Examples
23.3 SCUBA Diving and Snorkeling with Sharks, Rays, Large Fishes and Marine Turtles: International Examples
23.4 Materials and Methods
23.5 Results
23.6 Discussion and Conclusions
References

Citation preview

Coral Reefs of the World  18

Vassil N. Zlatarski · John K. Reed Shirley A. Pomponi · Sandra Brooke Stephanie Farrington Editors

Coral Reefs of Cuba

Coral Reefs of the World Volume 18 Series Editors Bernhard M. Riegl, Nova Southeastern University, Dania Beach, FL, USA Richard E. Dodge, Nova Southeastern University, Dania Beach, FL, USA

Coral Reefs of the World is a series presenting the status of knowledge of the world's coral reefs authored by leading scientists. The volumes are organized according to political or regional oceanographic boundaries. Emphasis is put on providing authoritative overviews of biology and geology, explaining the origins and peculiarities of coral reefs in each region. The information is so organized that it is up to date and can be used as a general reference and entry-point for further study. The series will cover all recent and many of the fossil coral reefs of the world. Prospective authors and/or editors should consult the Series Editors B.M. Riegl and R.E. Dodge for more details. Any comments or suggestions for future volumes are welcomed: Dr. Bernhard M. Riegl/Dr. Richard E. Dodge Nova Southeastern University Dania Beach, FL 33004 USA e-mail: [email protected] and [email protected]

Vassil N. Zlatarski • John K. Reed • Shirley A. Pomponi • Sandra Brooke • Stephanie Farrington Editors

Coral Reefs of Cuba

Editors Vassil N. Zlatarski Independent Scientist Bristol, RI, USA Shirley A. Pomponi Harbor Branch Oceanographic Institute Florida Atlantic University Fort Pierce, FL, USA

John K. Reed Harbor Branch Oceanographic Institute Florida Atlantic University Fort Pierce, FL, USA Sandra Brooke Coastal and Marine Lab Florida State University St. Teresa, FL, USA

Stephanie Farrington Techglobal Inc Rockville, MD, USA

ISSN 2213-719X ISSN 2213-7203 (electronic) Coral Reefs of the World ISBN 978-3-031-36718-2 ISBN 978-3-031-36719-9 (eBook) https://doi.org/10.1007/978-3-031-36719-9 # The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover Image: Satellite image of the reticulate coral reef system Grand Banco de Buena Esperanza, SE Cuba. # Spalding MD, Rivilious C, Green EP (2001) World Atlas of Coral Reefs. Univ California Press. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Foreword

Cuba is the largest island in the Caribbean and a hotspot of biological diversity. Its vast land area with wonderfully diverse and rich habitats is matched by its wealth in marine resources— among them one of the biggest coral reef systems in the Caribbean. For coral reef scientists, Cuba has long been a promised, yet for many forbidden, land and final frontier of marine research and exploration. The vagaries of politics have long caused a relative isolation of this biodiverse, culturally and scientifically rich island with a vibrant academic culture and strong research. While information about its reefs and the work of its scientific community was of course available, the sheer biogeographic and ecological importance as well as the vast academic capability of this island’s scientists merits to be highlighted in a special contribution to this book series. This is exactly what this present volume, “Coral Reefs of Cuba,” which is number 18 in the series “Coral Reefs of the World,” aims to do. This book was produced with the goal of providing an overview of the status of knowledge of Cuban coral reefs and to provide a baseline from which further investigations can depart. Importantly, it opens much of the knowledge that has been developed inside Cuba and its institutions in an easily accessible way to a wider international, English-speaking, audience. It is a scholarly review as opposed to a status report and there is no claim that the materials presented are complete and will satisfy everybody’s interest. Nonetheless, it should become a key work of reference of where to start reading when interested in Cuba’s reefs and the citations contained within will also serve as a guide to an important body of literature that is not always so easy to come by. Experts from within and without Cuba have collaborated to present in this volume the widest possible overview of present knowledge of these most interesting reef systems. Cuba has been a cradle of modern reef research, which is highlighted by V. Zlatarski’s first chapter giving a rich overview of what can be learned from Cuban reefs. Zlatarski himself is one of the pioneers of modern coral reef research who has been investigating these reefs since the 1970s. The research history of Cuba’s reefs is treated in a separate chapter by S. González-Ferrer (Chap. 2), himself a key player. Part III is a special section of the volume dedicated to the physical description of the reef edifices as a whole. The physico-geographic properties of the modern reefs (Chap. 3 by Estrada Estrada et al.), the relevance of the uplifted marine terraces for modern reefs (Chap. 4 by Iturralde-Vincent and Hine), and the use of remote sensing to describe extent and geomorphology of the reefs (Chap. 5, Andréfouët and Bionaz) are discussed. Part IV treats the distribution and nature of biota, and these chapters should be of key interest to anybody working anywhere in the Caribbean, given Cuba’s central role to connectivity. Among the benthos, the macrophyte flora (Chap. 6, Suárez and Martínez-Daranas), the sponge fauna (Chap. 7, Busutil and García-Hernández), the stony corals (Chap. 8, González-Ferrer, Cairns and Zlatarski) and octocorals, so important on Caribbean reef systems (Chap. 9, Rey-Villiers et al.), are treated in detail. Chapters on reef mollusks (Chap. 10, Espinosa and González-Ferrer), herbivory and herbivores (Chap. 11, Duran et al.), the Lionfish invasion (Chap. 12, Chevalier Monteagudo et al.), and ecological and fisheries aspects of sharks and rays (Chap. 13, Pina-Amargós et al.) provide a more process-oriented view for the reader. v

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Chapter 14 by Reed et al. describes the rich mesophotic (deep) reefs surrounding Cuba. Part V leads us through the ecology of these reefs as relevant for conservation and management. We are informed on the status of coral reefs in Cuba (Chap. 15, Pina-Amargós et al.), the population genetics of corals (Chap. 16, Ulmo-Díaz et al.), and stressor effects in NW Cuba (Chap. 17, González-Díaz et al.). More regional aspects are described in chapters on the coral reefs in Guanahacabibes National Park (Chap. 18, Cobián-Rojas et al.), the Cienaga de Zapata Biosphere Reserve (Chap. 19, González-Méndez et al.), and in Cuba’s marine protected areas in general (Chap. 20, Perera-Valerrama et al.). These chapters reveal the strong approach taken in Cuba to the conservation of these critically important ecosystems. The final Part VI of the volume deals with economic valuation of reefs (Chap. 21 by Figueiredo-Martín et al. and Chap. 22 by Ferro-Azcona et al.) and the fish that live within (Chap. 23 by Figueiredo-Martín and Pina-Amargós). Even after just glancing cursorily over the chapters presented in this volume, there will no doubt in anybody’s mind about the importance of Cuba’s coral reefs and the hard work and competence of its scientists. As editors of the book series, we thank authors and volume editors for the hard work that has gone into the production of this beautiful contribution. It should have a place of honor in any library or any interested person’s bookshelf and we are certain that it will be amply cited. Dania Beach, FL, USA March 2023

Bernhard Riegl Richard Dodge

Preface

Cuban coral reefs are uniquely valuable and important, as demonstrated by Hawthorne L. Beyer and 20 colleagues (Risk-sensitive planning for conserving coral reefs under rapid climate change. Conservation Letters. 2018;11:e12587. https://doi.org/10.1111/conl.12587). They identified a global portfolio of 50 bioclimatic units (BCUs) for conservation investment that maximizes the chances these coral reefs are secure in the future. Five of the 50 BCUs in the entire world are in Cuba (10%), as are five of the total of six in the Caribbean (83%)! This volume represents the first English language overview of Cuban coral ecosystems, which are among the healthiest in the Caribbean. It is our hope that this information will be of value to others who work on these fragile and endangered ecosystems. This project was initiated in 2020 at the suggestion of Dr. Bernhard Riegl, Editor of the Series “Coral Reefs of the World.” It was an exciting and timely project, but it began under the difficult conditions of the COVID-19 pandemic. Since most of the authors spoke Spanish as their native language, the first challenge was to find qualified Editors who were also fluent in Spanish and English. From the original Editorial Board, Vassil Zlatarski and Sergio GonzálezFerrer developed the book structure and engaged the chapter authors. More than 100 colleagues enthusiastically joined the project and overcame the challenges of quarantine and disrupted communications to produce the first chapter drafts in Spanish and received editorial suggestions from Vassil and Sergio. Unfortunately, Sergio had to withdraw as Editor in January 2021, but continued to play a critical role in communicating with the Cuban authors and assisting with the editorial process as an “unofficial” Editor until the end of the project. The next challenge was the translation of the Spanish manuscripts to English with limited financial resources. Some authors recruited translators, but additional assistance was generously provided by the Harte Research Institute (Texas Agricultural and Mechanical UniversityCorpus Christi) and the Environmental Defense Fund. During the review period in October 2021, another member of the original editorial board (Benjamin J. Greenstein) had to withdraw, leaving the sole remaining Corresponding Editor to recall the words of Dr. Yossi Loya: “During the process of editing the mesophotic book, some authors got married, some divorced, and others had personal reasons for delaying their contribution. There were fires in California, Hurricanes demolishing labs, you name it.” The dictum that there is always somebody who has the interest and is willing to devote their efforts to a worthwhile cause led to the present Editorial Board with four new members: John K. Reed, Shirley A. Pomponi, Sandra Brooke, and Stephanie Farrington. During 2022, John, Shirley, and Sandra contacted the reviewers to complete their reviews, edited the chapters for content and language, and finalized the reviews of each chapter’s multiple revisions. Special thanks to Stephanie for organizing the large number of documents and correspondence. The Editorial Board is now happy to present the volume “Coral Reefs of Cuba” with deep gratitude for the tireless contributions of the enthusiastic authors, patient translators, generous

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Preface

facilitators, eminent experts, volunteer reviewers, and both former Co-Editors, all during extraordinary circumstances. Bristol, RI, USA Fort Pierce, FL, USA Fort Pierce, FL, USA St. Teresa, FL, USA Rockville, MD, USA

Vassil N. Zlatarski John K. Reed Shirley A. Pomponi Sandra Brooke Stephanie Farrington

Contents

Part I 1

Insights from Cuban Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vassil N. Zlatarski

Part II 2

Introduction

History

Research History of Corals and Coral Reefs in Cuba . . . . . . . . . . . . . . . . . . . Sergio González-Ferrer

Part III

Physical-Geographic Characteristics of Cuban Reefs . . . . . . . . . . . . . . . . . . . Reinaldo Estrada Estrada, Gustavo Martín Morales, Joán Hernández-Albernas, Rodney Borrego Acevedo, Jorge Olivera Acosta, Yudelsy Carrillo Betancourt, Idalmis Almeida Martínez, and Lourdes Coya de la Fuente

4

Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf and Uplifted Marine Terraces of Cuba: Tectonic and Sea-Level Control of Present-Day Coral Reef Distribution . . . . . . . . . . . . . . . Manuel A. Iturralde-Vinent and Albert C. Hine A Remote Sensing Appraisal of the Extent and Geomorphological Diversity of the Coral Reefs of Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serge Andréfouët and Océane Bionaz

Part IV

19

Description

3

5

3

51

75

93

Biota

6

Macrophytes Associated with Cuban Coral Reefs . . . . . . . . . . . . . . . . . . . . . . 111 Ana M. Suárez and Beatriz Martínez-Daranas

7

Sponges: Conspicuous Inhabitants of the Cuban Coral Reefs and Their Potential as Bioindicators of Contamination . . . . . . . . . . . . . . . . . . . . . . . . . . 127 Linnet Busutil and María R. García-Hernández

8

Species List of Cuban Stony Corals: Class Anthozoa, Order Scleractinia; Class Hydrozoa, Suborders Capitata and Filifera . . . . . . . . . . . . . . . . . . . . . . 147 Sergio González-Ferrer, Stephen D. Cairns, and Vassil N. Zlatarski

9

Octocoral Forests: Distribution, Abundance, and Species Richness in Cuban Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 Néstor Rey-Villiers, Leslie Hernández-Fernández, Hansel Caballero, Mayilen Triana López, Alejandro Pérez Angulo, and Yunier Olivera Espinosa

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Contents

10

Current State of Knowledge of Reef Mollusks in Cuba . . . . . . . . . . . . . . . . . . 185 José Espinosa and Sergio González-Ferrer

11

Herbivory on Cuban Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 Alain Duran, Patricia González-Díaz, Rodolfo Arias, Dorka Cobián-Rojas, Pedro Chevalier, Tamara Figueredo, Alain García-Rodríguez, Ariagna Lara, Yunier Olivera, Susana Perera-Valderrama, and Fabián Pina

12

Chronology of the Lionfish Invasion in Cuba and Evaluation of Impacts on Native Reef Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Pedro Pablo Chevalier Monteagudo, Dorka Cobián Rojas, Raúl Igor Corrada Wong, Alexis Fernández Osoria, Hansel Caballero Aragón, Delmis Cabrera Guerra, Zenaida María Navarro Martínez, and Leandro Rodríguez Viera

13

Sharks and Rays in Cuban Coral Reefs: Ecology, Fisheries, and Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 Fabián Pina-Amargós, Yunier Olivera-Espinosa, Alexei Ruiz-Abierno, Rachel Graham, Robert Hueter, Juan Fernando Márquez-Farías, Aracelys Hernández-Betancourt, Raidel Borroto-Vejerano, Tamara Figueredo-Martín, Alejandra Briones, Yureidy Cabrera-Páez, Ariandy González-González, Consuelo Aguilar-Betancourt, and Gaspar González-Sansón

14

Mesophotic Coral Ecosystems of Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253 John K. Reed, Patricia González-Díaz, Joshua D. Voss, Linnet Busutil, Cristina Diaz, Shirley A. Pomponi, Stephanie Farrington, Dorka Cobián-Rojas, Andrew David, Beatriz Martínez-Daranas, M. Dennis Hanisak, Juliett González Mendez, Alexis B. Sturm, Patricia M. González Sánchez, María Rosa García Hernández, Jorge Viamontes Fernández, Mingshun Jiang, Jack H. Laverick, and Vassil N. Zlatarski

Part V

Ecology, Conservation and Management

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Status of Cuban Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283 Fabián Pina-Amargós, Patricia González-Díaz, Gaspar González-Sansón, Consuelo Aguilar-Betancourt, Yandy Rodríguez-Cueto, Yunier OliveraEspinosa, Tamara Figueredo-Martín, Néstor Rey-Villiers, Rodolfo Arias Barreto, Dorka Cobián-Rojas, Rodolfo Claro, Susana Perera-Valderrama, Zenaida María Navarro-Martínez, Enrique Reynaldo-de la Cruz, Alain Durán, Yenizeys Cabrales-Caballero, Leonardo Espinosa-Pantoja, Zaimiuri Hernández-González, Hansel Caballero-Aragón, Pedro Pablo Chevalier-Monteagudo, Juliett GonzálezMéndez, Leslie Hernández-Fernández, Susel Castellanos-Iglesias, Ariagna Lara, Alain García-Rodríguez, Linnet Busutil, Carlos Luis Reyes Suz, Joán Irán Hernández-Albernas, Aloyma Semidey Ravelo, and Pedro Alcolado Prieto

16

Population Genetics of Cuba’s Scleractinian Corals . . . . . . . . . . . . . . . . . . . . 309 Gabriela Ulmo-Díaz, Jessy Castellanos Gell, Didier Casane, Alexis Sturm, Joshua Voss, and Erik García-Machado

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Multiple Cumulative Effects on Coral Reefs of the Northwestern Cuban Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 317 Patricia González-Díaz, Gaspar González-Sansón, Consuelo Aguilar-Betancourt, Néstor Rey-Villiers, Alain Duran, Orlando Perera Pérez, and Sergio Álvarez Fernández

Contents

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18

Guanahacabibes National Park: Research, Monitoring, and Management for the Conservation of Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339 Dorka Cobián-Rojas, Susana Perera-Valderrama, Pedro Pablo ChevalierMonteagudo, Juan J. Schmitter-Soto, Raúl Igor Corrada Wong, Elena de la Guardia Llansó, Juliett González Mendez, Alain García-Rodríguez, Joan Hernández-Albernas, Lázaro Márquez-Llauger, Zaimiuris Hernández-González, Leonardo Espinosa Pantoja, Patricia González-Díaz, and Hansel CaballeroAragón

19

Ciénaga de Zapata Biosphere Reserve: Integrating Science with the Management of Coral Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359 Juliett González-Méndez, Susana Perera-Valderrama, Hansel Caballero-Aragón, Dorka Cobián-Rojas, Pedro Chevalier Monteagudo, Tania Piñeiro Cordero, Reynaldo Santana Aguilar, Leyaní Caballero-Thiert, Raúl I. Corrada Wong, Silvia Patricia González-Díaz, Dinorah Millán Caballero, and Jorge Luis Jiménez Hernández

20

Coral Reefs in Cuban Marine-Protected Areas . . . . . . . . . . . . . . . . . . . . . . . . 375 Susana Perera-Valderrama, Juliett González-Méndez, Aylem Hernández-Ávila, Reinaldo Estrada-Estrada, Dorka Cobián-Rojas, Adonis Ramón-Puebla, Elena de la Guardia-Llansó, Hakna Ferro-Azcona, Joán Hernández-Albernas, Zaimiuri Hernández-González, Leonardo Espinosa-Pantoja, Ariagna Lara, Fabián PinaAmargós, Patricia González-Díaz, Pedro Pablo Chevalier-Monteagudo, Néstor Rey-Villiers, Jorge Antonio Tamayo-Fonseca, and Hansel Caballero-Aragón

Part VI

Economic Valuation

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Economic Valuation of the Coral Reefs of Jardines de la Reina and Punta Francés National Parks, Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395 Tamara Figueredo-Martín, Laura López-Castañeda, and Fabián Pina-Amargós

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The Economic Value of Coral Reefs in the Context of Marine-Protected Areas: Experiences of the South Cuban Archipelago Project . . . . . . . . . . . . . . . . . . . 415 Hakna Ferro-Azcona, Gloria de las Mercedes Gómez-País, Susana PereraValderrama, Dorka Cobián-Rojas, Alberto González-Tejeda, Berta LizanoMachado, Adaris Calderín González, Orlando Acosta-Rodríguez, Raisa Escalona-Domenech, and Adonis Ramón-Puebla

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Fish Can Be more Valuable Alive than Dead . . . . . . . . . . . . . . . . . . . . . . . . . 429 Tamara Figueredo-Martín and Fabián Pina-Amargós

Part I Introduction

1

Insights from Cuban Coral Reefs Vassil N. Zlatarski

Abstract

In the 1970s, the Cuban coral reefs were the object of extensive exploration. The results were published in Spanish, Russian, French, and German but unfortunately remain relatively unknown. This chapter provides, for the first time, English translations of excerpts from the study of Cuban reefs conducted in 1970–1973. The obtained knowledge had been planned to be published in a monograph and colored atlas of photos of the underwater scleractinians, but only the monograph was published (in Russian, French, and Spanish). Recent specialists’ opinions and the author’s reflections offer present-day insights from Cuban coral reefs regarding different aspects of the pioneer investigation: sampling and material, methodology, area study, terminology, variability, hybridization and morphogenesis, invasion, state of Scleractinia knowledge, reef zonation, reticulated reefs on muddy bottom, and reef health and care.

knowledge. The results were published in Spanish, Russian, French, and German but unfortunately remain relatively unknown (Zlatarski 1980, 1982, 2018b; see Chap. 2, Gonzalez-Ferrer). This chapter is an attempt to share some insights from a longtime researcher’s interest in coral reefs and to consider how the results of the study conducted in Cuban waters in 1970–1973 resonate after all these many years. Since the original publication of the monograph “The scleractinians of Cuba” (Zlatarski 1980), some species and genus names have been synonymized, and new taxa were described. In these cases, the International Code of Zoological Nomenclature term auct. is used. This is a Latin term meaning “of authors,” often given to indicate that a name is used in the sense of a number of subsequent authors and not in its (different) sense as established by the original author. The next two sections on reef types and ecological zonation (Sect. 1.2) and scleractinian ecology (Sect. 1.3) present, for the first time, English translations of excerpts from the monograph on Cuban scleractinians.

Keywords

Coral reef types · Reef zonation · Ecology · MCEs · Scleractinia · Variability

1.1

Introduction

Cuban reefs represent the largest coral reef system in the Caribbean. It is therefore of great importance for a complete understanding of coral and reef dynamics in the Caribbean basin but compared to other more frequently visited locations like Jamaica or Mexico, information is still relatively sparse. However, much information has been collected over the years, even though it may not have been available in English. In the 1970s, Cuban reefs were the object of systematic international studies resulting in considerable ecosystem V. N. Zlatarski (✉) Bristol, RI, USA

1.2

Coral Reef Types and Ecological Zonation of the Scleractinians

1.2.1

General Notes

Reefs abound in the Cuban archipelago. Of the three classic types of reefs (fringing, barrier, and atoll), only the first two were found, and a special type was found on the muddy bottom, observed only in the Gulf of Guacanayabo (Fig. 1.1). The studied profiles are distributed according to reef type (Fig. 1.2): on the muddy bottom, six profiles; fringing reefs, 15; barrier reefs, 20; and three transitioning between the last two. The following reasons were decisive for reef construction in Guacanayabo: the hydrodynamic pattern, the substrate, and the lack of competitors that inhabit the bottom (greater detail is available in Sect. 1.2.4).

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_1

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Fig. 1.1 Types of reefs in the Cuban archipelago

The data on salinity, temperature, and direction of the currents in Cuban waters (Núñez Jiménez 1970) signal that these physical–geographic conditions did not influence the formation of fringing and barrier reefs. A decisive factor behind the formation of these two types was the shelf width or, more exactly, the location of the limits between the shelf and the continental slope. There, the scleractinians have optimal living conditions, permanently clear and moving waters, and waters are rich in food particles. Further, the water–temperature variation in that area conditions the rapid accretion of sediment. Once the barrier has emerged by itself, it helps to further highlight the differences among nearby areas. Thus, when the shelf is wide, barrier reefs form, and when it is narrow or does not exist, fringing reefs arise. The width of the platform changes gradually as long as the boundaries between the two types of reefs are conditioned.

By scuba diving, it was established that reef formation also continues far from the breaker indicated by the reef barrier and away from the coasts abundantly covered with coral, occupying a larger space than that indicated for the shelf and the continental slope. Often, the continental slope at depths of 55–65 m and deeper was covered by scleractinians. Further, by penetrating deeper toward caverns along the bottom and caves, coral life was discovered in places that dredges could not reach. The presence of reef formations, including small reef bodies with dimensions from a few meters to 10 or 20 m and with irregular contours, were observed in the sand channels in the reef barrier, in some lagoons, or in front of the fringing reefs. They are called cabezos (from “head” in Spanish; known as “coral head” or “coral knoll,” in English; “massif corallien,” in French) and were made up of huge

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Insights from Cuban Coral Reefs

Fig. 1.2 Reef types of the Cuban archipelago (profile numbers refer to Fig. 1.1)

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Fig. 1.3 Types of fringing reefs in Cuba and their zonation as seen in the 1970s; (a) fringing reef on steep rocky coast; (b) fringing reef on a slightly sloping sandy beach; (c) transition between fringing and barrier reefs

groups of coral colonies, which concentrate a world of organisms from the less inhabited nearby environment. The specific content was very diverse. Almost all the scleractinian species found in Cuban waters were found here. Modern Cuban reefs are not considered thick (not exceeding 30 m). This is due to frequent changes in the sea level during the geological past. Many times, reefs emerged above sea level, and as a result of denudation, the colonies that formed them accumulated in terraces. These are the Pleistocene and Holocene coastal terraces that border much of the island. Reef formation was established during the Miocene, near the cities of Santiago de Cuba and Matanzas and in the former Province of Las Villas (this latter location also shows signs of reef formation during the Paleogene, with coral banks dating from the Upper Cretaceous).

1.2.2

Fringing Reefs

Fringing reefs emerged in the coastal areas between the large archipelagos (Fig. 1.1). In these locations, shelf width is negligible or simply virtually absent. In the latter case, the continental slope descends from the coast. We confirmed that, in all their extension, the various fringing reefs are not inhabited by the same species of

Scleractinia due to the heterogeneity of the ecological environment. In one case, the shore rock abruptly drops vertically to a depth of 4 m or more (Fig. 1.3a; Profile 33, Playa Siboney; Profile 39, Punta Seboruco). Sea waves crash upon these rocks, hindering the growth of the corals and leaving rocks uninhabited. Only few flat colonies of Siderastraea radians were able to affix themselves and exist, strongly anchored to the bottom. Sometimes Agaricia agaricites forma massiva and Diploria auct. appeared. The community of Scleractinia in those poorly populated areas was dominated by Siderastraea. Moving away from the coast, colonies of Acropora palmata appeared. First, they were flat, short, and well affixed to the bottom; they increased in height with increasing depth and distance from the coastal waves. The branches appeared not only on a single plane, but instead, their colonies successively undertook isometric development. Here, the hydrodynamic movement was less intense. We can often find fragile colonies of Agaricia agaricites forma bifaciata. The community Acropora palmata–Agaricia agaricites forma bifaciata was followed by dense and graceful “shrubs” of Acropora cervicornis. This is why the community was also named after this species. Among them, perpendicular to the coastline, there were grooves or channels with depths of 1–3 m along which the compensatory

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Insights from Cuban Coral Reefs

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Fig. 1.4 Barrier reef zonation in Cuba as seen in the 1970s

movement of the bottom waters took place. Due to this movement, the bottom had bare rock or detrital sand. On the slopes, massive or spherical colonies arose, most often Montastraea auct. and Diploria auct. The community is named after Diploria–Montastraea. At greater depths, the conditions for coral life of fringing reefs and barrier reefs are similar; thus, the communities were similar, for which reason they will be discussed later. When the coast was not made up of steep rock, but rather a slightly sloping sandy beach so that the impact of abrasion was much less, corals inhabited the bottom in the immediate vicinity of the coast (e.g., Fig. 1.3b and Profile 6, 14 km). Sometimes, a narrow (1–3 m) channel with sand at the bottom abutted the shore. In this case, the Siderastraea community did not take hold, and in general, the scleractinians were not found there. Behind the channel, moving away from the beach, Acropora palmata colonies again took on a predominant role, growing in dense, almost massive, low formations with short branches. Among them, in this profile, there were “oases” of A. cervicornis. Sometimes, the surface

of the reef was covered by the massive growth of the soft coral Zoanthus. As the shelf widened, a lagoon channel was formed, and the zonation became increasingly similar to that of the barrier reef (Fig. 1.3c and Profile 16, Institute of Oceanology). An extreme case was that of a wide band of coral sand that descended smoothly without coral inhabitants and a particular development of the fringing reef in front of Varadero (Profile 35). There, the sand bottom influenced the absence of the first communities. It only appeared on the outer slope of the reef and on the continental slope, where scleractinian life was more abundant. In the transition type, the bottom of the outer slope of the reef and the continental slope also had the same number of scleractinian communities (see Figs. 1.3a and 1.4) as those of the fringing and barrier reefs, due to similar conditions. As the coast grew increasingly remote, communities of Diploria–Montastraea and Agaricia agaricites forma unificata continued.

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1.2.3

V. N. Zlatarski

Barrier Reefs

Barrier reefs emerge on a relatively wide shelf, i.e., when the continental slope is quite far from the coast. They grow near the edge of the shelf. This is why the reefs are facing the archipelagos of Los Colorados, Sabana-Camagüey, Jardines de la Reina, and Los Canarreos (Fig. 1.1). In the cuneiform area of the shelf, the previously mentioned type of transition was observed between the fringing and barrier reefs (Fig. 1.3c). In barrier reefs, unlike fringing reefs, a clearer ecological zonation of coral communities was established. The change of conditions on shelf edge with continental slope causes the barrier to appear, creating great variability in the ecological environment in front of, above, and behind it. Therefore, certain communities of Scleractinia were ascribed to the different elements of the underwater relief (continental slope, outer slope, outer edge, reef flat, lagoon slope) (Fig. 1.4). All of them were found in the complexes “which correspond to homogeneous conditions: morphological, hydrodynamic and sedimentological, bionomic” (Battistini et al. 1975, p. 25). The following complexes were established in the barrier reef: prelitoral, front reef, epireef, and postreef (the last two were absent in fringing reefs). The creation of complexes was based on the particularities of the reefs—the only community in the physical, geographical, and organic environment. Therefore, despite being defined according to the particular types of reef formation (near the island of Madagascar), complexes also occur in Caribbean waters. Local conditions of Cuban waters prepared the subdivisions of the complexes described here as zones with their corresponding communities (Fig. 1.4). The continental slope creates a very asymmetrical ecological environment, because the rock on which the corals grow is very inclined, in some places almost steep and the sun’s rays barely shine there. This favored the growth of flat colonies, laterally affixed to the rock by one of their ends, with their distal surfaces sticking out perpendicularly toward the penetrating light. The absence of active movements of water masses explains the fragility of the colonies and the decrease of food at that depth; it also explains the need for a wider area to capture food, i.e., a greater distance between individuals in the colonies. The growth of the colonies on thin plates is carried out, optimally, through serial intracalicinal budding. These are, then, the main characteristics of Agaricia agaricites forma unifaciata, whose colonies more frequently inhabited the continental slope. Thus, the community was named after this form. At times, the continental slope was completely covered by Agaricia. Next to it occurred Helioseris cucullata, Mycetophyllia reesi, Mycetophyllia lamarckiana, Montastraea cavernosa, M. annularis auct., Mussa angulosa, Scolymia lacera, and others. When the continental

slope was inclined or tiered, the community was not as homogeneous. We found sand and mud banks with coral knolls, as well as steep slopes with a greater presence of coral. When the transition of the underwater relief, from the continental slope to the outer slope of the reef, was even steeper, the change in the scleractinian community was clear. Since the activity of the water masses on the outer slope was immense, the bottom was hardly slanted and well illuminated, and the environment was abounded with food. Thus, colonies were tightly attached, firm, massive, and quite often spherical; their diameter sometimes exceeded half a meter, reaching 2 and 3 m. When the bottom was rocky, it was greatly inhabited; when it was sandy, colonies are grouped around the first inhabitant that managed to attach itself, starting with the aforementioned knolls. The active movement of water masses with abundant food particles was favorable both for scleractinians and all sessile filters. There were many octocorals and sponges. The growth of the sponges damaged the scleractinians because they lived on the coral colonies and killed the polyps. In some cases, their aggression caused a small but regular damage to the distal surface of the colonies. The morphological characteristics of the coralla varied as if another species had been found. This pathological phenomenon was conditioned by ecological factors having to do with the aggressor, not the coral. Therefore, in areas favorable to sponges, affected scleractinian colonies were more frequent. In other cases, the sponges of the Cliona genus penetrated the colonies, creating a thick network of small channels, destroying the coralla internally. Yet, mutualistic relationships with sponges were not always antagonistic; they could also be favorable for both parties, when some sponge species covered the lower surfaces of the coral colonies, so that the edges of the colonies could arch in an undulating way. In this case, the sponges had a larger habitable area, without competitors, and the feeding of the corals was facilitated by a strong water current caused by the sponges; furthermore, their lower surfaces were protected from the aggressive Cliona sponges. In the limits of the external slope of the reef, two communities were established; the lower one occupied 95–99% of the width of the area, having greater specific diversity. Almost all species were found there, and their colonies were generally massive (spherical, cylindrical, bifacial, incrustation). It is difficult to point out a predominant species, which is why the area was called Diploria– Montastraea. The part of the outer slope, below the outer slope, was almost always inhabited by Millepora alcicornis forma alcicornis and thus the name of the community. Scleractinia were rarely found. The outer edge of the reef barrier is pounded by waves. The colonies of Millepora alcicornis forma complanata, with

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Insights from Cuban Coral Reefs

their fine vertical lamella, strongly welded together in polygonal constructions made up parts of the barrier and were the only attached animals that withstood the pounding of the waves and vibrations. The area was very narrow from 1 to 3–4 m and was only inhabited by these hydrozoans. The reef flat is the shallowest area. The maximum depth does not exceed 60–70 cm. At low tide, the upper parts of Acropora were temporarily exposed. The width of the zone varied (from 30–40 to 150 m); it was inhabited by stable Acropora palmata colonies, enormous like trees (diameters: 1.5 m or more), sometimes welded together massively, which is where the name of the community was derived. These “forests” of acroporid coral were sometimes impassable. Sometimes, small colonies of Favia fragum and few spherical colonies of the species, known for inhabiting the outer slope, developed at their base. Also, there were isolated patios between A. palmata, totally inhabited by P. porites and areas covered by Zoanthus. In some places, the reef barrier was crossed by channels, through which passed compensatory movements by water masses produced by local and tidal currents. Bottoms were excavated, rocky, and sometimes sandy with pebbles, and large, partially destroyed coral colonies. According to their active hydrodynamic nature, the channels were located at the bottom of the outer slope of the reef, with scarce inhabitants of the Diploria–Montastraea community. The inner slope of the reef was the most protected area from the waves; it was a narrow strip of medium width from 5 to 20 m, inhabited by the community of dense bushes of “staghorn coral,” Acropora cervicornis (thus the name of the community). In the same community also occurred Favia fragum, Dichocoenia stockesi, Agaricia agaricites forma bifaciata, Mycetophyllia lamarckiana, Siderastraea radians, Mussa angulosa, and others. In the lagoon, we usually found fragmented and overturned colonies of Acropora palmata that managed to regenerate by continuing their growth, but in opposite directions, as if “their feet were facing upward.” The presence of some colonies of A. palmata in the lagoon were indicative of the destructive force of past hurricanes. While the detritic sandy characteristic of the lagoon’s sediments could be explained by the destructive force of the waves, the establishment of the oolitic sandy accumulations, for example, in the Archipelago of Los Canarreos, illustrated what caused the formation of oolite: there the coldest oceanic water mixed with warm lagoon water. In the sandy “deserts” of the lagoon, knolls were made up of Siderastraea, Diploria auct., Meandrina auct., Mycetophyllia, Dichocoenia, Montastraea auct., and others. Their diameter rarely exceeded 2 or 3 m. The absence of a firm base caused incrustations to be more frequent in that area, which were usually the remains of other

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dead scleractinians, other organisms, and even pieces of different objects. The most frequent inhabitant of the lagoon was the rose coral, Manicina areolata, which gave its name to the community. They “floated” at the surface of the muddy and sandy sediments, allowing their reproduction in this habitat. However, their growth was limited by their critical weight, when the coral “escaped” only with difficulty from the sediments. For this reason, this species and other inhabitants of the lagoon were dwarf sized as a result of pedomorphism. In the lagoon, there were sometimes wide fields covered by Thalassia grass, often inhabited by the sea star Oreaster reticulatus and large gastropods. A part of the coral sand carried to the lagoon washed up to accumulate on the coasts, forming beaches or beach rock.

1.2.4

Reefs on Muddy Bottom

In the Gulf of Guacanayabo (Fig. 1.1), reefs with heights between 20 and 25 m grew on mud (now referred to as reticulate reefs—Zlatarski and Gonzáles Ferrer 2017; Zlatarski and Greenstein 2020) and were inhabited by species of Scleractinia that are not known as reef builders, such as Oculina spp., Cladocora arbuscula, Madracis decactis, Porites porites forma divaricata, and a new form of Eusmilia fastigiata with fine and small corallites. The colonies of Acropora cervicornis and the hydrozoan Millepora alcicornis, also usually found here, were smaller and more branched. The geological conditions and hydrodynamic nature of the gulf, i.e., muddy bottom and almost motionless turbid waters, were all very unusual and contraindicative for the formation of contemporary reefs. The dimly lit, still, and turbid waters were unsuitable for a rich benthic community. With visibilities of only 20–30 cm underwater, we were able to verify that the vertical ridges of the reefs were made up of small, highly branched colonies of the genus Oculina (also called ivory coral) and Cladocora arbuscula; these were the only ones that, with their light, highly branched colonies, managed to take hold on the soft muddy bottom. The absence of competitors and the wide uninhabitable areas at the bottom made it possible for them to disperse greatly and, together with the exuberant increase of sponges, “weave” these paradoxical “gelatinous” reefs. This latter name arises from the fact that divers are able to move large portions of these reefs almost effortlessly due to the presence of sponges. The body of these reef constructions was poor in specific diversity. No clear zonation or communities were apparent. In the upper part (depth of 2–5 m), the reef slopes were welded by a strip composed usually of Acropora cervicornis. The reef flat, with variable width (from 10 to 50 m), was often

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a desert covered by dead branches of Oculina spp., Acropora cervicornis, and Porites spp. or was heavily inhabited by Thalassia grass (these parts of the sea bed were locally known as “seibadal”). It is likely that the reef flat of the Gulf of Guacanayabo suffered serious destruction because of Hurricane Flora (1963). Often, “coral cemeteries” in this gulf recalled parts of the destroyed reef flat, covered in the Canarreos Archipelago or other Cuban reef locations, generally with the remains of Acropora palmata. Less frequently, on the reef slopes in Guacanayabo, Porites spp., Millepora alcicornis, Mussa angulosa, Mycetophyllia lamarckiana, Madracis spp., Siderastraea radians, and Eusmilia fastigiata forma guacanayabensis occurred. The deviations in the morphology of gulf inhabitants were considered ecoforma and pedoforma, due to the specific hydrological and sedimentological conditions of this isolated area of the Cuban shelf. Understanding this special type of reef is important not only for the most complete knowledge of the modern reefformation process. The observation of a living model of reef construction on the soft muddy bottom holds special interest for paleontologists and lithologists, who often find reefs “standing” over clay, which were built by small, highly branched colonies.

1.3

General Review of the Ecology of the Scleractinians of the Cuban Archipelago

Figure 1.5 depicts the presence of scleractinians according to their profiles, stations, associations, communities, and their role in reef construction as observed in Cuba of the 1970s. Data on the left outline the total number of profiles where the taxon was found and then the number of profiles according to the type of reef where it was encountered (i.e., fringing reefs, barrier reefs, reefs in transition between the previous two, and reefs on muddy bottom). To a great extent, these values represent the frequency with which the taxa were found in the reefs. Values on the right show the station in which each taxon was identified. First number is station and then number of stations according to the presence of the taxon in each association: present (found it locally), dominant (predominant of numbers or coverage of substratum, and highly dominant (predominant by more than half in numbers and coverage of substratum). Interestingly, only five taxa in 12 stations are highly dominant; 13 taxa are dominant in 52 stations; and all taxa are found as present. The presence of each taxon in the community is expressed semiquantitatively: casual, constant, and characteristic of the community. “Characteristic” taxa are indicated when they make up the community entirely or > 80%. These species

tended to be absent or very rare when found outside this community. Among these were Acropora cervicornis, Acropora palmata, Agaricia agaricites forma unifaciata, Manicina areolata, and Siderastraea radians. In such cases, the communities are named after the characteristic taxon. An original case occurred with Astrangia solitaria, which independently and massively covered the dark walls of the coastal reefs. “Constant” taxa are indicated when a single taxon did not strongly predominate in the community and superiority is shared among some quite permanent ones. These include Pseudodiploria strigosa, P. clivosa, Montastraea cavernosa, and Orbicella annularis, for the outer slope of the fringing and barrier reefs. Acropora palmata and Agaricia agaricites forma bifaciata were constant in the fringing reefs as they moved away from the coast. The other taxa (casual) were present in the community. According to their specific diversity, the communities could be monotaxonic or polytaxonic. Communities were more differentiable on barrier reefs. With different fringing reefs, there were frequent deviations due to the nature of the coast; on muddy bottom reefs, the zonation was hardly visible and communities were not clear. In the analysis of the morphology of the inhabitants of different communities, we note that the borders between them coincided with the isophenes (the lines that indicate the distribution of taxa of similar morphology). For example, in the barrier reefs in the lagoon, the inhabitants were most often nanoforms and pedoforms. The inner slope of the reef was inhabited by finely branched bushy colonies. The reef flat was covered by large, firm, tree-shaped colonies. The outer slope was “armed” with lamellar hydrozoans. On the outer slope, the most frequent were the spherical and bifacial representatives, and the continental slope was covered by fine monofacial colonies. The similarity in conditions caused the last two zones to be inhabited by the same scleractinians as in the fringing reef. Therefore, the isophenes in the two reef types coincided here. On the fringing reef, in the direction of land, finely branched shrub colonies appeared and then large colonies like trees. The absence of a barrier was the reason for the reverse distribution of isophenes in these parts of the fringing reef, in relation to barrier reefs. A convergent development also occurred in the muddy bottom habitat. Only Manicina areolata and Meandrina meandrites brasiliensis, with conical coralla, were present on the soft bottom. They never grew to large dimensions. The first has been shown to have the ability to absorb water by decreasing its specific weight, thus helping it to “stand” and “come out” of the mud (Zlatarski 2018b, p. 96). In accordance with their participation in reef building, the taxa were classified in four groups (Fig. 1.5): first-degree reef builders, second-degree reef builders, third-degree reef builders, and solitary participants.

Fig. 1.5 Presence of Scleractinia in the profiles, stations, associations, and communities studied (details in the text)

1 Insights from Cuban Coral Reefs 11

12

By themselves, first-degree reef builders made up the reef body in complete zones; these were Acropora cervicornis, Acropora palmata, and Agaricia agaricites forma unifaciata. Second-degree reef builders, together with others, played a decisive role in reef construction; these were Diploria auct., Montastraea annularis auct., Oculina diffusa, Agaricia agaricites forma bifaciata, and Porites porites forma divaricata. Third-degree reef builders were often present in reefs, but because of their small size or quantity, they were not very significant in reef construction. It was possible for a taxon to be characteristic for a community but at the same time being a third-degree reef builder in the area (e.g., Manicina areolata). Solitary participants were taxa in which one or some specimens have been found in the reefs or inhabit the rock caverns or caves. In such a group, we can even find a taxon characteristic for some community (e.g., Astrangia solitaria). Solitary participants usually had coralla with insignificant dimensions. The composition of the associations was related to depth. Polytaxonic associations did not inhabit shallower areas (inner slope and reef flat), nor greater depths. The causes of monotaxonity and the exceptional richness of coralla closest to the sea surface and those living at great depth were different. In the former, this was linked to changes in depth. After the last glaciation, only Acropora palmata and A. cervicornis adapted to living at the highest parts of the reef. In the latter, this was due to the homogeneity of deep ocean environments and some unfavorable conditions. The distribution of the Cuban scleractinians with depth in the 1970s is depicted in Fig. 1.6. Many of the Scleractinia inhabited greater depths than expected, but our research situation hindered our ability to study up to the inferior bathymetric limit. The diversity of taxa at depths is shown in Fig. 1.7. Generic variability was hardly influenced by depth. Specific diversity varied sharply closer to the abscissas (20 m depth), unlike the variability of the representatives of the infrasubspecific category, whose numbers remained almost constant up to 30 m depth, and then decreased up to 60 m. Below this depth, the three curves were vertically oriented, that is, the number of taxa did not change. Diversity was greatly controlled by the effects of urbanization near the water. Thus, for example, scleractinians rarely occurred in the waters off Havana at depths to 8–10 m. They did occur below this limit, up to 20 and 40 m and some at a depth of 55 m. For such cases, the so-called anthropogenic subclimax or disclimax was established. Depth did not influence the distribution of the sciaphilic species. They were equally found at significant depth, as in caverns and dark caves or on the lower surface of other scleractinians. With an extremely photophobic characteristic, these were Gardineria minor, Coenocyathus bartschi,

V. N. Zlatarski

Caryophyllia smithi, and Astrangia solitaria. The colonies of Tubastraea coccinea were found in blocks of steep rocks and overhanging rock, i.e., they show less photophilia. This species was found only on the southeastern coast of Cuba. It is probable that it is a species that now quickly emigrates and has not yet settled in all Cuban waters. Phyllangia americana is the only species that inhabited very turbid and brackish waters.

1.4

Contemporary Perspective

The author started to study fossil Scleractinia in the Lower Cretaceous, finding rich localities in Bulgaria that urged massive phenotype collecting. The applied systematic sampling discovered rich biodiversity and many new taxa and proved to be the adequate way to study unknown material (Zlatarski 1968). This sampling strategy appeared reliable also for the investigation of modern Cuban scleractinians, because, since the first encounters with them, it became clear that some samples “did not fit in the drawers” of described discrete species and showed intermediate characters. It was evident that sampling should be the first decisive step for well-grounded coral taxonomic and ecological conclusions. The result was the largest Atlantic scientific stony coral collection curated and available now for study in the Acuario Nacional, Havana. It prompted leaving the typological species approach and led to the study of species dynamics. In addition, along with the stony corals, the associated organisms were collected, described, and curated (Martínez Estalella 1980, 1982, 2018). The study is thoroughly explored by scuba all parts of the coral reef ecosystem accessible at that time. The underwater observations, collecting and profiling, were not regionally fragmented or limited to shallow reef zones (e.g., reef flat, front reefs). They were not stopped by the conventional doctrine, which pronounced that in tropical waters coral life disappeared deeper than 40 m (Cousteau and Diolé 1971, p. 268). The stony coral distribution and participation in reef formation received qualification, and Cuban scleractinians are continuing today to provide rich regional Caribbean information about the different zones of the coral reefs and about the upper part of the mesophotic zone, with no faunal break between shallow-water reefs and Mesophotic Coral Ecosystems (Zlatarski 1980, 1982, 2018a, b; Pyle and Copus 2019; see Chap. 14, Reed et al.). The 1970 study covered the largest part of all zones of Cuban coral reefs ever to be the object of one project investigated using the same methodology. Regrettably, for some immense reefs, like the unique reticulated Gran Banco de Buena Esperanza in the Gulf of Guacanayabo, the information from that time is the only available, because it was never subsequently the object of stony coral investigation

Fig. 1.6 Bathymetric distribution of Cuban Scleractinia

1 Insights from Cuban Coral Reefs 13

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Fig. 1.7 Diversity of genera, species, and infrasubspecific categories in depth. 1 generic diversity; 2 species diversity; 3 infrasubspecífic diversity; n number of the taxa; m depth

(Hernández-Fernández et al. 2013; Zlatarski and Gonzáles Ferrer 2017; Zlatarski and Greenstein 2020). The field work in Cuban waters was performed in maximal geographic and bathymetric ranges over a relatively short period (1970–1973), and as a result, the information produced provided a snapshot of Cuban coral reefs half a century ago. The past decades have brought considerable alteration of coral reefs with shift of zones and degradation. The 50-yearold documentation together with the physical collection itself offer a rare possibility for comparative studies and longtime ecosystem analysis. The data presented here from 1970 to 1973 from 44 profiles and 194 stations, together with the large coralla collection to a depth of 70 m and the scientific documentary, provide solid documentation of the status of reefs in the early 1970s (Zlatarski and Martínez Estalella 1980, 1982, 2018). As a result, today’s researchers have solid older information at their disposal, and revisiting the documented places would offer the knowledge that is needed to support the conservation of the deteriorating coral reefs. For example, in the 1970s, openly branching Acropora colonies with fused branches were rarely observed and always in the proximity of A. palmata or A. cervicornis. For these reasons, the species status of A. prolifera was treated with reserve and marked with a question mark (Zlatarski 1980, 1982, 2018b). Decades later, genetics proved that it is hybrid of both mentioned species (van Oppen et al. 2000). Interestingly, on a visit to Cuban waters during this century,

V. N. Zlatarski

the hybrid was observed more frequently, even in some lagoonal areas where its fragments formed the basis of buildups (Zlatarski 2010; Zlatarski et al. 2004). The opportunistic nature for potential survival of this scleractinian hybrid was proved by the fact that, since 1972, A. prolifera inhabits Gran Banco de Buena Esperanza with only one of its parents’ species, A. cervicornis, and was evidenced by the presence of more than one F1 generation and its fertility (Zlatarski and Gonzalez-Ferrer 2017; Zlatarski and Greenstein 2020). The observed proliferation of the hybrid deserves special attention, especially in the present crisis in coral reef ecosystem, and any attempt to analyze the Caribbean acroporid corals by ignoring the established hybridization (Cramer et al. 2020) presents an incomplete picture. In the early 1970s, the invasive species Tubastraea coccinea was found only on the southeast coast of Cuba (Zlatarski 1980, 1982, 2018b). Half a century later, it is present in different remote parts of the Cuban Archipelago– Canarreos Archipelago, northwest and north central coast (see Chap. 8, Gonzalez-Ferrer et al.). This urges study of the invasion process and the need for its control. In the early 1970s, Cuban reefs were documented as a hot spot of scleractinian life in the Atlantic. Only few incidences of scleractinian abnormalities were observed—the presence of corallum hyperplasia, partial discoloration of the colonies, and damselfish chimneys—but not the existence of any epizootic phenomenon (Zlatarski et al. 2004). A clear negative urbanistic anthropogenic impact was documented in front of the Havana coast (Zlatarski 1980, 1982, 2018b). During the following decades, the reefs in Cuban waters underwent noticeable negative change, but the species richness remained high. There are ongoing serious efforts for reef conservation, which naturally should start with care for the reef-building coral species. In the Caribbean, the Cuban coral reefs are the largest, and they are centrally located, and the most diversified geographically and bathymetrically and with regard to bioconstructor zonation. The Cuban archipelago is only in part victimized by considerable anthropogenic deterioration. Some reefs are in good conditions. All of these factors make the Cuban reefs crucial for saving this threatened, vitally important ecosystem in the Caribbean. What is the best way to approach this goal? Many years of rambling in search of the most appropriate researcher’s path lead to the highly Parnassian verse by Antonio Machado: Wayfarer, there is no way, you make the way as you go. (Machado 1982, p. 143)

Acknowledgments The manuscript benefited a great deal from the thoughtful constructive suggestions of the reviewer Bernhard Riegl. I also thank John Reed for the editorial improvements and Sergio González-Ferrer for his assistance with the figures. Also appreciated is

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Insights from Cuban Coral Reefs

the help of Dan Whittle, Valerie Miller, and Michael Pickard with the English translation of text previously published in Spanish.

References Battistini R, Bourouilh F, Chevalier J-P, Condray J, Denizot M, Faure G, Fisher J-C, Guilcher A, Harmelin-Vivien M, Jauber J, Laborel J, Montaggioni L, Masse J-P, Mauge LA, Peyrot-Clausad M, Pichon M, Plante R, Plaziat J-C, Blessis YB, Richard G, Salvat B, Thomassin BA, Vasseur P, Weydert P (1975) Éléments de Terminologie Réciface Indopacifíque. Téthys 7(1):1–111 Cousteau J-Y, Diolé PH (1971) La vie et la mort des coraux. Flammarion, Paris Cramer KL, Jackson JBC, Donovan MK, Greenstein BJ, Korpanty CA, Cook GM, Pandolfi JM (2020) Widespread loss of Caribbean acroporid corals was underway before coral bleaching and disease outbreaks. Sci Adv 6:eaax9395 Hernández-Fernández L, Olivera Y, González-De Zayas R, Salvat Torres H, Guimarais Bermejo M, Ventura Díaz Y, Pina-Amargós F (2013) Caracterización fisicoquímica e inventario de especies del Gran Banco de Buena Esperanza, golfo de Guacanayabo, Cuba. Rev Investig Mar 33(2):43–57 Machado A (1982) Selected poems (trans: Trueblood AS). Harvard University Press Martínez Estalella N (1980) Data of organisms associated with Scleractinia of Cuba. In: Zlatarski VN, Martínez Estalella N (eds) The scleractinians of Cuba with data of associated organisms. Editions de l’Académie bulgare des Sciences, Sofia, pp 252–275. (in Russian) Martínez Estalella N (1982) Données les organismes associés aux Scléractiniaires de Cuba. In: Zlatarski VN, Martínez Estalella N (eds) Les Scléractiniaires de Cuba avec des données sur les organismes associés. Editions de l’Académie bulgare des Sciences, Sofia, pp 401–435 Martínez Estalella N (2018) Datos sobre los organismos asociados a los escleractnios de Cuba. In: Zlatarski VN, Martínez Estalella N (eds) Los escleractinios de Cuba con datos sobre sus organismos asociados. Harte Research Institute for Gulf of Mexico Studies at Texas A&M University, Corpus Christi, pp 404–435. http://www. harteresearchinstitute.org/project/legacy-book-los-escleractinios-decuba-corals-cuba Núñez Jiménez A (ed) (1970) Atlas Nacional de Cuba. Acad. Ciencia de Cuba, Acad. Cienc. URSS, La Habana Pyle RL, Copus JM (2019) Mesophotic coral ecosystems: introduction and overview. In: Loya Y, Puglise KA, TCL B (eds) Mesophotic coral ecosystems, coral reefs of the world, vol 12. Springer, Switzerland, pp 3–27. https://doi.org/10.1007/978-3-319-92735-0_1

15 van Oppen MJH, Willis BL, van Vugt HWJA, Miller DJ (2000) Examination of species boundaries of the Acropora cervicornis group (Scleractinia, Cnidaria) using nuclear DNA sequence analysis. Mol Ecol 9:1363–1373 Zlatarski VN (1968) Paleobiology of the Urgonian Scleractinia of the Central Fore-Balkan. Dissertation, Geol. Inst., Bulg Acad Sci & Committee Geol, Sofia Zlatarski VN (1980) The scleractinians of Cuba, Chapters 1–6. In: Zlatarski VN, Martínez Estalella N (eds) The scleractinians of Cuba with data of associated organisms. Editions de l’Académie bulgare des Sciences, Sofia. (in Russian) Zlatarski VN (1982) Les Scléractiniaires de Cuba, Chapitres 1–6. In: Zlatarski VN, Martínez Estalella N (eds) Les Scléractiniaires de Cuba avec des données sur les organismes associés. Editions de l’Académie bulgare des Sciences, Sofia Zlatarski VN (2010) Palaeobiological perspectives on variability and taxonomy of scleractinian corals. Palaeoworld 19(3–4):333–339 Zlatarski VN (2018a) Investigations on mesophotic coral ecosystems in Cuba (1970–1973) and Mexico (1983–1984). CICIMAR Oceánides 33(2):27–43 Zlatarski VN (2018b) Los escleractinios de Cuba, Capitulos 1-6. In: Zlatarski VN, Martínez Estalella N (eds) Los escleractinios de Cuba con datos sobre sus organismos asociados. Harte Research Institute for Gulf of Mexico Studies at Texas A&M University, Corpus Christi. http://www.harteresearchinstitute.org/project/legacy-booklos-escleractinios-de-cuba-corals-cuba Zlatarski VN, Gonzáles Ferrer S (2017) Gran Banco de Buena Esperanza: unique Caribbean coral reef system. Reef Encounter 32(1):60–62 Zlatarski VN, Greenstein BJ (2020) The reticulate coral reef system in Golfo de Guacanayabo, SE Cuba. Coral Reefs 39(3):509–513. https://doi.org/10.1007/s00338-020-01933-7 Zlatarski VN, Martínez Estalella N (1980) The scleractinians of Cuba with data of associated organisms. Editions de l’Académie bulgare des Sciences, Sofia. (in Russian) Zlatarski VN, Martínez Estalella N (1982) Les Scléractiniaires de Cuba avec des données sur les organismes associés. Editions de l’Académie bulgare des Sciences, Sofia Zlatarski VN, Martínez Estalella N (2018) Los escleractinios de Cuba con datos sobre sus organismos asociados. Harte Research Institute for Gulf of Mexico Studies at Texas A&M University, Corpus Christi. http://www.harteresearchinstitute.org/project/legacy-booklos-escleractinios-de-cuba-corals-cuba Zlatarski VN, Alcolado PM, González Ferrer S, Kramer P (2004) Archipiélago Jardines de la Reina, Cuba. Thirty years later, species richness of scleractinian corals remains high. Reef Encounter 32:30– 32

Part II History

2

Research History of Corals and Coral Reefs in Cuba Sergio González-Ferrer

Abstract

This chapter summarizes the history of the scientific knowledge regarding Cuban coral reefs. The work of Antonio Parra, published in 1787, is the first book to refer to species of reef life in Cuba. In the nineteenth century, the works of Alexander von Humboldt, Charles Darwin, Ramón de la Sagra, Felipe Poey, and Rafael Arango are the most prominent, along with the debates on the origins of reefs by William O. Crosby and Alexander Agassiz. During the twentieth century, William M. Davis and William Smith considered the reefs on the southern coasts of Cuba to be the longest in the West Indies. Also in this century, multiple institutions devoted to marine studies emerged and research grew exponentially. The monograph by Zlatarski and Martínez Estalella, published in 1980, demonstrated the variety of the Cuban scleractinian and the presence of intermediate phenotypes of acknowledged species. Information is presented concerning “true” coral barrier reefs in Cuba, the contribution of projects such as AGRRA, the early warning voluntary monitoring in coral reefs, the studies of the mesophotic coral ecosystems, and the economic value of the Cuban coral reefs, with references to the most notable publications in the last 30 years. Keywords

Coral reefs · History of coral reef science in Cuba · CIM · IDO · CITMA

S. González-Ferrer (✉) La Habana, Cuba

2.1

First Traces of Interest in Corals and Coral Reefs

Great naturalists and worthy scientists have left their mark in the research of Cuban coral reefs. This chapter provides a summary of the main moments in the the discovery of knowledge on these ecosystems in the largest island of the Antilles. Reference is made to the publications of some of the different zoological groups that form these reef systems; however, according to Alcolado et al. (2003), “stony coral” and “coral reef” are terms necessarily and closely linked, at least when writing about their recent history. We start off with traces of the first human settlements on the Isle of Cuba, where the coral reefs in some coastal areas represented good sites for fishing, harvesting, and sheltering from the actions of the waves. If we want to find the beginnings of interest in these ecosystems in Cuba or point to their discoverers, we must go back to the pre-agro-pottery (protoarchic) and agro-pottery groups who inhabited the coast of Cuba since 6000 years BC. Perhaps, the only proof of this are the anthropomorphic figures, seemingly of a religious nature, funerary objects, or possible mortars, used to macerate foods or ritual substances, prepared by those cultures from stony corals (González-Ferrer 2004a, Fig. 2.1).

2.2

The Eighteenth Century: The “Interesting Stones” of Don Antonio Parra

In the eighteenth century, we find the first evidences of scientific curiosity regarding some of the organisms belonging to coral reef ecosystems. Cuba had only around 171,620 inhabitants (ONEI 2019) which begin to open up to the Enlightenment, a philosophical, artistic and scientific movement coming from Europe. According to García Sanchez (2011), this movement considered reason as the only force capable of ensuring progress, and, at the same time, its

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_2

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Fig. 2.1 Idols sculptured on stony coral by natives in the Cuban isle. These pieces can currently be found in the Museo Antropológico de la Universidad de La Habana (Anthropological Museum of the University of Havana)

defenders felt a great love for nature and a preference of useful things over beautiful ones. In Garcia’s (1859) opinion, it is with the establishment of the Real y Pontificia Universidad de San Gerónimo de La Habana in 1728 and the Colegio de la Compañía de Jesús around the same time that the Cuban Enlightenment had its beginnings. It is in this atmosphere of thought and with the end of the war of 1763 between Spain and England that Don Antonio Parra Callado, of Portuguese origin, arrived in Cuba as a simple soldier of the Mallorca infantry regiment at the order of King Carlos III. He soon retires from the army and forms a family, while his interest for natural history leads him to gather collections, especially of “marine productions” (García 2016). The publication Descripción de Diferentes Piezas de Historia Natural las mas del Ramo Maritimo, Representadas en Setenta y Cinco Laminas (Description of Different Pieces of Natural History most of them of the Marine Branch, Represented in 75 Pictures) by Parra (1787) constitutes the catalog of his collection (Fig. 2.2). This book was published by the Imprenta de la Capitanía General (General Captaincy Press) in Havana and is the first in Cuba to be illustrated with engravings of the collections made by hand by the author’s son on copper etching, with a layer of wax over the metal (Saco 1858a; García 2016). Those collections came from the coasts, fishing nets, and limestone quarries from which material for colonial constructions were extracted, as well as from the Escambray Sierra. Parra groups in this collection and in his later work 71 species of fish and 25 crustaceans, many of them of reef life, besides reptiles and a considerable number of species of echinoderm, sponge, gorgonian, algae, polychaete, as well as stony coral, which he calls “interesting stones” (GonzálezFerrer 2004a, Fig. 2.3).

Fig. 2.2 Cover of Don Antonio Parra Callado’s book (1787)

All of these exhibits, identified by their popular names, constituted the first Science Cabinet of Havana (Presas 1866) which, because of its frequent public exhibition, could be considered the main predecessor to the natural science museums of Cuba. Parra’s collection later traveled to Spain to form part of the Royal Cabinet of Natural History, for which King Carlos IV assigned him a tidy annual pension (Parra 1799). Some of the specimens of this collection are still kept in the Museo Nacional de Ciencias Naturales de Madrid and maintain their “original” shape and color (García 2016). Years later, several scholars set themselves to classifying Parra’s exhibits, a work that can be considered among the first classification of Cuban reef-life organisms. Thus, the Spanish geographer and naturalist Don José Cornide, one of the better-known members of the Enlightenment in Galicia, made a classification of several of the fishes, which was later amended by Saco (1858a). Similarly, the French sages Georges Cuvier and Achille Valenciennes, in their publication Histoire Générale et Particulière des Poissons, assign scientific names to Parra’s common names (Jimeno 1858), not without some errors corrected later by the Cuban scholar

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Research History of Corals and Coral Reefs in Cuba

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Fig. 2.3 Some “interesting stones” published by Parra (1787)

Felipe Poey y Aloy in his work Enumeración de los Peces Descritos y Figurados por Parra (List of the Fishes Described and Illustrated by Parra), published in the Proceedings of the Philadelphia Academy in 1863 (García 2016). Presas (1866) tells us how French zoologist Henri Milne-Edwards used Parra’s data of crustaceans in his publication Historia Crustacea of 1837. In addition, the Frenchman Guérin-Méneville (1856), the great German scientist resident in Cuba Juan Gundlach in 1917 (García 2016), and more recently Sanchez-Valero et al. (2009) classified some echinoderms, one annelid, one bryozoan, some sponges, and two corals. However, among these latter “interesting stones,” it is not difficult to distinguish Parra’s drawings at least five genus (González-Ferrer 2004a). Parra’s book is undoubtedly a scientific relic (Fernández 1943) and contains the first noteworthy approximation to

Cuban coral reefs, although as Parra himself said concerning his collections “serving as a sample, they will make the lion known by its nail.”

2.3

The Nineteenth Century: Beginnings of Marine Sciences in Cuba

The nineteenth century begins with the visits to Cuba by Alexander von Humboldt in the years 1800 and 1804. This German explorer and scientist is considered by many as the second discoverer of the Americas, given his extensive and varied work in these lands and because, according to Acosta (2005), he did not have the biased colonialist view of many European scholars. In his work Political Essay on the Island of Cuba published in 1826 (Fig. 2.4a, b), he refers to reefs

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Fig. 2.4 (a) Alexander von Humboldt, painted by Joseph K. Stieler in 1843 (Wikipedia 2021); (b) cover of his book Political Essay of the Island of Cuba, published in 1826

and reef obstacles to navigation. In his Chart of the Island of Cuba of 1820, it is possible to see mapped many currently known reef areas (Fig. 2.5) with greater accuracy than other previously made maps.

Humboldt acknowledged coral accretion in stating “When sitting near Havana at the foot of the Castillo de la Punta (Castle of the Point) on banks of cavernous rocks carpeted both by ulvas that turn green and live octopuses, one can see

Fig. 2.5 Geographic-topographic map of the Isle of Cuba prepared according to Alexander von Humboldt’s notes by the Royal Corps of military geographers of France (Humboldt 1827)

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Research History of Corals and Coral Reefs in Cuba

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Fig. 2.6 Distribution of the different kinds of coral reefs, considered by Darwin (1842)

enormous masses of madrepores and other lithophyte corals squeezed into the fabric of these banks, it is reasonable to admit that all of this limestone rocks, which most of the island of Cuba consists of, is the effect of an uninterrupted operation of nature, of the action of organic forces and of partial destructions, and which continues to our days in the ocean bottom . . .” and goes on “These same coral rocks of the Castillo de la Punta are again found inland in the high mountains, along with petrification of bivalve sea shells very different from those currently found in the coasts of the Antilles” (Humboldt 1827). Another interesting publication in these years, also among the first to classify Cuban reef-life organisms, can be found in the Encyclopédie méthodique - Histoire naturelle des zoophytes, ou animaux rayonées (Lamouroux et al. 1824). In its reference is made to the sponge Spongia robergii on Cuban coasts; to the coelenterates Antipathes ligulata, Pocillopora damicornis, Muricea elongata, Plexaura flexuosa; and the macroalgae Acetabularia crenulata, identified as an animal, whose registry is considered the first mention of a Cuban macroalgae (Suárez et al. 2015). In Volume IX of 1836 of the publication The Magazin of Natural History, one reads “It is not probable that many of your readers are acquainted with the northeastern portion of the island from whence I write (Cuba); for I believe that very little has been known or written on this district, and that, possibly, it has never been visited by men of scientific research.” Thus wrote R.C. Taylor, member of the Pennsylvania Geological Society, as part of the introduction to his description of “The reef stretches eastward, parallel with the shore, for several miles, from the entrance of the fine bay of Gibara (. . .)” (Taylor 1836). Taylor writes his personal experiences while at the same time gathering very summarized information of his observations of the current

emerged reefs in that area, as well as of the mangrove swamps and lagoons. By his notes it is known that he gathered illustrative samples of the solidification process of conchiferous limestone; concerning high reefs he considered that “it would appear that this inner reef was produced under different circumstances of relative elevation of sea and land; indicating a slight depression in the one case, or an elevation in the other.” He also makes note of the presence of species of corals, sponges, mollusks, crustaceans, fishes, and birds, and of the capture and collection of marine species by means of freediving. His observations on reef accretion are also noteworthy “Within the reef new colonies are rearing slowly their habitations, and miniature reefs arise around. But here they appear to have nearly completed their labors, and nature has set limits to their combined operations. They have almost brought their work to that elevation beyond which they are unable to proceed, and henceforward the structure must receive its increase in its breadth alone.” In 1842, the great English naturalist and geologist Charles Darwin published his theories on the structure and distribution of coral reefs with a later second edition in 1874 (Darwin 1842, 1874). This book contains the first geographical notion of coral reefs at a global scale (Fig. 2.6). The information relating to Cuba was compiled by Darwin based on the analysis of nautical charts of the times, on the notes by Humbold and Taylor, as well as on the maps of the ports prepared by Captain Owen. Darwin, referring to the reefs of the Sabana-Camagüey (north central of the island), puts Cuba as an example with a reef “(. . .) which extends parallel to the shore at a distance of between one half and one third of a mile, and encloses a space of shallow water, with a sandy bottom and tufts of coral,” which supports his theory that “on coasts where the sea deepens very suddenly the reefs are much narrower, and their limited extension seems

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evidently to depend on the high inclination of the submarine slope; a relation which, as we have seen, does not exist in reefs of the barrier class.” He considered that this geographic barrier that borders the northern coast of Cuba was, like the banks of the Great Bahamas and Cay Sal, the result of the accumulation of sediments generated by an upward movement of the waters, with coral formations only on the external edge. Referring to the northwestern tip, he stated that Los Colorados were similar to the banks of the southern tip of Florida, with a lagoon with a depth of from two to three fathoms which increased to twelve fathoms in the southern part, where he considered were no coral islets or reefs. On the southern coast he only acknowledges reefs close to the Isle of Pines and makes notes that he was unable to obtain a detailed description of the large group of banks and “keys” more to the east on the southern part of Cuba, “(. . .) within them there is a large expanse, with a muddy bottom, from eight to twelve fathoms deep: although some parts on this line of coast are represented in the general charts of the West Indies, as fringed, I have not thought it prudent to color them. The remaining portion of the south coast of Cuba appears to be without coral-reefs” (Fig. 2.7a, b). Finally, he concludes that on the coasts of Cuba, as well as the rest of the West Indies and the Gulf of Mexico, the reefs were only coastal ones (also known as fringing reefs). In the mid-nineteenth century, when the largest island of the Antilles reached 1 million population, the work Historia Física, Política y Natural de la Isla de Cuba (Physical, Political and Natural History of Cuba) (1838–1857) was published. It comprised 13 volumes compiled by the Spanish naturalist and economist Ramón de la Sagra, can be considered of great value in its time, although the author ignored the contribution that Cuban naturalists of the stature of Felipe Poey y Aloy (Saco 1858b). Thus, the identification of marine species, was in the hands of important French naturalists (Guichenot identified the fish (Guichenot 1843), D’Orbigny the mollusks (D’Orbigny 1845) and foraminifera (D’Orbigny 1840), Guérin-Méneville the crustaceans and Montagne the algae). This work speaks of coasts covered with corals, coral islands and banks; the latter described as “buildings raised Fig. 2.7 (a) Charles Darwin at Down House, his house in Kent, around 1880 (NGH 2021); (b) detail of the distribution of coral reefs considered by Darwin for Cuba and other nearby islands, with the names edited for their better presentation at this scale

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from the bottom of the sea to the surface of the waters by numerous animals,” thus connecting some of the species described to the coral reefs. In Volume V of 1845 (Fig. 2.8), D’Orbigny defines the coral reefs as “gigantic works of the smallest and most imperfect beings.” In that same period, some Cuban geography books place emphasis on considerable reef formations, as is the case with Belasco and Echeverría (1842), where reference is made to “Los Colorados and the dangerous reefs of Santa Isabel, masses of which form an almost uninterrupted chain of reefs (. . .).” Also, the book by Pichardo (1854), in whose preliminary explanations the author discusses the use of the words bank and reef, as synonyms of bottom, connected with reef formations, and further on, in his descriptions he talks about reefs, reef chains, coastal reefs and channels formed by reefs, although he also refers to coral formations that surface or whose nearness to the surface can hinder navigation. Section 2.5, about animals, calls one’s attention to, for example, the magpie shell (Cittarium pica), “(...) whose base is stuck to the marine reefs” or “rocks (. . .) carpeted with live octopuses and enormous masses of madrepores and other lithophyte corals squeezed into them,” which is considered “proof of the superimposition and growth of modern tertiary ones, deducting from what is seen that an equal operation is executed in the same rocks of the sea bottom repeated until raising those keys above the surface of the waters (. . .).” Similarly, we find some geographic references to some Cuban reefs in the Crónicas de las Antillas (Chronicles of the Antilles) by Don Jacobo de la Pezuela, where he also makes comments about the oscillatory movements of the coralliferous limestone as proof of the elevation in recent periods of some stretches of the coast, stating that “The Isle of Cuba can be considered as one of the classical countries of zoophytes” (Pezuela 1871). Also, in the second half of the nineteenth century, some publications continued the studies of current coral reefs, dealing with classification of species or the study of fossil registries. Felipe Poey, one of the most important Cuban naturalists of this period, stands out when he refers to the habitat of some species of fishes described in his Repertorio

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Research History of Corals and Coral Reefs in Cuba

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Fig. 2.9 Rafael Arango y Molina (EcuRed 2021) published the first taxonomic reference to many marine invertebrates of Cuba

Fig. 2.8 Cover of Volume V of Historia Física, Política y Natural de la Isla de Cuba (de la Sagra 1845)

Físico-Natural de la Isla de Cuba (Physical-Natural Repertoire of the Isle of Cuba) of 1865–1866 or in his masterpiece, Ictiología Cubana (Cuban Ichthyology), presented manuscript in Amsterdam in 1883 (Poey and Aloy 2000). Although it was not yet possible for him to perceive the magnitude of the reefs as an ecosystem, he was able to point out some species that “do not stay away from the reefs” or “live in the caves along the reefs from where it goes out to seek its food.” References to stony corals of Cuba and emerged reefs are also in the works of Milne Edwards and Haime (1848–1951), Duchassaing and Michelotti (1861), Duncan (1876), and Brüggemann (1877). Poey created the Museo de Historia Natural de la Universidad de La Habana in 1842, which in 1913 received the name of its founder to this day (Suárez et al. 2020). It is not difficult to presume that many collections from the coral reefs where first exhibited in this museum and later in the Museo de Historia Natural y Anatomía Patológica (Museum of Natural History

and Pathological Anatomy) founded in 1874 by the Academy of Medical, Physical, and Natural Sciences of Havana. In 1877, in the science journal Anales de la Real Academia de Ciencias Médicas, Físicas y Naturales (Annals of the Royal Academy of Medical, Physical, and Natural Sciences of Havana), Rafael Arango y Molina, a Cuban scientist celebrated for his contribution to the knowledge of land and river mollusks of Cuba, published Radiados de la Isla de Cuba (Radiata of the Isle of Cuba) (Fig. 2.9), where he presents information on 30 species of echinoderms and 66 stony corals. This material is the first publication about the taxonomy of these Cuban invertebrates. Part of the species named there are from collections made by Felipe Poey and by Arango himself, received from fishermen or found by the shores. These samples were identified by the prestigious Swiss taxonomist and naturalist Sr. Count L. F. Pourtalès and were added to the species found by him and also Swiss Louis Agassiz, in the dredges carried out in La Chorrera and Cojímar during 1867 and 1870 (González-Ferrer 2004a). In this work, Arango acknowledges Pourtalés as the first person to make known the “polyps of Cuba” (Arango 1877). An important moment arises from the discussions concerning the origin of the coral reefs of the Cuban coasts at the end of the nineteenth century. William Otis Crosby stands out for having suggested in 1884 that the coastal inlets of Cuba “are not the work of the sea, but rivers in a period in which the land was higher than now,” due to which he called the harbors “half-drowned valleys,” and because he was the first to acknowledge the discordant contact of the reefs with their main rocks (Davis 1928). This American geologist and engineer describes the dimensions of emerged reef terraces and observes that the reefs reach heights of up to 2000 feet, with clear evidence of having been built by stony corals

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(Crosby 1884). This leads us to conclude that their origin could not have been in deep waters as, according to him, was to be supposed from the theories of the eminent zoologist Alexander Agassiz. Crosby also supposed, taking into account the breadth of the Cuban emerged reefs, that they could not have been formed during a progressive rise of the land layer, but their formation was rather due to a slow subsidence, thus supporting the theory put forward by Darwin (1842). He also considered the Yunque de Baracoa peak (literally the Baracoa Anvil due to its shape) in the province of Guantánamo, “a monument” of a subsidence of up to 2000 feet, which must have reduced the Greater Antilles to a few lines of small islands. Alexander Agassiz (Fig. 2.10) measured a large part of the Cuban coasts in 1893, in which he also studied fossil corals, emerged marine terraces, and he furthermore described, seemingly for the first time, the geographic trajectory of many current coral reefs of our coastal shelf. He comments, “Outward of the high coral reef, which may be said to border almost all of the coast of Cuba, we find the growing reef, which extends to ten or twelve fathoms or more wherever there is an insular shelf of any breadth between the high reef and the 100-fathom line” (Agassiz 1894). Thus, in his tour, he speaks of beaches of “coral sands derived from the live peripheral reefs,” of fringe or coastal reefs, coral patches, and banks surrounded by reefs and reef chains. Agassiz is the first

Fig. 2.10 Alexander Agassiz in 1896 (Cortés 2009)

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we have reference of to have called our attention to the coral barrier of Cuba, when he describes a “formidable coral barrier” referring to the Canarreos Archipelago and terms the Colorados Archipelago “the great coral barrier.” Further on, in his section Algunas Opiniones Recientes Sobre la Teoría de la Formación de los Arrecifes de Coral (Some Recent Opinions on the Theory of the Formation of Coral Reefs), based on his explorations in The Bahamas, Florida, Cuba, and the Windward Islands, Agassiz defends his hypothesis, stating that the formation of reefs has not always been influenced by the subsidence of the sea bottom. He also comments on the “limestone barriers which form the basement of the coral reefs of Florida, Cuba, and other reefs of the Antilles (Mexico and the Caribbean)” unlike the sea bottoms that bear the reef constructions of other latitudes.

2.4

The Twentieth Century: Birth of the Marine Research Centers, Protected Areas, and First Collaboration Agreements

Linked to the University of Havana, the Sociedad Cubana de Historia Natural (Cuban Society of Natural History) was founded in 1913. Its records, under the supervision of the zoologist and anthropologist Don Carlos de la Torre, also include some works that make reference to coral reefs. Thus, we find in his Volume II, where they speak about the Oak Alligator (Roig 1917) that “grows on the reefs of the Sierra de Guanes,” making references to emerged reefs, or in another study on the sea urchin (Delgado 1917), referring to Diadema setosum (likely Diadema antillarum) as “very common in coral reefs.” It is in Volume IV of these reports (Fig. 2.11) where the first reference to a work dedicated to the current coral reefs of Cuba is found: Antonio Pastor Giraud’s Ph.D. thesis, of which the Cuban anthropologist, naturalist, and physician Arístides Mestre considers “undoubtedly deserves special mention” (Mestre 1919). During the first half of the twentieth century, there are several studies which mention Cuban reefs, such as Vaughan (1919, 1933), Sánchez-Roig (1928), Boone (1928, 1933), Wells (1934, 1941), and Rivero and Bermúdez (1938). William Morris Davis, responsible for the concept of the “cycle of erosion,” refers to the reefs of Cuba in his study The Coral Reef Problem (1928), where he considers that the reef barriers of Cuba are the best developed in the West Indies, and he believes them to be real barriers “in the sense of skirting deep waters.” Davis, studied the nautical charts and raised the existence of a long coral reef on the north coast, included in the area of the Sabana-Camagüey Archipelago, with a lagoon of “a few fathoms” and joined by a coastal reef to another barrier section in the Los Colorados Archipelago. On the south coast, he takes into account the

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Research History of Corals and Coral Reefs in Cuba

Fig. 2.11 Cover of Volume IV of the Reports of the Cuban Society of Natural History (de la Torre 1922), where the Ph.D. thesis of Antonio Pastor Giraud is mentioned, the first such work devoted to coral reefs in Cuba

largest barrier in the West Indies, with a discontinuous development and a length of 150 miles, referring to a section included in the area of the Jardines de la Reina. Similarly, Walton Smith in 1948 considers the Los Colorados reef, from Cape San Antonio to Bahia Honda, to be a barrier reef “more closely allied to reefs of the true coral seas” and considers the reefs of the south, from the Isle of Pines to Cape Cruz, the longest in the West Indies (Walton-Smith 1948). In 1949, Pedro Pablo Duarte Bello defended his doctoral thesis at the University of Havana with the work Contribution to the Study of the Madrepores of the Coasts of Cuba (Duarte-Bello 1949). In this work, 44 species of Cuban corals are presented, and topics such as their physiology, reproduction, ecology, and distribution are addressed. Unfortunately, this work was not published as a scientific work, but only

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informally in 1960 (republished in 1963), as the second number of the educational series of the Aquarium and the University of Havana. This can be considered the first illustrated publication, showing the stony corals that inhabit the waters of Cuba. According to Suárez et al. (2020), in the 1950s, people began to think about the practical application of marine research, and thus, in 1952, the Fisheries Research Center (CIP) was founded, west of Havana, sponsored by the Agricultural and Industrial Development Bank, which later in 1955 was forced to close due to lack of official support. Around this time, the Universidad Católica de Villanueva opened a marine laboratory under the influence of the Institute of Marine Sciences of the University of Miami, while the University of Oriente, in Santiago de Cuba, created a marine laboratory managed by Manuel Díaz Piferrer (Suárez et al. 2020). With the triumph of the Cuban Revolution in 1959, a period of creation of facilities for marine research began throughout the island, which starts with the reopening of the CIP and the inauguration of the National Aquarium of Cuba (ANC) in 1960. In 1963, Duarte Bello, who became an ichthyologist at the ANC, published the book Corales de los Arrecifes Cubanos (Corals of Cuban Reefs) (Fig. 2.12), the first popular work on this topic, which includes photos and descriptions of the most common coral species of the northern reefs of Pinar del Río and Havana, accompanied by their scientific names and names from popular jargon or designed by the author for a better understanding. In these years, new equipment that would allow autonomous diving began to be available to researchers, opening a new stage in marine research in which divers and specialists begin more direct observations of coral reefs. In 1965, the Institute of Oceanology (IDO), currently Institute of Marine Sciences (ICIMAR), of the Academy of Sciences of Cuba was born (Claro 2010) and in 1970 the Center for Marine Research (CIM) of the University of Havana. These two centers, together with the ANC, have been the leaders in research related to coral reefs, as well as in the training of specialists in this field, from its emergence to the present. In 1968, the first numbers of the IDO magazine, called Serie Oceanológica, began to be published; and later, in 1972, the Science magazine (Series 8) published the first issue of the Marine Research series, which, according to Suárez et al. (2020), constituted the genesis of the CIM’s Journal of Marine Research in 1980. In these journals, you can find innumerable works over 50 years on Cuban coral reefs. Also in 1980, the law for theprotection of the environment and the rational use of natural resources was approved, which represents a significant legal framework for the protection of coral reefs (see Chap. 12, Duran et al.). In this period, we also have some references to Cuban scleractinian species and coral reefs in the works of the

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Fig. 2.12 Cover of the book Corales de Los Arrecifes Cubanos (Duarte-Bello 1963)

eminent Soviet geologists Zenkovich (1965, 1969, 1972), Ionin (1966, 1967, 1975a, b), Ionin et al. (1972a–d, 1977), and Zenkovich and Ionin (1969), who provided invaluable information about the origin, age, and characteristics of the low littoral terraces and the coral reefs and keys located behind them. Other Soviet scientists published on the subject (Bogdanov 1968; Ionin and Pavlidis 1968, 1971a, b), as well as the Czech researchers Seneš (1966), Balon and Seneš (1967, 1971), Kukal and Naprstek (1967), Ivan (1967), and popular publications by Anonymous (1967), Gruner (1967), Wagner (1967a–d), Richter (1967), Kramer (1967, 1968), Kaden (1968), and Taege and Wagner (1975). Other research projects, oriented toward the study of the biocenosis of the coral reefs of Cuba, were carried out in IDO expeditions in collaboration with the Humboldt University of Berlin, such as the ones led by Kühlmann (1970a–c, 1971a– e, 1974a–c) which addressed the morphology, origin, ecology, as well as the description of complex reef systems (barrier reefs, crossbar reefs, cluster reefs, complex reefs,

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coral canyons, incision reefs, spur and groove reefs, and horst reefs) based on the substrate and other environmental factors (Kühlmann 1974a). We also find in the study of Kühlmann (1974a, b) the first news of the mesophotic reefs of Cuba (see Chap. 15, Reed et al.). It is also in this period that some media began to spread as news that Cuba had the second largest coral reef in the world, in the area of the Sabana-Camaguey Archipelago, with a length of more than 400 km, only surpassed by the Great Barrier Reef in Australia, information that, according to Del Cueto (1974), came from school geographic sources. One of the most exuberant and scrupulous books in the history of the knowledge of stony corals and coral reefs also saw the light in 1980 with the name Los escleractinios de Cuba (The Sclaractinians of Cuba) (Fig. 2.13). This arose from a collaboration between the Bulgarian Academy of Sciences and the IDO, in the hands of the Bulgarian geologist Vassil Zlatarski and the Cuban biologist Nereida Martínez Estalella (Zlatarski and Martínez Estalella 1980, 1982, 2018). For its preparation, the scleractinians and reefs from 194 sites, along the south and northwestern coasts, were studied. This study included the collection of 5924 samples from the seabed up to 70 m deep together with direct observations of reefs as deep as 90 m. The collected samples were thoroughly described, which included 436 thin sections of skeletal microscopic structures and descriptions of the accompanying organisms. In addition, they studied 1230 fossil samples corresponding to 96 localities, of which 22 thin sections and 38 polished sections were prepared for the study of skeletal microscopic structures. This material is notable for the quality of the descriptions, accompanied by excellent photographic presentations of the skeletal characteristics of the corals, which demonstrated the variability of Cuban scleractinians and the presence of some intermediate phenotypes of the recognized species (see Chap. 1, Zlatarski). There is abundant information on coral distributions in the reef profiles which, in addition, broadened knowledge about the mesophotic zone (see Chap. 15, Reed et al.). In addition to the book, this study left a vast collection of stony corals, unmatched in the Atlantic, currently in the ANC (Fig. 2.14), which was digitized in multimedia form (Anonymous 2009), and a 37 min film with many details of the research (https://youtu.be/DMa-82bIwU). In addition, this book acknowledged three types of coral reefs in Cuba: fringe reefs, barrier reefs, and a lagoonal coral reef system on muddy bottom (see Chap. 1, Zlatarski). The latter, until now, is the only one of its kind in the world. It is located in the Gulf of Guacanayabo (SE Cuba) and forms the Great Bank of Good Hope. Even today, there remain many questions about the origin and geomorphology of these ecosystems (Zlatarski and González-Ferrer 2017). To Vassil Zlatarski, we owe the effort of more than 50 years calling international attention to Cuban corals and coral reefs (Fig. 2.15a), which

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Research History of Corals and Coral Reefs in Cuba

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Fig. 2.13 Cover of the book Los escleractinios de Cuba in its thirrd edition in Spanish (Zlatarski and Martínez Estalella 2018)

is why he is considered an illustrious Cuban and dean of the new generations of scholars of these invertebrates in the island (González-Ferrer 2004a). The contribution of the renowned photographer, Alberto Korda, throughout the project was also very valuable, since his photographs documented data collection in the field (Fig. 2.15b) and every living colony just before it was withdrawn from the ocean bottom. This photographic work helped build the

Fig. 2.14 (a) Nereida MartínezEstallela next to the coral collection in the Institute of Oceanology, towards the end of last century (Zlatarski 2004); (b) view of the condition of said collection in 2005 after its arrival at the National Aquarium of Cuba

catalog which, although never published, was finished in 1981 (Zlatarski 2004). Also in the 1980s, the great Cuban geographer and explorer Antonio Núñez Jiménez estimated that there were around 3966 km of reefs in Cuba, with 2150 km along the north coast and 1816 km along the south coast (Núñez Jiménez 1984a). Referring to possible reefs that have become atolls, he comments “In the extreme north-west of the Gulf of

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Fig. 2.15 (a) Vassil Zlatarski during the collection of corals used in the publication of the book Los escleractinios de Cuba in 1980 (González-Ferrer 2004a); (b) photo of Zlatarski, taken and signed by Alberto Korda on Jan. 19, 1973 and dedicated to Zlatarski’s mother with the following: “Dear Mom: As a memory of who shared with your son this beautiful stage of his life, I want to make you this present”

Cazones an annular coral reef rises, similar to a submerged atoll, 13 km in diameter, around which the Diego Pérez and the Macío keys extend, also the southern coast of the Zapata Peninsula, as well as the Cobo, Cancamán, Palomo and Calvario keys.” He also refers to the Paloma Cave on the south coast, 1.5 km south-southwest of the key with the same name, that it is “an underwater bank surrounded by a circular atoll with a double coral ring.” He also summarizes “After our excursion through the marvelous underwater world, let us now see, in tight synthesis, the coral formations that surround the Cuban Archipelago and form barrier reefs, living and fossil coastal reefs, ring reefs and coral banks (. . .) that constitute one of the most extensive coral reefs in the world” (Núñez Jiménez 1984b). Zlatarski and Martínez Estalella (1980, 1982, 2018), Núñez Jiménez (1984a, b), Martínez Estalella (1986), and Wells (1988) all reported that the Los Colorados reef is a barrier reef. They also pointed to another one on the outer edge of Sabana-Camagüey island group and two to the south in the Jardines de la Reina and Los Canarreos island groups. Wells (1988) estimated the Jardines de La Reina coral reef extends along 400 km and considered the existence of another barrier to the west of the Isle of Youth, between the San Felipe and Los Indios keys. She also refers to other formations such as the “crossbar reefs” similar to bank reefs but that extend in front of the bays and the “cluster reefs” that are a form of patch reefs found in small prominences on a flat substrate and that can form miniature atolls. Núñez Jiménez (1984a) also considered that as a whole the northern reefs constituted a great coral reef, whose 2150 km made it “the second great coral barrier of the world.” From the institutional point of view, in 1985, the National Enterprise for the Protection of Flora and Fauna under the Ministry of Agriculture was created. Its contribution to research and monitoring of coral reef ecosystems is much more recent, as in the Los Colorados marine protected area, currently in the approval process. Then, in the 1990s of the

twentieth century, another important moment began in the birth of provincial environmental service centers, currently categorized as research centers, with economic and legal independence. These have made it possible to gradually aid specialists interested in coral reefs and to increase scientific studies on reefs, as well as to increase education of populations in coastal areas and the staff of tourist and diving centers. Among common objectives are the following: to offer environmental scientific and technical services that the sustainable management of the marine resources of each territory may demand, as specialized guidance for recreational purposes, as well as for the execution of projects of the national science and technological innovation system, or of those from international collaboration on environmental topics. Thus, there is, attached to the Ministry of Science, Technology, and the Environment (CITMA), the Coastal Ecosystems Research Center (CIEC) in the province of Ciego de Ávila, founded in 1991, which stands out for its many research projects in the coral reefs of the SabanaCamagüey and Jardines de la Reina archipelagos; the Eastern Center for Ecosystems and Biodiversity (BIOECO), established in 1993 in the Santiago de Cuba province; the Alejandro de Humboldt National Park in the provinces of Holguín-Guantánamo, established in 1996 and a world heritage site since 2001; the Center of Environmental Research in Camagüey (CIMAC) since 1997; the Center for Environmental Studies of the province of Cienfuegos (CEAC) and the Center for Research and Environmental Services (CISAT) of Holguín, both of which were created in 1999; the Ecovida Environmental Services Center, which also sees the light in 1999 in the westernmost province of Pinar del Rio, and where the Guanahacabibes National Park and the Environmental Analysis and Monitoring Station of Sandino have been pillars of many of the most recent research and monitoring projects; and last but not least, the Center of Environmental and Marine Studies (CESAM), the youngest of these, inaugurated

2

Research History of Corals and Coral Reefs in Cuba

in the province of Villa Clara in 2001. Importantly, the National Center for Protected Areas (CNAP), founded in 1995, was also recently categorized as research center. CNAP has been foundational in working with many partners to assemble a well-designed system of MPAs, often with a focus on the protection of coral reefs or systems with associated biotic connectivity such as mangroves and seagrasses. Among the many publications referring to the reef environment in Cuba in the second half of the twentieth century, we have Zlatarski (1981), Franco-Alvarez (1983), FrancoAlvarez et al. (1981, 1983a, b), Socorro et al. (1986), and Rodríguez et al. (1989) on reef geology and fossil corals; Miravet et al. (1999) on coral reef microbiological indicators; Suárez (1989) on macrophytobenthos; Trelles et al. (1997) on reef macroalgae communities; Zlatarski et al. (1973), Zlatarski (1975), Martínez-Estalella (1986), Herrera-Moreno and Martínez-Estalella (1987), Martínez-Estalella and Herrera-Moreno (1989), and Herrera et al. (1991) on scleractinian communities; Alcolado (1978, 1979, 1985, 1989, 1990, 1994, 1999a), Alcolado and Herrera-Moreno (1987), and Alcolado and Gotera (1985) on reef sponge communities; Guitart-Manday (1959), Behety (1975), Alcolado (1981), Alcolado et al. (1980), Herrera-Moreno and Alcolado (1983, 1985, 1986a, b, 1988), García-Parrado and Alcolado (1996, 1998), and Herrera-Moreno et al. (1997) on reef octocoral communities; Martínez-Iglesias and GarcíaRaso (1999) on crustacean reef communities; Espinosa (1992), Espinosa and Rams (1987), Espinosa et al. (1994) and Ferrer & Alcolado (1994) on mollusk reef communities; Mochek and Valdés-Muñoz (1983), Valdés-Muñoz and Garrido (1987), Bosh Méndez et al. (1988), García Coll et al. (1988), Finalé-Gómez et al. (1989), Claro et al. (1990, 1994), Claro and García-Arteaga (1994), Aguilar et al. (1997), González-Sansón et al. (1997a, b), and Claro et al. (1998) on reef fish communities; Alcolado (1984, 1992), Herrera-Moreno and Alcolado (1985, 1986a, b), HerreraMoreno (1990, 1991), and Alcolado et al. (1994) on coral reef biomonitors; Ibarzábal (1993) on reef vagile macrofauna; Martínez-Estalella (1984), de la Guardia and González-Sansón (1997a–c), González-Sansón et al. (1997c), and González-Díaz (1999) on reef fauna symbiotic and coral reef associations; Herrera-Moreno (1983) on the meiobenthos of polluted reef; Plante et al. (1989) on redox potential in coral reefs; Alcolado et al. (1997, 1999), Lang et al. (1998) and Quirolo (1998) on the status of Cuban coral reefs; and Núñez Jiménez (1982) and Alcolado (1999b) on legislation relating to marine resources and threats.

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2.5

The Twenty-First Century: First Steps

Some international initiatives and projects directly or indirectly related to coral reefs were developed since the end of the twentieth century, linked to the National Program of Science and Technology of the Ministry of Science, Technology, and the Environment (CITMA). These include the Caribbean Coastal Marine Productivity (CARICOMP 2000), Reef Check, International Coral Reef Action Network (ICRAN), Coast and Beach Stability Project (COSALC), National Geographic Society’s Committee on Research and Exploration, Ocean Research and Education Foundation, Caribbean Planning for Adaptation to Climate Change Program (CPACC), UNDP/GEF Sabana-Camagüey, and Atlantic and Gulf Rapid Reef Assessment (AGRRA 2021; Kramer and Lang 2003). The latter emerged in 1997, directed and supported by the prestigious Robert N. Ginsburg, through his foundation The Ocean Research and Education Foundation (ORE), and was perhaps the most prominent, due to its geographical scope and for involving scientists from different countries and different Cuban research centers. The AGRRA initiative initially had two expeditions with international teams. The first expedition explored the coral reefs in the southern and eastern Gulf of Batabanó in March 2001, and the second did so on the coral reefs of the Jardines de la Reina Archipelago in August of the same year (Fig. 2.16). The third expedition had a national team that covered the coral reefs of the Sabana-Camagüey Archipelago, a year later. AGRRA’s initial goals were to provide a standardized assessment of key structural and functional indicators that could be applied to reveal spatial and temporal patterns of regional reef condition (AGRRA 2021). After these expeditions, many institutions adopted or adapted the AGRRA protocol to their research. Also with the arrival of the new century, the reefs of Cuba are addressed in the remarkable work World Atlas of Coral Reefs by Spalding et al. (2001). It documents the presence of coral reef systems along practically the entire edge of the Cuban shelf and indicates large areas offshore that “resemble barrier reefs,” separated from the main island by wide lagoons. Thus, they considered the reef of the SabanaCamagüey as the longest barrier system, with about 400 km. On the south coast, they documented a reef that extends for more than 350 km, the Jardines de la Reina area. They state that “Unlike true barrier reefs, the lagoons behind these stretches of reefs are very shallow.” In this book, a reef area of 3020 km2 was estimated. It provides information on topics such as urbanization in relation to pollution, hurricane activity, algae cover in relation to diseases and depletion of the Diadema antillarum urchin populations, as well as fishing and collection of reef organisms. In addition, they refer to the

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S. González-Ferrer

Fig. 2.16 Members of the first expedition of the Atlantic and Gulf Rapid Reef Assessment Project (AGRRA), in 2001. From left to right: on the upper level Phillip Kramer, Ken Mark, and James; in the second level Paul Blanchon, Fabián Pina Amargós, and Karel Cantelar; in the third level Ramón, Eric, and Macho (crew), Miguel Hernández, Pedro Alcolado, Elena de la Guardia, Abel (crew), Sergio GonzálezFerrer, Vladimir Kosminin, and Robert Gingsburg; and at the base level Alexis (crew)

regulations for the protection of reefs and considered 46% of these reefs are threatened. Alcolado et al. (2003) well summarize the outlook for Cuban coral reefs. These authors account for the presence of approximately 526 species of marine macroalgae, 160 sponge species, 41 species of stony corals, and 68 gorgonian species and consider turtles, dolphins, and fishes as the major vertebrates associated with Cuban coral reefs. They likewise comment on coral bleaching and diseases and summarize the main natural disturbances that affect these communities such as sedimentation, cyclones and cold fronts, natural supply of nutrients, climate change impacts and El Niño-Southern Oscillation, anthropogenic sedimentation and pollution, as well as on the impacts of tourism, and coral reef fisheries issues. These authors considered the entire edge of the platform to be carpeted with fore reefs (>95%) that they extend approximately 3781 km (3181 km in Alcolado 2006) and considered calling “reef crests or reef flat” what Zlatarski and Martínez-Estalella (1980, 1982) and Kühlmann (1974a) called barrier reefs and Wells (1988) and others called reef banks. Corales Pétreos: Jardines Sumergidos de Cuba (Stony Corals, Submerged Gardens of Cuba) was another largeformat book published in this period (González-Ferrer 2004e, Fig. 2.17). Starting with a basic summary of the characteristics of corals and coral reefs, it updates the list of hermatypical corals, this time with photos of almost all living species, ecological details and threats (González-Ferrer 2004b, c; González-Ferrer et al. 2004a), as well as Cuban environmental management and legislation involving reefs (Alcolado et al. 2004a, b). In addition, it includes a list of

ahermatypic corals (González-Ferrer and Cairns 2004), experiences with corals in the marine aquarium (Caballero

Fig. 2.17 Cover of the book Stony Corals, Submerged Gardens of Cuba (González-Ferrer 2004a–e)

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Research History of Corals and Coral Reefs in Cuba

2004), a summary of many of their associated organisms (González-Ferrer et al. 2004b), connections between reefs and tourism (Alcolado 2004a), and protective measures (González-Ferrer 2004d). This book for the first time discusses the natural history of Cuban corals and their formations (González-Ferrer and Iturralde Vinent 2004) and is published with a public-friendly format, including satellite images (Lorenzo-Sánchez et al. 2004). It also considers the value of corals and coral formations in medicine, the arts, and architecture (González-Ferrer and Villar Zaldivar 2004) and in the historical reconstruction of climate trends (Jiménez and Rixen 2004) and information on the Cuban collections of stony corals (Zlatarski 2004). In this book, the existence of barrier-type formations in Cuba is defended, in opposition to the publications of Spalding et al. (2001) and Alcolado et al. (2003) who did not consider them “true barrier reefs.” González-Ferrer et al. (2004a), besides direct observations, relied on nautical charts, satellite images, and the characteristics of reef barriers in other latitudes, to support the hypothesis of the existence of two coral reefs of this type in Cuba, with reef lagoons deeper than a few meters and developed along the border of the insular shelf. The first one, the Los Colorados Barrier Reef facing the Gulf of Mexico, has an extension of 50 km and a lagoon with maximum depths of 30 m. The second, in the southeastern part of Cuba facing the Caribbean Sea, which they termed “The Great Barrier Reef of Los Jardines de la Reina,” has an extension of 240 km, with a lagoon that reaches 40 m depth. They also suggested that the latter, due to its length, could be considered the longest in the Caribbean region, somewhat longer than the Belize Barrier Reef, estimated 230 km in length by Spalding et al. (2001). Later publications support this statement (Alcolado 2004b; see Chap. 3, Estrada et al. and Chap. 5, Andréfouët and Bionaz). However, Andréfouët and Cabioch (2011) and Goldberg (2013) considered these formations coastal barrier systems or bank barriers, although they did not have many of the aforementioned publications. Burke and Maidens (2005) estimated the area of coral reefs in Cuba to be 3290 km2 (13% of the reefs of the Wider Caribbean). According to these authors, The Bahamas (3580 km2) and Cuba are almost tied in first place in reef extension in the tropical western Atlantic. Later on, in La Biodiversidad Marina de Cuba (Marine Biodiversity of Cuba), Claro (2006) points out that more than 98% of the approximately 3215 km of the edge of the Cuban insular shelf is bordered by frontal reefs, which extend approximately 1440 km in the north and 1675 km in the south, for a total of 3115 km in length. This IDO book, with many partner institutions, includes a CD with a large amount of information across differing habitat and ecosystem types, including species lists for many marine and coastal taxa in Cuba, including coral reef fauna and flora.

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Also in 2004, the Manual de Capacitación para el Monitoreo Voluntario de Alerta Temprana en Arrecifes Coralinos (Training Manual for Voluntary Monitoring of Early Warning in Coral Reefs) was published (Fig. 2.18), as part of the organization and training of the Monitoring Network of that name. This initiative arose from the conclusions of the Symposium “Coral Reefs: United for their Conservation” that took place in Cayo Coco, north Cuba in September 2002, sponsored by the Ministry of Tourism and the UNDP/ GEF Sabana-Camagüey Project and co-sponsored by the ProNaturaleza and Environmental Defense (Alcolado 2004b). It had a broadly inclusive character since not only biologists but also diving instructors and government staff from marine protected areas participated, which enabled early ideas a national summary of the occurrence of coral bleaching by year during peak months of this phenomenon (Alcolado 2007a–c, 2008, 2009, 2010, 2011, 2012, 2013, 2014a–c, 2015). The Monitoring Network operated from 2003 to 2016, the latter being the year when Pedro M. Alcolado (1948–2017) presented his last master lecture at the Ibero-American Forum on Healthy Marine Ecosystems (Hernández-Zanuy 2018). A tireless defender of environmental sustainability, Pedro was the most agglutinating of Cuban scientists related to coral reefs in recent decades (Fig. 2.19). His prolific activity and multidisciplinary and multi-institutional vision became palpable with the gestation and execution of projects such as that of the GEF/UNDP Sabana-Camagüey or AGRRA in this period. Pedro studied, oriented, stimulated, trained, and guided several generations of scientists and researchers on the marine and coastal biodiversity of Cuba and groups of marine benthic organisms, including sponges, gorgonians, polychaetes, anemones, hydrozoans, crustaceans, sea squirts, mollusks, macroalgae, phanerogams, echinoderms, and corals, among others, not only from the point of view of taxonomic knowledge but also in terms of ecological connections (Hernández-Zanuy 2018). Reactivated on the first anniversary of Pedro’s death, currently the Monitoring Network continues to function, directed from CIM, Universidad de Habana, in collaboration with ICIMAR. In the report of the National Program for Biological Diversity for the period 2016–2020, one of the goals was to implement measures to reduce anthropogenic pressures on reefs, including the diving carrying capacity of several sites, the use of anchoring systems, as well as the banning of parrotfish and surgeonfish spearfishing, which are key herbivorous species for the recovery of coral reefs. Enhancing resilience in the coral reefs of Cuba was proposed by Alcolado et al. (2016) in response to one of the priority messages of the fifth National Report of Cuba to the Convention on Biological Diversity, which stated “Prioritize the rehabilitation and restoration of ecosystems to avoid fragmentation, increase resilience and connectivity, and

34

S. González-Ferrer

Fig. 2.18 Cover of Training Manual for Voluntary Monitoring of Early Warning in Coral Reefs (Alcolado 2004a, b)

contribute to adaptation and mitigation of climate change and extreme events.” The proposal entailed actions aimed at promoting the adaptation of reef ecosystems to climate change, including the biological control of benthic algae by urchins and parrotfish, the control of the invasion of lionfish and the protection of large carnivorous fish, banning of bottom trawling, setting of nets and traps, as well as assigning fishing rights or quota to local fishermen and compensation to those affected by these measures, among other legal instruments. These authors considered that, despite the impacts to date, these Cuban reef ecosystems are still “an oasis in the Caribbean.” Another notable action occurred in 2017, coordinated by Prof. John Reed from the Harbor Branch Oceanographic Institute, Florida Atlantic University, and Patricia González from the Centro de Investigaciones Marinas (Marine Research Center) of the University of Havana (Reed et al. 2018, and Chap. 14, Reed et al.; Fig. 2.20). According to

Reed et al. (see Chap. 14), in 2015, a Joint Declaration on Cooperation in Environmental Protection was developed between the United States and the Republic of Cuba, which enabled the US National Oceanic and Atmospheric Administration (NOAA) to sign a memorandum of understanding between the US National Park Service and the Cuban National Center for Protected Areas. It established a “Sanctuary-Sister” relationship between the Guanahacabibes National Park and the Banco de San Antonio Marine Sanctuary in western Cuba and the Florida Keys National Marine Sanctuary (FKNMS) and the Flower Garden Banks National Marine Sanctuary (FGBNMS) in the United States. This project recognizes that all of these regions are inextricably linked through ocean currents. In support of these agreements, a joint research expedition was carried out on the ship F.G. Walton Smith of the University of Miami. The objective of this joint expedition was to determine the distribution of the mesophotic coral reef

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Research History of Corals and Coral Reefs in Cuba

Fig. 2.19 Pedro Manuel Alcolado (1948–2017), the most agglutinating of Cuban scientists related to coral reefs in recent decades

ecosystems and associated fish communities of Cuba, their biodiversity, health status, and connectivity (physical,

Fig. 2.20 Members of the research expedition to study the mesophotic reefs around the entire coast of Cuba in 2017. From left to right: top row, Ulises Fernández Gómez (CITMA International Relations Director), Dr. Beatriz Martínez Daranas (CIM-UH), MSc. Patricia GonzálezSánchez (ANC), MSc. Mayelin Carmenate (ICIMAR), Dr. Alain Muñoz Caravaca (CEAC), Dr. Dorka Cobián Rojas (GNP), MSc. Juliett González Méndez (CNAP), MSc. Linnet Busutil López (ICIMAR),

35

genetic, and ecological) among themselves, with shallow reef systems and with reef systems located in marine protected areas of the southestern United States and the Gulf of Mexico. Work was carried out with the University of North Carolina Mohawk ROV at 36 stations around the entire island of Cuba from depths of 30 m to 150+ m where samples of water and marine organisms, photographs, videos, and oceanographic and topographic data were collected (Reed et al. 2018; Suárez et al. 2020; Chap. 14, Reed et al.). In addition to the aforementioned references, among other publications from this last century, on the study of fauna and flora associated with Cuban reefs are Perera-Montero and Rojas-Consuegra (2005), Medina-Batista (2007), Zlatarski (2007, 2010, 2017), and Iturralde-Vinent and Inne (see Chap. 4) on reef geology and fossil corals; Miravet et al. (2000) and Loza et al. (2003) on coral reef microbiological indicators and phytoplankton; Trelles et al. (2001), Valdivia (2004), Valdivia-Acosta and de la Guardia (2004b), Suárez et al. (2015), Martínez-Daranas et al. (2016), and Suárez and Martínez-Daranas (see Chap. 6) on reef macroalgae communities; Núñez-Luis et al. (2018) on foraminifera as reef bioindicators; Alcolado (2002, 2007d), Alcolado et al. (2004a), Caballero et al. (2009b), Nuez et al. (2011), Busutil and Alcolado (2012), and Busutil and García-Hernández (see Chap. 7) on reef sponge communities; Castellanos-Iglesias

Prof. John Reed (FAU, USA), Dr. Patricia González-Díaz (CIM-UH), MSc. Carlos Díaz (CNAP), Michael Studivan (FAU, USA), and Andrew David (NOAA Fisheries, USA); on the lower level MSc. Jorge Viamontes Fernández (GEOCUBA), Daniel Estrada Pérez (GEOCUBA), Lic. Alain García Rodríguez (ICIMAR), and Dra. Cristina Díaz (FAU, USA)

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(2018) on coral reef hydroids; Varona and Varela (2005), Varona et al. (2004, 2005) Varela and Orozco (2006), Hernández-Fernández and Alcolado (2007), Alcolado et al. (2008), Varela et al. (2008a), and Rey-Villiers (2009) on reef octocoral communities; de la Guardia (2000, 2006), Zlatarski et al. (2004), Zlatarski (2009), Zlatarski and Stake (2012), González-Ferrer et al. (2007a, 2021 in prep.), SemideyRavelo (2008), Hernández-Fernández et al. (2008a), González-Ferrer (2009), Alcolado et al. (2009a), GonzálezDíaz (2010), Perera-Valderrama (2010), Ferrer-Rodríguez et al. (2016), Hernández-Fernández and Bustamante-López (2017), Zlatarski (2018a), Hernández-Fernández et al. (2019), Caballero et al. (2020), González-Ferrer et al. (see Chap. 8), and Caballero et al. (see Chap. 10) on stony coral communities; González-Ferrer (2006) on coral spawning; Perera (2012), Alcolado-Prieto et al. (2012), Alcolado-Prieto (2015), Hernández-Fernández and Bustamante-López (2019), and Ramos-Romero et al. (2019) on coral recruits; Espinosa et al. (2012), Espinosa and Ortea (2007), and Espinosa and González-Ferrer (see Chap. 10) on mollusk reef communities; Nodarse-Konnorov (2001) and MartínBlanco et al. (2010) on reef echinoderm communities; Baisre (2000) and Claro et al. (2001), Claro and Cantelar (2003), González-Sansón and Aguilar (2003), Claro and Lindeman (2003), Paris et al. (2005), Chevalier and Cárdenas (2005a, b, 2006), García et al. (2005), Hernández et al. (2008a, b); PinaAmargós (2008), Claro et al. (2009), Cobián and Chevalier (2009), Durán and Claro (2009), Cobián (2010), Cobián et al. (2011, 2016), Pina-Amargós et al. (2014), Kough et al. (2016), Navarro-Martínez et al. (2017), Baisre (2018), Durán et al. (2018), Claro et al. (2019), Chevalier et al. (see Chap. 12), and Pina-Amargós et al. (see Chap. 13) on reef fishes, highly connected to coral reefs (e.g., as nutrient exchangers across associated mangrove and pelagic food webs). Also worth mentioning are systematic and ecological studies such as Aguilar et al. (2000), de la Guardia and González-Sansón (2000a, b), Lalana et al. (2001), de la Guardia and González (2002), de la Guardia et al. (2003, 2004a, b, 2005, 2006a, b), González-Díaz et al. (2003, 2008, 2010), Valdivia et al. (2004), Valdivia-Acosta and de la Guardia (2004a, b), Caballero et al. (2004, 2005a, b, 2009a), de la Guardia (2005), González-Ontivero et al. (2007), González-Ontivero and de la Guardia (2008), PinaAmargós et al. (2008), Varela et al. (2008b), Caballero and Alcolado (2011), Hernández-Fernández et al. (2002, 2016a, 2018), Alcolado et al. (2001a, 2010a, b), Alcolado and Durán (2011), Hernández-Fernández et al. (2011), Alcolado-Prieto et al. (2013), Perera-Valderrama et al. (2013), Busutil et al. (2016), Rey-Villiers et al. (2016), Zlatarski and GonzálezFerrer (2017), and Zlatarski and Greenstein (2020) on reef bentonic fauna structure and conditions; Zlatarski (2018b) on

S. González-Ferrer

mesophotic coral ecosystems; Caballero and de la Guardia (2003), Caballero et al. (2005b) on reef collection zone evaluation; Caballero et al. (2013) and González-Dias et al. (2013) on reef monitoring protocol; de la Guardia et al. (2004b), Hernández-Fernández et al. (2008b, 2016b), and Figueredo-Martín et al. (2015) on the impact of diving and load capacity for diving activities; Alcolado et al. (2009b), Caballero et al. (2012), and González-Cano et al. (2006) on coral reefs and hurricanes; Armenteros et al. (2008) and González-Díaz et al. (2014) on climate reconstruction; Alcolado et al. (2013) on reef resilience; Alcolado et al. (2004b), Caballero et al. (2010), Gómez et al. (2011), Lara et al. (2012), Perera-Valderrama et al. (2017, 2018), González-Díaz et al. (see Chap. 17), Cobián-Rojas et al. (see Chap. 18), González-Méndez (see Chap. 19), PinaAmargós et al. (see Chap. 15), and Perera-Valderrama et al. (2005, see Chap. 20) on marine protected areas and management; Burke and Maidens (2005) and Bowdery et al. (2015) on legislation, threats, and good practices; Woodley et al. (2000), Alcolado et al. (2000, 2001b–d, 2010a, b), GonzálezFerrer et al. (2003, 2007b), Castellanos-Iglesias et al. (2004), Caballero et al. (2007), Creary et al. (2008), Wilkinson (2008), Wilkinson and Souter (2008), Caballero and Perera (2014), González-Diaz et al. (2018), Caballero et al. (2016, 2019), Gil-Agudelo et al. (2020), and Pina-Amargós et al. (see Chap. 15) on the status of Cuban coral reefs. To end this brief tour through the history of the study of coral reefs in Cuba, a scientific discipline is presented that we consider one of the most recently introduced on the island: the study of Cuban coral reefs from the economic perspective including the valuation of ecosystem goods and services. This area of economics began to arise in Cuba at the end of last century, mainly within the university academic field. This resulted in the creation of several graduate and doctoral theses, some of which were promoted through the SabanaCamagüey project (Figueredo-Martín et al. 2014a, b; see Chap. 21, Rangel Cura et al.; Chap. 22, Figueredo-Martín et al.). With direct emphasis on coral reef valuation, the UNDP/GEF Archipelagos del Sur is the most important project so far. Ferro-Azcona et al. (2014) included three case studies that explore the contribution to well-being of coral reefs in three marine protected areas (see Chap. 22, Figueredo-Martín et al.; Chap. 23, Ferro-Azcona et al.). By 2012, capacities were extended, both in specialists and resources, for studies of economic valuation of ecosystem goods and eervices and environmental damage. These studies developed from academic initiatives or international projects to becoming a foundational need and a line of work for the country’s environmental policy (see Chap. 21, Rangel Cura et al.). Castañeda and Angulo-Valdés (2016) pointed out how the Marine Research Center of the University of Havana incorporated these studies into the Marine Resources

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Research History of Corals and Coral Reefs in Cuba

Management and Conservation Group in the last decade and proposed lines of research, including the estimation of reef values, the evaluation of ecological tourism alternatives and of damages, and the development of procedures to organize these processes (see Chap. 22, Figueredo-Martín et al. and Chap. 23, Ferro-Azcona et al.). Many are the developing and pending topics that involve the coral reefs of Cuba. The island’s history of marine science has countless beginnings, but never an end. Sometimes, it sleeps when one of its muses sails, but at some point, a new plunge takes place and it wakes up again. Acknowledgments I would like to thank Ana María Suarez, Beatriz Martínez Daranas, Tamara Figueredo-Martín, Patricia González Díaz, Vassil Zlatarski, José Espinosa, Reinaldo Estrada, Fabián Pina Amargós, and John Reed, for their valuable input and revisions in the making of this chapter.

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Research History of Corals and Coral Reefs in Cuba

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42 García-Parrado P, Alcolado PM (1996) Catálogo de Octocorales (Cnidaria) de Cuba, con comentarios sobre su taxonomía. Avicennia 415:41–45 García-Parrado P, Alcolado PM (1998) Nuevos registros de Octocorales para la plataforma cubana. Avicennia 8(9):105–108 Gil-Agudelo DL, Cintra-Buenrostro CE, Brenner J, González-Díaz P, Kiene W, Lustic C, Pérez-España H (2020) Coral reefs in the Gulf of Mexico large marine ecosystem: conservation status, challenges and opportunities. Front Mar Sci 6:807. https://doi.org/10.3389/fmars. 2019.00807 Goldberg WM (2013) The biology of reefs and reef organisms. The University of Chicago Press. 401 p Gómez BM, Helguera PY, Cabrales CY, Rosell AG, Hernández BY, Caballero AH, Chevalier MP, Pérez AA, Pérez HA (2011) Línea base flora y fauna marina en el área protegida: CienfuegosGuajimico. Informe del proyecto: Aplicación de un enfoque regional al manejo de áreas costeras y marinas protegidas en los archipiélagos del sur de Cuba. Centro de Estudios Ambientales de Cienfuegos y Acuario Nacional de Cuba. 27 p González-Cano J, Ibarra-Navarro R, Vega-Zepeda A, Huitrón-Baca JC, Najera Hillman E (2006) Rehabilitation program of corals damaged by hurricane “Wilma” in October 2005, Isla Mujeres-Cancun National Park, Mexico. Proc Gulf Caribb Fish Inst 59:459–464 González-Díaz P (1999) Comunidades de esponjas, corales y gorgonias en un arrecife coralino costero de Ciudad de La Habana. Trabajo de Diploma, Universidad de La Habana. 25 p González-Díaz P (2010) Efecto acumulativo de agentes estresantes múltiples sobre los corales hermatípicos de la región noroccidental de Cuba. Tesis presentada en opción al grado científico de Doctor en Ciencias Biológicas. Universidad de La Habana, Centro de Investigaciones Marinas, Ciudad de La Habana. 100 p González-Díaz P, de la Guardia E, González-Sansón G (2003) Efecto de efluentes terrestres sobre las comunidades bentónicas de arrecifes coralinos de Ciudad de La Habana, Cuba. Rev Investig Mar 24(3): 193–204 González-Díaz SP, González-Sansón G, Piloto CY, Cabrales S, Álvarez F (2008) Estructura de las poblaciones de Acropora palmata, Porites asteroides y Agaricia agaricites forma masiva (Cnidaria; Scleractinia) en el arrecife de Playa Baracoa, Cuba. Rev Investig Mar 29(3):213–224 González-Díaz P, González-Sansón G, Álvarez-Fernández S et al (2010) High spatial variability of coral, sponges and gorgonian assemblages in a well preserved reef. Rev Biol Trop 58(2):621–634 González-Díaz P, Martínez YB, Perera O, Álvarez S (2013) Estimación de indicadores ecológicos a nivel de comunidad y población de corales hermatípicos en arrecifes con grado diferente de impacto. UCE Ciencia 1:112–132 González-Díaz P, Nuñez-Luis JY, Hernández-Zanuy A (2014) Bases metodológicas para las reconstrucciones paleoclimáticas en corales hermatípicos y foraminíferos. En Métodos para el estudio de la biodiversidad en ecosistemas marinos tropicales de Iberoamérica para la adaptación al cambio climático RED CYTED BIODIVMAR, pp 50–71 González-Díaz P, González-Sansón G, Aguilar-Betancourt C, ÁlvarezFernández S, Perera-Pérez O, Hernández-Fernández L, Ferrer Rodríguez VM, Cabrales Caballero Y, Armenteros M, de la Guardia LE (2018) Status of Cuban coral reefs. Bull Mar Sci 94:229–247. https://doi.org/10.5343/bms.2017.1035 González-Ferrer S (2004a) Introducción. In: González-Ferrer S (ed) Corales pétreos, Jardines sumergidos de Cuba. Editorial Academia, La Habana, pp 19–25 González-Ferrer S (2004b) Capítulo I: Generalidades. In: GonzálezFerrer S (ed) Corales pétreos, Jardines sumergidos de Cuba. Editorial Academia, La Habana, pp 27–43 González-Ferrer S (2004c) Capítulo III: Corales pétreos de Cuba. Catalogo de corales hermatípicos de aguas cubanas. In: González-

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2

Research History of Corals and Coral Reefs in Cuba

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46 coral reefs and associated fish communities. Rev Investig Mar 38(1): 56–125 Rey-Villiers N (2009) Características de las comunidades de octocorales y ensayo de su uso como bioindicadores en un gradiente de contaminación en el oeste de La Ciudad de La Habana, Cuba. Tesis de Diploma. Instituto de Oceanología. 69 p Rey-Villiers N, Alcolado-Prieto P, Busutil L, Caballero H, PereraPérez O, Hernández-Fernández L, González-Díaz S, Alcolado PM (2016) Condición de los arrecifes coralinos del Golfo de Cazones y el Archipiélago de Los Jardines de la Reina, Cuba: 2001-2012. In: Rey-Villiers N (ed) Línea base ambiental para el estudio del cambio climático en el golfo de Cazones y el archipiélago Jardines de la Reina. Instituto de Oceanología, CITMA, La Habana, Cuba, pp 93–149 Richter R (1967) Unsere Arbeitsmethoden (Kubanische Korallen für Berlin). Poseidon 71(11):486–489 Rivero LH, Bermúdez PJ (1938) Expedición biológica a los Mares de Cuba concertada entre las Universidades de Harvard y de La Habana. Publ. Rev. Univ, Habana. 24 p Rodríguez R, Sánchez-Arango JR, Toucet S (1989) Pronóstico de estructuras arrecifales y salinas en la plataforma marina septentrional de Cuba. Serie Geológica. Centro de Investigaciones del Petróleo, pp 92–129. Número Especial 2 Roig JT (1917) Plantas nuevas o poco conocidas de Cuba. Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey”, vol II, pp 109–123. 1916–1917. No. 4. Julio-Agosto, 1916 Saco JA (1858a) Noticia sobre la Obra del Señor Parra. Colección de papeles científicos, históricos, políticos y de otros ramos Sobre la Isla de Cuba, ya publicados, ya inéditos, vol 1. Imprenta de D’Aubusson y Kugelmann, Paris, pp 331–341 Saco JA (1858b) Historia Física Política y Natural de la Isla de Cuba. Colección de papeles científicos, históricos, políticos y de otros ramos Sobre la Isla de Cuba, ya publicados, ya inéditos, vol 1. Imprenta de D’Aubusson y Kugelmann, Paris, pp 331–341 Sánchez-Roig M (1928) Instituto Nacional Científico y Museo de Historia Natural, La Habana. 220 p Sánchez-Valero MV, Sánchez JI, Muñoz J, Yagüe F (2009) El gabinete perdido. Pedro Franco Dávila y la Historia Natural del Siglo de las Luces: un recorrido por la ciencia de la Ilustración a través de las “producciones marinas” del Real Gabinete (1745–1815). CSIC, Madrid, pp 993–1027 Semidey-Ravelo A (2008) Variaciones espaciales y temporales en comunidades de corales de arrecifes de cresta del norte de La Habana, Cuba. Trabajo de Diploma. Universidad de La Habana. 48 p Seneš J (1966) Recent facies of Guanabo shelf (Cuba). Geol Sbornik, Bratislava, XVII 2:283–304 Socorro R, Sánchez-Arango JR, Kuznetsov V (1986) La búsqueda de arrecifes en Cuba. Serie Geológica, vol 3. Centro de Investigaciones del Petróleo, pp 64–73 Spalding MD, Ravilious C, Green EP (2001) World atlas of coral reefs. Prepared at the UNEP World Conservation Monitorie Centre. University of California Press, Berkeley, USA. 424 p Suárez AM (1989) Ecología del macrofitobentos de la plataforma de Cuba. Rev Investig Mar 10:187–206 Suárez AM, Martínez-Daranas B, Alfonso Y (2015) Macroalgas marinas de Cuba. Editorial UH, La Habana. ISBN: 978-959-721144-0 Suárez AM, García T, Ortíz M (2020) 50 Años, 1970–2020. Centro de Investigaciones Marinas, Universidad de La Habana. El CIM-UH nuestro orgullo, el mar, nuestra razón. Imagen Contemporánea, 262 p Taege M, Wagner J (1975) Taucher am Korallenriff. VEB. F. A. Brockhaus Verlag, Leipzig. 156 p Taylor RC (1836) Notes on natural objects observed while staying in Cuba. Mag Nat Hist J Zool Bot Mineral Geol Meteorol IX:449–457

S. González-Ferrer Trelles J, Suárez AM, Callado-Vides L (1997) Macroalgas del arrecife de la Herradura, Costa NO de La Habana. Rev Investig Mar 18(3): 191–192 Trelles J, Suárez AM, de la Guardia E (2001) Macroalgas dominantes de Playa Herradura, plataforma noroccidental de Cuba: Caulerpales y Dictyotales. Rev Investig Mar 22(1):1–6 Valdés-Muñoz E, Garrido OH (1987) Distribución de los peces en un arrecife costero del litoral habanero. Rep Inv Inst Ocean, Acad Cien Cuba 61:1–22 Valdivia A (2004) Variación espacial y temporal de las asociaciones de algas en zonas del sublitoral norte habanero, Cuba. Tesis de Maestría, Centro de Investigaciones Marinas, Universidad de La Habana. 121 p Valdivia A, de la Guardia E, González-Díaz P, Aguilar C (2004) Inventario de los componentes más comunes de la flora y la fauna de los arrecifes coralinos de la Península de Guanahacabibes, Pinar del Río, Cuba. Rev Investig Mar 25(2):113–122 Valdivia-Acosta A, de la Guardia E (2004a) Estructura de la comunidad de corales en el arrecife costero Boca de Canasí, La Habana, Cuba. Rev Investig Mar 25(1):15–22 Valdivia-Acosta A, de la Guardia E (2004b) Variaciones espaciales y temporales de la comunidad de algas en el arrecife costero Boca de Canasí, La Habana, Cuba. Rev Investig Mar 25(2):123–132 Varela C, Orozco MV (2006) Nuevo hallazgo de Lophogorgia cardinalis (Gorgonacea) en aguas cubanas. Cocuyo 16:5–6 Varela C, Orozco MV, Varona G (2008a) Registros nuevos de octocorales (Cnidaria: Anthozoa: Octocorallia) para Cuba. COCUYO 17:10–11 Varela C, Castellanos S, Hernández L (2008b) Registros nuevos de invertebrados (Cnidaria y Crustacea) para Cuba. COCUYO 17:12– 14 Varona G, Varela C (2005) Primer hallazgo de Eunicea pinta Bayer y Deichmann 1958 (Cnidaria, Gorgonacea) en aguas cubanas. COCUYO 15:3–4 Varona G, Caballero H, de la Guardia E (2004) Estructura ecológica de la comunidad de octocorales en la costa oriental de Bahía de Cochinos. Rev Investig Mar 25(3):209–218 Varona G, Caballero H, de la Guardia E (2005) Estructura ecológica de la comunidad de gorgonáceos en la costa norte oriental de La Habana. Rev Investig Mar 26(1):27–36 Vaughan TW (1919) Fossil corals from Central America, Cuba, and Porto Rico with an Acount of the American tertiary, Pleistocene, and recent coral reefs. US Nat Mus Bull 103:VI 189–VI 524. 68–152 pls Vaughan TW (1933) Report on species of corals and larger foraminifera collected in Cuba by O.E. Meinzer. J Wash Acad Sci 23:352–355 Wagner J (1967a) In den Riffen von Arroyo Bermejo. (Kubanische Korallen für Berlin). Poseidon 68(8):346–350 Wagner J (1967b) Wir tauchen mit Freunden. Poseidon 70(10):456–459 Wagner J (1967c) Erste Bilanz (Kubanische. Korallen für Berlin). Poseidon 70(10):460–462 Wagner J (1967d) Tick-tack Test in Kubanische Riffen. Poseidon 71(11):508–509 Walton-Smith FG (1948) Atlantic reef corals. A handbook of the common reef and shallow waters corals of Bermuda, Florida, the West Indies and Brazil. University of Miami press. 112 p, 41 pls Wells FG (1934) Eocene corals, pt. 1, from Cuba; pt. 2, a new species of Madracis from Texas. Bull Amer Pal 20. 20 p, 3 pls Wells JW (1941) Upper cretaceous corals from Cuba. Bull Am Paleontol 26(97):1–16. 282–298 Wells SM (1988) Coral reefs of the world. In: Atlantic and Eastern Pacific, vol 1. UNEP & IUCN, Nairobi, Gland. 373 p Wikipedia (2021) Wikimedia commons. File: Stieler, Joseph KarlAlexander von Humboldt-1843.jpg. https://es.wikipedia.org/wiki/ Alexander_von_Humboldt#/media/Archivo:Stieler,_Joseph_Karl__Alexander_von_Humboldt_-_1843.jpg

2

Research History of Corals and Coral Reefs in Cuba

Wilkinson C (2008) Status of the coral reefs of the world: 2008. Global Coral Reef Monitring Network and Reef and Rainforest Research Centre, Townsville, Australia. 296 p Wilkinson C, Souter D (2008) Status of Caribbean coral reefs after bleaching and Hurricanes in 2005. Global Coral Reef Monitoring Network, and Rainforest Research Center, Townsville. 152 p Woodley JD, Alcolado PM, Austin T, Barnes J, Claro-Madruga R, Ebanks-Petrie G, Estrada R, Geraldes F, Glasspool A, Homer F, Luckhurst B, Phillips E, Shinn D, Smith R, Sullivan-Sealey K, Vega M, Ward J, Wiener J (2000) Status of coral reefs in the northern Caribbean and western Atlantic. In: Wilkinson C (ed) Status of coral reefs of the world: 2000, global coral reef monitoring network. Australian Institute of Marine Science, pp 239–226 Zenkovich VP (1965) Costas coralinas de Cuba. Vokrug sveta 12:12– 13. (in Russian) Zenkovich VP (1969) Zonas someras de Cuba occidental y sus sedimentos. Okeanologia, IX 2:256–270. (in Russian) Zenkovich VP (1972) En el lejano mar azul. Varna. 309 p (en búlgaro) Zenkovich VP, Ionin AS (1969) Breve resumen sobre las investigaciones de la estructura y dinámica de la zona litoral de la isla de Cuba. Ser. Oceanol, vol 8. Acad. Cien. de Cuba, La Habana. 22 p Zlatarski VN (1975) On the coral fauna of Cuba (in Russian), vol 202. Ancient Cnidaria, Trans. Inst. Geol. & Geoph., Acad. Sci. U.S.S.R, Siberian Branch, pp 231–236 Zlatarski VN (1981) Publications on the Cuban fossil corals (Quarternary excluded). Fossil Cnidaria, Brisbane 10(2):31–32 Zlatarski VN (2004) Capítulo V: La colección más valiosa de los escleractinios del Atlántico. In: González-Ferrer S (ed) Corales pétreos, Jardines sumergidos de Cuba. Editorial Academia, La Habana, pp 253–257 Zlatarski VN (2007) The scleractinian species – a holistic approach. In: Hubmann B, Piller WE (eds) Fossil corals and sponges. Proceedings of the 9th international symposium on fossil Cnidaria and Porifera, vol 17. Österr. Akad. Wiss., Schriftenr. Erdwiss. Komm, Wien, pp 523–531 Zlatarski VN (2009) Need for a more integrative approach to scleractinian taxonomy. In: Riegl B, Dodge R (eds) Proceedings of the 11th international coral reef symposium, Ft. Lauderdale, Florida, pp 1412–1416

47 Zlatarski VN (2010) Palaeobiological perspectives on variability and taxonomy of scleractinian corals. Palaeoworld 19(3–4):333–339 Zlatarski VN (2017) The actuopaleontological studies on Cuban scleractinians and coral reefs of half a century ago are not over. Geologica Balcanica 46(2):111–116. https://www.geologicabalcanica.eu/journal/46/2/ Zlatarski VN (2018a) Morphology or genetics – the case of Caribbean branching Porites corals. Reef Encounter 33(1):55–57 Zlatarski VN (2018b) Investigations on mesophotic coral ecosystems in Cuba (1970–1973) and Mexico (1983–1984). CICIMAR Oceánides 33(2):27–43. http://oceanides.ipn.mx/index.php/cicimaroceanides/ article/view/230 Zlatarski VN, González-Ferrer S (2017) Gran Banco de Buena Esperanza: unique Caribbean coral reef system. Reef Encounter 32(1):60–62 Zlatarski VN, Greenstein BJ (2020) The reticulate coral reef system in Golfo de Guacanayabo, SE Cuba. Coral Reefs 39(3):509–513. https://doi.org/10.1007/s00338-020-01933-7 Zlatarski VN, Martínez Estalella N (1980) The scleractinians of Cuba with data of associated organisms. Editions de l’Académie bulgare des Sciences, Sofia. (in Russian) Zlatarski VN, Martínez Estalella N (1982) Les Scléractiniaires de Cuba avec des données sur les organismes associés. Editions de l’Académie bulgare des Sciences, Sofia Zlatarski VN, Martínez Estalella N (2018) Los escleractinios de Cuba con datos sobre sus organismos asociados. Harte Research Institute for Gulf of Mexico Studies at Texas A&M University, Corpus Christi. 471 p, 1 anexo. http://www.harteresearchinstitute.org/ project/legacy-book-los-escleractinios-de-cuba-corals-cuba Zlatarski VN, Stake JL (2012) The scleractinian corals: a perspective. Geol Belg 15(4):370–375 Zlatarski VN, Chevalier JP, Duarte-Bello PP, Geyer OF, Gill G, Krasnov EV, Morycowa E, Russo A, Wells JW et al (1973) Glossary of equivalent terms for scleractinian (Madreporaria) studies in English, German, French, Italian, Spanish, Polish, Russian, Bulgarian. Fossil Cnidaria, International Newsletter, CNRS, Paris, pp 34–55c Zlatarski VN, Alcolado PM, González Ferrer S, Kramer P (2004) Archipiélago Jardines de la Reina, Cuba. Thirty years later, species richness of scleractinian corals remains high. Reef Encount Mag Int Soc Reef Stud 32:30–32

Part III Description

3

Physical-Geographic Characteristics of Cuban Reefs Reinaldo Estrada Estrada , Gustavo Martín Morales , Joán Hernández-Albernas , Rodney Borrego Acevedo, Jorge Olivera Acosta, Yudelsy Carrillo Betancourt, Idalmis Almeida Martínez , and Lourdes Coya de la Fuente

Abstract

Cuban reefs were mapped following an expert-driven approach using satellite imagery and complemented by previous maps, studies, and fieldwork. Mapping classes were defined initially according to their geographical characteristics (such as surface area, lengths, general and patch spatial distribution, average depths, and other general descriptions), the geomorphology on which they develop, and also on regionalization and typology. The analyses were made up to a maximum depth of 20–35 m (depending on water transparency sunlight penetration). Current results were compared with others existing for Cuba. Keywords

Geography · Coral reefs · Cartography · Marine ecosystems · Cuba

R. E. Estrada (✉) Fundación Antonio Núñez Jiménez de la Naturaleza y el Hombre, La Habana, Cuba G. M. Morales Escuela Nacional de Estudios Superiores Unidad Mérida (ENESUNAM Mérida), Mérida, Mexico J. Hernández-Albernas Refugio de Fauna Cayo Santa María, Villa Clara, Cuba R. Borrego Acevedo Universidad de Queensland, Brisbane, Australia e-mail: [email protected] J. O. Acosta Instituto de Geofisica y Astronomia, La Habana, Cuba Y. C. Betancourt · I. A. Martínez Instituto de Geografía Tropical, La Habana, Cuba L. C. de la Fuente Direccion General de Medio Ambiente. CITMA, La Habana, Cuba e-mail: [email protected]

3.1

Introduction

There are numerous biological and geographic characterizations of the Cuban reefs, comprehensively described by Alcolado et al. (2003), González-Ferrer et al. (2004), and González-Ferrer (see Chap. 2). A recent study, by Andréfouët and Bionaz (see Chap. 5), classifies Cuban reefs from Landsat satellite images following the Millennium Coral Reef Mapping Project methodology. Recent advances in image availability, processing techniques, and the multidisciplinary approach applied by Cuban experts from different disciplines allowed the mapping of coral reefs for this study. In this chapter, we characterized the regional and typological subdivision of Cuban coral reefs based on an expert-driven mapping approach and some of the most relevant metrics (4.77% of the Cuban platform and seamount are covered with coral).

3.2

Data and Methodology

3.2.1

Datasets

The groundwork for this chapter is the recent completion of a new map of the Cuban reefs (Estrada Estrada et al. 2022). Maps showing the location of coral reefs in Cuba were created using a number of different sources to delineate their extent and general shape. A total of 144 publicly available satellite images (64 Landsat 8 Operational Land Imager (OLI) and 80 Sentinel 2) were selected to map and characterize Cuban coral reefs. Images were filtered by their lowest cloud coverage, sun-glint, and highest water clarity in areas of interest and downloaded from USGS (2020) and the Copernicus Open Access Hub (ESA 2020). Image corrections were done to minimize undesirable atmospheric effects by image providers (analysis ready data (ARD) approach). Additional image manipulation techniques were applied by the authors to further enhance the visual

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_3

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appearance of the scenes. Sentinel 2 images were corrected at 1C level including geometric alignment and radiometric correction at surface reflectance (SR) by ESA (2020). Landsat 8 scenes were corrected against geometric and radiometric effects down to SR by USGS (2020). To have the scenes standardized at the same level of correction was advantageous as this ensured higher accuracy in image interpretation. Scene details used in this study are available at https://github. com/RSRCsupport/Cuban-Coral-Reefs. Sources of ancillary information were used to support image interpretation including free high spatial resolution satellite imagery, Bing (2022), Google Earth (2022), Nautical Charts (Navionics 2020), and existing maps from domestic cartography. Extensive fieldwork was carried out over the years by the authors which significantly increased the knowledge of coral reefs in this study. Photo transect (Roelfsema and Phinn 2010) was the survey method, also used to collect field data for the international initiative Allen Coral Atlas (2020) which aimed to map coral reefs globally. Photo transects were adjusted to Cuban conditions such as using cell phones as GPS mounted in plastic bottles instead of floating devices. Coral reefs in this study were mapped following the 30 m depth contour derived from national tidal corrected bathymetric charts purchased for a previous project (Geocuba 2006).

3.2.2

Outline of Methods

Over the years, the authors in this study thought about generating maps of sea bottom types including those in turbid

waters with seagrass beds. We started by visually exploring optical satellite imagery from different sensors and found that, in most of the scenes, sea bottom structures could be seen from space, due to the light penetration characteristics of Cuban waters. On June 2, 2020, we started to map shallow coral reefs in Cuba. Table 3.1 depicts main steps in this study. We held two workshops in July and October 2020 with Cuban geographers, biologists, and ecologists to define mapping classes, zonal distribution patterns, and six mapping areas (Table 3.1a). Authors in this chapter (hereafter experts) were assigned to each mapping area, one expert per area, based on their knowledge, experience, and extensive fieldwork carried out on local coral reefs. We constantly followed up the progress of the workshop via social media when available, as the Internet is limited for Cuba. A resulting action from the workshops was to review literature to get insights of the geographical and typological characteristics of Cuban coral reefs. We identified coral structures down to 43–54 m deep in some cases, values close to the maxima reported in the literature (Gordon and Mccluney 1975). This was possible due to the highest water transparency in some areas, mainly seen on Landsat 8 images, using a true color composite combination of spectral bands 4, 3, 2, and 1 for this sensor, or in shallow areas above 20–25 m, Sentinel 2 scenes with spectral bands 4, 3, and 2 were predominantly used in our workflow as they provided the appropriate spatial resolution to identify coral reef formations. Selected Sentinel 2 and Landsat 8 images were visually inspected and clipped to areas of interest (AOI) per mapping area. We selected AOI that underwent image segmentation

Table 3.1 Methodological steps in the expert-driven mapping approach for the physical and geographic characterization of coral reefs in Cuba (a) Expert discussion forums during research design . Two multidisciplinary workshops (July/October 2020) . Definition of mapping classes . Division of Cuba in six mapping areas based on local expertise . Workshop following up via online meeting . Action: A broad literature review for geographical and typological characteristics of coral reef

(b) Satellite image filtering and acquisition . Landsat 8 OLI (64) and Sentinel 2 (80) corrected from source . Selection of areas of interest in imagery

(c) Image segmentation via Object Based Image Analysis (OBIA)

Data processing

(d) Segment labelling/manual digitalization/quality assurance-control (QA/QC)

(e) Accuracy assessment

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Physical-Geographic Characteristics of Cuban Reefs

53

research design, we assessed different automated supervised and unsupervised classification algorithms (Jensen 2005b) in their capacity to generate thematic maps that accurately depict coral reef features. These algorithms were discarded due to unsatisfactory results. We assessed the accuracy of the coral reef map (Estrada Estrada et al. 2022) generated from the expert-driven approach followed in this chapter (Table 3.1e). Traditional accuracy assessment of thematic maps involves reporting errors in a matrix defined as confusion or error matrix, where rows and columns represent the agreement between the mapping categories (e.g., in pixels or polygons) with the actual category in the ground (i.e., reference data) (Congalton 1991). Together with the error matrix, further quantification of errors includes classes that were omitted in the map (omission error/producer’s accuracy) and classes that were erroneously included in the map (commission error/user’s accuracy). We generated 5052 random points distributed across the marine platform defined by the 30 m depth contour. A subset of 887 points was selected which coincided with corals from the map generated in this study and from maps previously published by other authors (see Table 3.2). These points were manually matched against additional Sentinel images and higher spatial resolution datasets from Bing (2022) and Google Earth (2022) used as references for presence/no presence of coral reefs. An error matrix and corresponding producer and user’s accuracy estimates were generated to assess map accuracy in this study.

using object-based image analysis (OBIA) to split images into objects (Table 3.1c). Object-based image analysis methods include the initial step of using specific software to divide images into objects or “segments” as a result of grouping pixels based on their spectral and contextual information (Blaschke 2010). Bands 2 and 4 were used in segmentation and proved very useful to delineate borders of fore reefs and reef crests. Experts in each mapping region labelled image segments with eight defined mapping classes (see Sect. 3.2.3) using QGIS open-source software (QGIS 2020), version 3.16 “Hannover” and SAGA 7.8 (Conrad et al. 2002; Bechtel et al. 2008). Image interpretation for labelling image segments required a flexible approach where experts tested (1) different spectral band combinations in imagery, (2) userdefined histograms for contrast enhancement, and (3) spatial filtering for smoothing pixel brightness. These image enhancement steps (Jensen 2005a) ensured coral reef features were properly identified during segment labelling. The use of ancillary information and fieldwork data (described in Sect. 3.2.1) significantly improved segment labelling by experts. We applied manual digitization to create additional polygons or to edit segments when segmentation did not produce homogenous or “pure” segments. A further quality assurance/quality control (QA/QC) process consisted of (a) experts undertaking cleanup of output maps by removing and editing non-homogeneous segments and (b) a senior expert (Estrada, R.) reviewing maps per mapping region editing segments as needed (Table 3.1d). As part of the Table 3.2 Cuba coral reef area (km2) per reef type and Cuban platforms Reef types

Great Cuban units Ecological reef zones Reef crest Fore reef Patch reef Atoll- like reef structures Other coral habitats Local reef structures Mud-bottom Guacanayabo reefs Ana María back reef slopes Seamount reefs Total km2 % Cuban reefs

Fringing reefs Cuban platforms

Bank reefs Total km2

% Cuban reefs

136.66 1079.82 89.37 554.06

0.25 1.95 0.16 1.00

386.1

0.70

241.34

241.34

0.44

15.82

15.82

0.03

142 2645.17

0.26 4.77

NW

NC

28.45 91.84 37

25.22 153.32 2.2

8.63 291.83 35.27 543.01

32.78 244.29 13.67 11.05

8.05

62.54

49.03

254.63

165.34 0.30

243.28 0.44

SE

1184.93 2.14

SW

556.42 1.00

North Hab-Mat

5.06 63.29 0.1

68.45 0.12

East

Cuba southcentral

South Guanahacabibes

36.02 180.12 1

0.09 21.98 0.13

0.41 33.15

11.56

0.29

228.7 0.41

22.49 0.04

33.56 0.06

Sea mounts

142 142 0.26

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3.2.3

Definition of Mapping Classes

Cuban reefs are traditionally defined as fringing and bank reefs (Seamounts) (Alcolado 2004; González-Ferrer et al. 2004). There is a current debate between Cuban scientists whether barrier reefs occur in the Cuban platform. The latest work by Andréfouët and Bionaz (see Chap. 5) defined actual outer and intra-barrier reef complexes out of nine categories used by the Millennium Coral Reef Mapping Project classification schema. The definition of classes in this study follows two main levels, fringing and bank reefs. We included the ecological zones from Alcolado (2004) and also considered one local structure (Ana María back reef slopes, following Kramer P. (1992)) review of Geister, in addition to mud-bottom Guacanayabo reefs defined previously from Zlatarski and Martínez Estalella (2018) within fringing reefs. Seamount reefs were also added in this work as the typical formations for bank reefs. Expert interaction during workshops, followup via online meetings (Table 3.1a), and recommendations from the reviewers enabled us to define geomorphological classes to map the coral reefs in Cuba. The reef crest, fore reefs, patch reefs, and mud-bottom Guacanayabo reef classes are well established in Zlatarski and Martínez Estalella (1980), Alcolado et al. (2003), and González-Ferrer et al. (2004). Other classes related to areas with some relevant percent of coral coverage (estimated between at least 5–10% of coral coverage, dead or alive) are established as “other coral habitats” (Kramer P. review recommendation): I. Fringing reef 1. Ecological zones (a) Reef crest (b) Fore reef (c) Patch reef (d) Atoll-like reef structures (e) Other coral habitats 2. Local structures (a) Mud-bottom Guacanayabo reefs (b) Ana María back reef slopes II. Bank reefs 1. Seamount reefs New Classes

(Purdy 1974, p. 15) and “rhomboid platform atolls or faroes” by James and Ginsburg (1979). These Atoll-like reef structures in the gulfs of Ana María and Cazones seems to be remains of ancient hills and watershed and/or old relict reefs, according to bathymetry and satellite images. They are usually round or oval-shaped atolls that rise from the bottom from 20 to 25 m up to few meters deep. In Ana María Gulf, they occur on sandy or sandymuddy bottom with the development of small reef crest and patch reefs on their surface (González P., com. pers.). In Cazones Gulf, they are small hills and possible relics of small fluvial systems, with very little coral development on their surface. We do not know currently the characteristics of their slopes, although we presume that there must be coral development in them, similar to what happens in coral structures in the Ana María Gulf. The class added in this work for local structures (I.2b above) and the one described as seamount reefs (II.1) for Cuba are as follows: I.2.b) Ana María back reef slopes: They are located on areas of the inner border of Jardines de la Reina as well as Levisa and Médanos de la Vela banks. They develop as linear structures located in shallow waters emerging from the Ana María Gulf. The back reef slopes represent a reef structure that supports the thesis of barrier reefs occurring in Jardines de la Reina, Levisa and Médanos de la Vela banks. II.1) Seamounts reefs: Seamounts are geological and geomorphological structures that rise from the seafloor and remain submerged. Six seamounts or deepwater knolls, also called banks in Cuba, can be identified: (1) San Antonio Bank, west of Los Colorados and apparently not related to this archipelago; (2) the Jagua banks; (3) Bucanero; (4) Paz; (5) Silvertown, in the deep waters south of Cuba’s central region; and (6) West Bank (defined for this work), located 18 km west of Cabo Cruz. According to bathymetry, we are not sure whether the latter is a seamount or a relict from the submarine terrace system in this region as it is close to the platform border. These formations are subjected to strong currents with coral covering their top surfaces. The topmost of these banks ranges from 7 to 43 m deep but most of them are deeper than 15 m, according to bathymetry maps. There is another possible seamount inside the Cazones Gulf (Sprague et al. 2019), but we did not include it in this class, as it is shallow and connected to the coast.

I. d. Atoll-like reef structures: According to P. Kramer, in this Chapter review, these reef structures are very similar to others that occur within mixed carbonate siliciclastic lagoonal settings in the Caribbean such as southern Belize. These have been termed “shelf atolls”

3.2.4

Estimation of Metrics for Cuban Coral Reefs

Metrics for Cuban coral reefs were estimated from the thematic map produced in this article by the Estrada Estrada

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Physical-Geographic Characteristics of Cuban Reefs

et al. 2022). Using a GIS approach, we estimated the total area of the Cuban insular platform, coral cover of the platform border, total linear length of coral reefs, as well as the total coral reef area of the platform. The length of the different reef classes was calculated as per their central line, according to the Voronoi skeleton algorithm with some manual editing (QGIS 2020). All processing operations were carried out using the plane coordinate grid system UTM zone 17 north. Platform length calculations were made on the 30-m depth contour. This contour outlines the border of the platform and the upper limit of the mesophotic reefs (Reed et al. 2018). The current map does not include mesophotic reefs (>30 m depth), which are extensive around Cuba (Reed et al. 2018). Therefore, at the few sites, we identified reef structures deeper than 30 m; we discussed them in relation to their adjacent mapped habitats. We also compared the metrics from Estrada’s thematic map with those derived from prior mapping approaches (i.e., Alcolado et al. 2003). Previous maps were developed manually drawing features over hard copies. We digitized these previous maps and intersected classes with the Estrada’s map using QGIS.

Fig. 3.1 General map of Cuban reefs

55

3.3

Results and Discussion

3.3.1

Description of the Main Regional Coral Reef Groups in Cuba

Coral reef systems in Cuba are located in four main platforms with associated archipelagos. These archipelagos are Los Colorados, Los Canarreos, Sabana-Camagüey (Jardines del Rey), and Jardines de la Reina. These four large areas are connected to each other by four narrow platforms. The rest of the coral formations in Cuba are located on six seamounts outside of the four main platforms (Fig. 3.1). (a) Cuba’s northwestern platform (NW) to Los Colorados Archipelago (Fig. 3.2) Los Colorados Archipelago develops on the northwestern platform of Cuba with corals covering 165 km2 (see Table 3.2). This formation extends approximately for 280 km depending on what the origin is, whether from Pasa Balandras, Pizarro’s Bank, or San Antonio Cape (west). Los Colorados ends up in Bahía Honda (east). Los Colorados Archipelago is divided into two large sections (western and east). The western section is separated from the mainland by

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Fig. 3.2 Cuba’s northwestern platform (NW) to Los Colorados Archipelago

The Sabana-Camagüey Archipelago (SCA) originates in the northcentral portion of Cuba, one of the most studied marine areas in the country. The SCA starts in the west of the Hicacos Peninsula, Matanzas province, and finishes at the entrance channel of Nuevitas Bay, Camagüey province. The SCA spreads over 510 km (coral area, 243 km2) toward northwest-southeast orientation with high exposure to trade winds and winter front systems, with limited fetch due to the Great Bahama Bank and Cay Sal Bank. Some variations occur at smaller scales in this orientation due to the bathymetry and geomorphology of the platform. This region has reef crest, fore reefs, patch reefs, and hard ground with high coral coverage. The reef crests are usually more than 400 m from the coastline, except in the easternmost zone, where large volumes of sandy sediments make up beaches on the outer coral keys. Occasionally, patch reefs are found over sandy areas with terrestrial vegetation and associated fauna. We divide SCA into four large sectors based on geomorphological patterns, the spatial distribution of the coral formations, and its geology:

a macro-lagoon (Guanahacabibes Gulf) with depth ranging from 25 to 30 m where patch reefs are abundant. This section, together with the Sancho Pardo and Pizarro’s Bank, resembles a small barrier reef (50–75 km long) with reef crests. The east section, from Buenavista keys to Bahía Honda, presents a shallower macro-lagoon (average 3–6 m). Deeper areas in the macro-lagoon (below 15 m) are associated with submarine river valleys and riverbeds. The east section shows a more consistent geomorphology with other areas in Cuba with a seaward sequence of coral keys, patch reefs, reef crest, and fore reefs. The border of the insular shelf in the platform of Los Colorados Archipelago follows a sinusoidal curving outline. The projected portions, known as crowns, extend up to 2 km wide and have been reported as important multispecies spawning spots (Claro 2006). During recent fieldwork with echo sounding, the first author of this chapter detected a submarine terrace at around 60 m between reef crests and the border of the insular shelf. Coral formations were ubiquitous in the border of the terrace but not mapped in Estrada Estrada et al. (2022), as they were not visible from satellite images.

• Hicacos Peninsula to Bahía de Cádiz key (Fig. 3.4)

(b) Northcentral platform (NC). Sabana-Camagüey Archipelago (SCA) (Fig. 3.3)

This sector is characterized by a large development of fore reefs on submerged terraces and in the platform border. Fore

3

Physical-Geographic Characteristics of Cuban Reefs

Fig. 3.3 Cuba’s northcentral platform (NC). Sabana-Camagüey Archipelago (SCA)

Fig. 3.4 Sabana-Camagüey Archipelago (SCA). Sectors from Hicacos Peninsula to Fragoso key

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Fig. 3.5 Sabana-Camagüey Archipelago (SCA). Sectors from Frances key to Nuevitas Bay

reefs are easy to identify from satellite imagery, due to water clarity and shallow depths. Typical structures of fore reefs are clearly visible such as rocky surfaces, sand banks, and spurs and grooves. There are reef crests in different stages of development, mainly in shallow waters around small coral keys. • Bahía de Cádiz key to Fragoso key (Fig. 3.4) There is a significant decrease in the development of fore reefs after the Bahía de Cádiz key. The platform changes its orientation toward NE and returns to the predominant NW-SE orientation at the end of this sector. • Francés key to Cruz key (Fig. 3.5) North of Frances key, the platform edge is again offset to the NE (Fig. 3.5). The fore reef could not be distinguished for 40 km seaward in this area, because the slope descends gently to >30 m. Nevertheless, the submarine profile has been determined with an echo sounder, which showed three terraces. The first terrace has a rocky bottom made of calcarenites from the “Jaimanitas” formation. Small reef crests and dropping edges were detected in this first terrace. The second terrace has an elongated shape at 17–23 m deep and oriented SW-NE with coral formations interspersed with

sandy areas. The third terrace occurs between 30 and 38 m deep with sandy bottom and coral outcrops sometimes buried in the sediments. The end section of the third terrace becomes the edge of the insular shelf, with dense sand layers that increase in thickness with depth (CESAM-VC 2017). North of Los Caimanes keys, the platform widens and the edge is oriented SE, forming a deep “Puntalón” (local name for a pronounced seaward extension of the platform’s edge). This section presents reef formations but were not detected from space. From this point on, deep fore reefs are detected again, almost uninterruptedly up to Cruz key. The vast width between the outer keys and the platform edge in this sector is mainly sand but also appears to exhibit patch reefs at different stages of development, despite dynamic sediment mobility over the last decades (CESAMVC 2017). The fore reef in this sector is weakly developed and less than 300 m wide. • Cruz key to Nuevitas Bay (Fig. 3.5) This sector starts around 9 km east from the southern tip of Cruz key, north of Romano key. It is different from the previously described sectors as it has a conspicuous reef crest that protects the coastline of the nearby keys. The reef crest occupies 70 of the 90 km between Romano key and Nuevitas Bay and continues at least 20 km east of

3

Physical-Geographic Characteristics of Cuban Reefs

Nuevitas Bay, which has traditionally been considered the eastern limit of the SCA. The back zones of these well-developed reef crests are shallow waters (less than 10 m deep), with chemical-physical characteristics similar to the surrounding sea (CESAM-VC 2017) forming a typical macro-lagoon adjacent to the main island of Cuba. Although the reef here displays some barrierlike characteristics, we have not classified it as a barrier reef because of its small surface and shallow depth. The fore reef in this sector is weakly developed and less than 300 m wide

59

We defined the reef formations in the southern portion of this platform as an actual barrier reef. The main systems forming this barrier reef include Jardines de la Reina Archipelago, Médanos de la Vela and Levisa banks, as well as the portion of the platform extending up to the Cabo Cruz (250–330 km long). The macro-lagoon between the barrier reef and the mainland shows depths between 20 and 30 m and up to 60 m in the Cauto submarine canyon, which belongs to the Guacanayabo Gulf. This Gulf has associated fault systems, and clearly defined slopes in the inner part of the “barrier.”

(c) Cuba’s southeastern platform (SE) to Jardines de la Reina Archipelago to Ana María and Guacanayabo gulfs (Fig. 3.6)

• Ana María Gulf (Fig. 3.7)

With an area of approximately 385 km long from the Ancon Peninsula (west) to Cabo Cruz (east), the Cuban southeastern platform consists of Ana María Gulf and Guacanayabo Gulf. Jardines de la Reina Archipelago borders the south portion of the Ana María Gulf. Coral extends approximately 1185 km2 in the SE platform (See Table 3.1), which represents the largest reef area in the country (45% of Cuban reefs). This is the most complex platform in Cuba regarding coral formations, due to their surface extent and geomorphological characteristics.

This Gulf spreads from the Ancon Peninsula to Pingues keys to Laberinto de las Doce Leguas. The Jardines de la Reina Archipelago and the Médanos de la Vela Banks limit the southern portion of the Ana María Gulf. Turbid waters are distinctive within the gulf, and coral reefs that are visible seem to originate from hill systems and remnant river valleys. Coral keys and associated with coral formations occasionally develop in this area (Ionin et al. 1977). Where they are present, coral formations are usually round and oval shaped and have patch reefs, little reef crests, and fore reefs. Médanos de la Vela Banks show a complex geomorphological structure, with abundant rocky, sandy, and seagrass-

Fig. 3.6 Cuba’s southeastern platform (SE) to Jardines de la Reina Archipelago to Ana María and Guacanayabo gulfs

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Fig. 3.7 Ana María Gulf

covered surfaces, often with coral predominance forming patch reefs, as well as fore reefs in the outer side. On its inner north side, it seems to attach to coral structures inside some areas of the Ana María Gulf and shows small fore reefs in its inner slope (Ana María back reef slopes). The southern seaward portion of the Jardines de la Reina Archipelago presents very well-developed reef crests, patch reefs, and fore reef systems. As previously mentioned, the Ana María Gulf is characterized largely for having inner fore reefs and isolated reef crests. One of these reef crests, Nuevo Eden, was considered one of the most important coral formations in Cuba; unfortunately, this structure died a long time ago (Alcolado com.pers). • Guacanayabo Gulf (Fig. 3.8) The division between the Ana María and Guacanayabo gulfs has been traditionally defined farther east, toward Carapacho key (Instituto de Geografía de la Academia de Ciencias de Cuba 1989; Hernández-Zanuy et al. 2019). We consider that the division is at the Pingues keys to Cabeza del Este key based on the bathymetry and the submarine geomorphology. The geological fault system Pingues Keys—Cabeza del Este key—may be considered as a transition zone between Ana María and Guacanayabo gulfs. One of the most acknowledged characteristics of the Guacanayabo Gulf is the occurrence of a very peculiar and

probably unique coral structure system known as mud-bottom Guacanayabo reefs (Great Bank of Buena Esperanza) (Zlatarski and Martínez Estalella 1980; González-Ferrer et al. 2004; Zlatarski and González-Ferrer 2017; Zlatarski Greenstein 2020). This system has possibly developed over remnant reefs, associated to the Cauto riverbed, now submerged and under sediments (Ionin et al. 1977). Dr. Vassil Zlatarski and González-Ferrer (2017) discovered these mud-bottom reefs. He provided an exhaustive description of these systems: “. . . . Gran Banco de Buena Esperanza (GBBE) is a lagoonal coral reef system in the Guacanayabo Gulf, SE Cuba. . . ., in the middle of the Caribbean reefs, but it is among the last unexplored such reefs. . .. . . . GBBE has complex irregular reticulated contours (Spalding et al. 2001) delineated by 20-25 m high biogenic dams with very steep to vertical external and internal escarpments surrounding deep patios and forming a maze offering refuge for many organisms. The reef tops are flat. . . . The real reef building height is approximately 75 m because it is embedded 50 m in grey clay. It stands not on solid substratum but compact red clay dated Late Pleistocene. . . . GBBE is in a tectonic scenario with faults of Cauto depression. The GBBE reefs are approx. 10k years old. . . Presumably, hydrodynamic conditions cause the vibration of reef tops because two-thirds of the reef height is embedded in clay but the remaining one-third is vertically reaching toward the water surface. . . .”

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Physical-Geographic Characteristics of Cuban Reefs

61

Fig. 3.8 Guacanayabo Gulf

The mud-bottom reefs extend to the west after the Pitahaya channel and integrate with the coral reefs of Ana María Gulf. A similar origin is presumed for coral reef systems on both gulfs, although the muddy reefs in the Guacanayabo Gulf seem to be more related to the evolution of the western margin and the Cauto River paleo-bed and its basin, possibly more active and excavated by this river. The fore reefs on the seaward margin of the Guacanayabo Gulf are among the broadest and best developed in Cuba. These fore reefs are related to two large adjacent structures: the Cauto River’s paleo-coast to paleo-bed and the Cabo Cruz’s marine terrace system, the world’s second largest marine terrace system (Cuba 1998). One or two coastlines can be seen in the shallower fore reef, showing aligned patch reefs which constitute the inner part of the possible “barrier” to the east of the Cauto River’s submarine canyon. This canyon, the country’s largest and deepest (up to 60 m on the platform), presents various submarine river mouths and paleo-coastlines, marked by the reef geomorphology and seafloor topography. The deepest portion, over 60 m, has a submarine delta on its fore reef terrace, clearly visible in the configuration of sediments and coral formations (Fig. 3.9). This submarine delta in the Cauto Canyon is reported for the first time in this article (Fig. 3.9). Also, there are at least 2–3 outlines of fore reefs associated with submarine terraces in the seaward side of the

Fig. 3.9 Canyon and submarine delta of the paleo riverbed (Cauto River). 1 and 2: former coast lines. 3: edge of the deep terrace with sand bottoms and fore reefs at near 60 m deep

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Fig. 3.10 Cuba’s southwestern platform (SW) to Los Canarreos Archipelago

Guacanayabo Gulf. We presume more of these outlines are present, due to the proximity with the marine terrace system of Cabo Cruz (World Heritage in the Desembarco del Granma National Park), where the number of terrace levels ranges from 4 to 11 according to Iturralde-Vinet and Hine (see Chap. 4).

This sector covers the area starting in San Felipe keys, all the south portion of the Isle of Youth and finishing in Campos key. It is characterized by having poor-developed fore reefs and reef crests, increasing toward its eastern part.

(d) Cuba’s southwestern platform (SW) to Los Canarreos Archipelago (Fig. 3.10)

The main feature of this sector is a change of direction of the insular shelf associated with a fault between Campos and Ávalos keys. An important reef system dominated by crests, front reefs, and coral outcrops is observed from Cabezo of Sambo to Guano del Este key.

Los Canarreos Archipelago expands across 640 km from Punta la Yana to Punta Navajas, with a total coral area of 556 km2 (see Table 3.2). The archipelago faces the Caribbean Sea (seaward direction) and it is bordered by the Cazones Gulf and the Bay of Pigs on the east. Deep waters in the eastern side influence some environmental conditions on this part of the platform. The main units in the archipelago comprise San Felipe keys, the Isle of Youth, the Batabanó Gulf, and the Broa’s Cove (Hernández-Zanuy et al. 2019). However, from the geomorphology and coral coverage points of view, there are six defined sectors: • La Yana Point (Punta La Yana) to Cayo Campos to Cabezo del Sambo (Fig. 3.10)

• Ávalos key to Guano del Este key (Fig. 3.10)

• Guano del Este key to Guanos Point (Puntalón) (Fig. 3.11)

From the southwest to the east of Guano del Este key, there is a large platform surface (Puntalón de Guanos) with depths between 10 and 20 m, mainly rocky and sandy. This sector has a similar coral coverage and shape of that of Jagua Bank’s; hence, Sprague et al. (2019) suggested a geomorphologic relationship between the two systems. Fore reefs are prominent toward the shelf’s edge of this sector. (Fig. 3.11).

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Physical-Geographic Characteristics of Cuban Reefs

63

Fig. 3.11 Los Canarreos Archipelago. East sector and the Cazones Gulf

• Guano del Este key to Navajas Point (includes the Cazones Gulf’s east and west borders and part of the west entrance to Bay of Pigs) (Fig. 3.11)

• Jardines and Jardinillos of Batabanó Gulf (Fig. 3.11) Inside the southeastern portion of the Batabanó Gulf, there is an extensive shallow (2–5 m deep) sandy zone called Jardines and Jardinillos with very clear waters and small scattered patch reefs.

This sector show the common distribution of reef crests and fore reefs, including some of the most renowned Cuban or Caribbean reef crests (Alcolado com. pers), due to their development and conservation, such as Nirvana, practically dead today (Grau 2020); Diego Pérez, Médanos de Vizcaíno (Cayo Blanco), Piedras key, and Palmillas. Some of the factors that make this development possible are (a) the platform’s edge oriented to NW-SE against oceanic deep waters and the Cazones Gulf, where strong currents with very oligotrophic waters occur. Water carrying nutrients from the Zapata Swamp is another contributing factor to this development. (Fig. 3.11).

The inner portion of the Batabano Gulf, between the main island of Cuba and the Isle of Youth, features muddy bottoms with seagrass beds (Thalassia sp.). As a general rule, there is a lack of coral formations in this zone evidenced in the map by Estrada Estrada et al. (2022). However, some isolated small reefs could be found in the area (Rodney Borrego’s field observations).

• North of Cazones Gulf (Fig. 3.12)

(e) Narrow platforms (Fig. 3.13)

The inner area of the Cazones Gulf, or Anillo de Cazones, is surrounded by two elongated reef systems with hill-shaped coral structures alternated with sandy-muddy bottoms. Some living corals can be observed in the edges and slopes in these systems, possible remnants from fluvial systems and relict hills from the period when this region was submerged (Fig. 3.12).

The abovementioned four large Cuban platforms are interconnected by another four narrow platforms, which also have coral development, although reduced due to their narrowness.

• Inner Batabanó Gulf (Fig. 3.10)

64

Fig. 3.12 North of Cazones Gulf

Fig. 3.13 Narrow platforms in Cuba

R. E. Estrada et al.

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Physical-Geographic Characteristics of Cuban Reefs

65

Fig. 3.14 North of Havana to Matanzas

• North of Habana to Matanzas (Fig. 3.14)

– Pilón to Chivirico (Fig. 3.15)

This platform is 230 km long located at the north of Artemisa, Havana, Mayabeque, and Matanzas. It is characterized by having fore reefs along most of the platform edge. Reef crest systems develop somewhat intermittently, mainly in two sectors: Honda Bay to Mariel and Guaynabo to Puerto Escondido.

Located south of the Sierra Maestra mountains and having the Bartlett Deep to the south, this area shows steep slopes and little terrace development. Therefore, with the exception of areas like Chivirico, there is very little reef crest. In the areas without terraces, in many cases, we have not been able to identify fore reefs either, and they may not exist due to the slopes and the sediment abundance and dynamics in the area.

• East (Fig. 3.15)

– Chivirico to Caleta (Fig. 3.15)

The east narrow platform covers the eastern provinces of Cuba, Santiago de Cuba, Guantánamo, Holguín, and Las Tunas, and the southern coast of Granma and the northeastern part of Camagüey. This section has a total extension of 888 km long (437 and 451 km in the southern and in the northern coast, respectively). Due to the different geomorphic features which characterize this east narrow platform, it can be further divided into the following minor units:

In the zone of Chivirico, karst structures and terraces appear and a series of reef crests and fore reefs develop. This pattern, although with less development, remains intermittent, alternating with greater slopes and apparently without reefs until the area of Caleta.

– Cabo Cruz to Pilón (Fig. 3.15) Here, there are at least four submarine terrace levels, probably more, associated with the Cabo Cruz Marine terrace system, the world’s second largest. There is significant fore reef development on these terraces and coral reef crests near Cabo Cruz and Pilón.

– Caleta to Yumurí (Fig. 3.15) This part is associated with the Maisí terrace system, the world’s largest (Cuba 1998). We have mapped at least three submarine terraces, but there may be more, according to Iturralde Vinent and Hine (2021). The terrace system and the predominant clear waters create the conditions for fore reef development. Other elements like reef crests are not developed, although in some spots there are rocky areas with coral coverage.

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Fig. 3.15 East

– Yumurí to Ensenada de Báez Cove (Fig. 3.15) This unit, influenced by the discharge of large rivers and with little development of submarine terraces, shows very little fore reef development and reef crests only in restricted areas (e.g., Mata Bay to Cajuajo). – Ensenada de Báez to Nuevitas (Fig. 3.15) A better development of submarine terraces begins in this sector, having fore reefs and reef crests, some among the longest in the country. • Southcentral Cuba (Fig. 3.16) This narrow platform extends over 182 km from Navajas Point, in the western coast of Bay of Pigs, to the start of the Ancon Peninsula, on the southern coast of Matanzas, Cienfuegos, and Sancti Spíritus. There is a good development of fore reefs but little reef crests and few patch reefs. • South of Guanahacabibes Peninsula (Fig. 3.17) The south of the Guanahacabibes Peninsula stretches across 159 km from San Antonio Cape to La Yana Point, in

the Pinar del Río province. It is a narrow platform with good fore reef development but few reef crests. (f) Cuban seamounts (Fig. 3.18) Six seamounts or knolls, probably of bedrock origin (Sprague et al. 2019), have been mapped for the first time in this research due to their depth range (7–50 m). They are the Jagua, Bucanero, Paz, and Silvertown banks in southcentral Cuba and the San Antonio Bank, northnorthwest of the San Antonio Cape, all far from the coast but within Cuban territorial waters. We also mapped for the first time the West Bank. We are unsure whether this is an actual geological seamount or an elevated relic from the submarine terrace system. The West Bank is very close to the platform (less than 1 km), and its maximum depth of 43 m matches the depth of one of the deepest and pronounced submarine terraces south of Ensenada del Real (Vuelta Grande). The Anillo de Cazones is considered a possible seamount in line with Sprague et al. (2019). The proximity to land and shallow depths create the conditions for well-developed reef crests, patch reefs, and fore reefs. The other six seamount surfaces are mostly sandy and rocky, with some coral development and at least two fore reef systems, possibly associated with submarine terraces.

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Physical-Geographic Characteristics of Cuban Reefs

Fig. 3.16 Cuba’s southcentral platform

Fig. 3.17 South of the Guanahacabibes Peninsula

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Fig. 3.18 Cuban seamounts

3.3.2

Estimating the Metrics of Cuban Reefs

The area of the Cuban platform is estimated as 55,412 km2 (including seamounts with 169 km2). The total coral reef area of the Cuban insular platform was calculated in this study as 2645.17 km2, representing 4.77% of the Cuban insular shelf (Table 3.2) (Estrada Estrada et al. 2022). Alcolado et al. (2003) and González-Ferrer et al. (2004) estimated the coral cover for the platform border at 95 and 98%, respectively. Our estimate of coral cover for this border is 85%. We consider our coral cover is underestimated because of the limitation of using optical satellite sensors for mapping reefs deeper than 30 m (i.e., the outline of the platform border is at 30 m isobath) (the 30 m-deep isobath is approximately the reference adopted for the upper edge of our platform).

3.3.2.1 Accuracy Assessment The map’s accuracy assessment (Estrada Estrada et al. 2022) shows a figure of 81.03% for objects classified as coral that are not actually so (18.96% commission error, 81.03% accuracy of the user). In addition, it indicates a 95.51% accuracy for corals not classified as such (4.48% omission error, 95.51% accuracy of the producer). In total, the overall accuracy is 88.27%, for a total of 887 points analyzed.

3.3.2.2 Comparison with Existing Coral Reef Maps of Cuba Spalding et al. (2001) estimated a coral reef area of 3020 km2 for Cuba and placed the country in the 21st place globally (Table 3.3). Other authors such as Burke and Maidens (2005) and Burke et al. (2011) estimated the figure at 3290 and 4, 920 km2 respectively. Alcolado et al. (2003) developed the first map of Cuban coral reefs. After digitizing the map, we found that it was very difficult to establish a comparison with the base map for this chapter due to the cartographic techniques applied at the time and its low spatial resolution. The reef area for the Cuban archipelago was overestimated in Alcolado’s map at 8, 306 km2. There is a good level of coincidence (29%) between both maps (Table 3.3). The map generated by Gerhartz and Estrada Estrada (2003) which was updated for the 2021–2030 Plan for the National System of Protected Areas (Sistema Nacional de Áreas Protegidas, SNAP in Spanish) showed 29% of coincidence with our map. The differences are mainly in the wrong allocation of large parts of fore reefs and different hard ground with high coral coverage classes (Table 3.3). Huggins et al. (2007) estimated an area of 1005 km2 in their map and showed a very good level of coincidence with our map (37.9%). The main differences rely on these authors only showing reefs located in shallow areas of the platform (reef crests, patch reefs, mud-bottom Guacanayabo reefs, and

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Table 3.3 Comparison between different coral reef maps/studies for Cuba, obtained from cited sources or standardized to UTM 17 N, WGS84, when a map exists

Maps Spalding et al. (2001) Alcolado et al. (2003) Gerhartz and Estrada Estrada (2003) updated by Estrada Estrada et al. (2022) Burke and Maidens (2005) Huggins et al. (2007) Burke et al. (2011) UNEP-WCMC et al. (2018) Allen Coral Atlas (2020) Schill et al. (2021) Andréfouët and Bionaz Chap. 5 Estrada Estrada et al. (2022)

Total area (km2) 3020 8306 2619

Coincident area (km2) – 1178 763

% coincidence other maps vs. Estrada Estrada et al. 2022 – 14.2 29.1

% coincidence Estrada Estrada et al. 2022 vs. other maps – 44.5 28.8

3290 1005 4920 2697 2447 2231 5807 2645

– 552 – 1216 360 861 – 2645

– 54.9 – 45.1 14.7 38.6 – 100.0

– 20.9 – 46.0 13.6 32.6 –

Ana María Gulf’s atoll-like reef structures) in contrast with our map that presented deeper features (Table 3.3). The United Nations Environment Programme World Conservation Monitoring Centre (UNEP-WCMC et al. 2018) estimated an area of 2697 km2 and showed the most accurate coincidence with our map (46%). The fore reefs in Los Colorados, north of Villa Clara and east of Jardines de la Reina, showed major differences between both maps. We found that these coral reefs are over- or underestimated in the UNEP-WCMC maps (Table 3.3). The estimates produced by Schill et al. (2021) coincide in 36% with our map. We considered that over- and underestimations of coral formations occurred in this map, mainly on the fore reefs in Los Colorados, Guacanayabo, and Ana María gulfs, and Los Canarreos and Sabana-Camagüey (Table 3.3). We found the lowest coincidence between the Allen Coral Atlas (ACA) Project (2020) and our maps (14%). The ACA is a project that aims to map coral reefs around the globe with a consistent method using Planet satellite data (Allen Coral Atlas 2020), and it is not meant to be a surrogate for local/ regional mapping efforts. This project is planned to continue adding further innovations from 2021 onward by using satellite imagery with higher radiometric quality. We compared our map and the first version of the ACA benthic layer for Cuba and detected overestimations of coral/algae in shallow waters as well as fore reef missing across the platform. It should be pointed out that, in Gerhartz and Estrada Estrada (2003) updated by Estrada Estrada et al. (2022), the inclusion of mesophotic reefs and seamounts helps coincidences, while the noninclusion of seamounts in the other references, with exception to Allen Coral Atlas (2020) and Schill et al. (2021) that include them partially, decreases the percentage of coincidence (Table 3.3).

% coincidence mean – 29.4 29.0

37.9 45.5 14.2 35.6 –

As can be observed, most of the results show a coral area from 2400 to 3300 km2 for Cuba.

3.3.2.3 Comparison with Other Countries According to the global coral reef map by UNEP-WCMC et al. (2018), Cuba ranks 11th in the world for reef area (3101 km2, Table 3.4). However, the UNEP-WCMC values were calculated in the Mercator projection. Recalculation in UTM 17 N, WGS84 changes Cuba’s reef area to 2697 km2, without altering its ranking.

3.3.3

Barrier Reefs in Cuba

There is a historic debate whether barrier reefs occur in Cuba. In this chapter, we agreed with González-Ferrer et al. (2004) and Alcolado (2004) that some Cuban reefs can be considered true barrier reefs, such as those of the Los Colorados Archipelago and the outer edge of the southeastern Cuban platform. They share the classical barrier reef definition of elevated linear structures, separated from the coast by a relatively deep lagoon. The potential barrier of Los Colorados starts in the west sector from the Buenavista keys (around 75 km long). The southeastern Cuban platform shows a barrier beginning in the Ancon Peninsula (or from the submarine canyon of the Machos de Afuera keys) and stretching over the shallows of Médanos de la Vela, Jardines de la Reina Archipelago, and Levisa Bank. This structure could reach Cabo Cruz (i.e., 250–330 km long). Andréfouët and Bionaz (Chap. 5 in this book) define a third barrier reef associated with Moa in the east coast (Fig. 3.15). Other authors such as Zlatarski and Martínez Estalella (1980), Núñez Jiménez (1984), Spalding et al. (2001), Andréfouët and Cabioch (2011), and Goldberg

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Table 3.4 Main world country reef areas according to UNEP-WCMC et al. (2018) Country Australia Indonesia Philippines France (Outré mer) Papua New Guinea USA Saudi Arabia Fiji Madagascar Micronesia Cuba Solomon Islands Egypt Maldives Bahamas Mozambique Malaysia Tanzania India Marshall Islands Kiribati Eritrea Seychelles British Overseas Territories Japan China Tonga Mexico Colombia Belize Tuvalu Honduras Mauritius Vanuatu

Spalding et al. (2001) area (km2) 48,960 51,020 25,060 14,280 13,840 3770 6660 10,020 2230 4340 3020 5750 3800 8920 3150 1860 3600 3580 5790 6110 2940 3260 1690 5510 2900 1510 1500 1780 940 1330 710 810 870 4110

(2013) consider the existence of other barriers in Cuba, especially in Los Canarreos and Sabana-Camagüey (see Chap. 2, González-Ferrer et al. 2004). Dr. Vassil Zlatarski stated a rule that can be used to categorize barrier and fringe reefs in Cuba: “When there is a broad platform, the barrier reefs are formed and when the platform is narrow or does not exist, fringe reefs are formed. The platform’s width changes successively as per the condition between the two types of reefs. That is the reason for the transition between the two types.”

Place 2001 2 1 3 4 5 16 8 6 25 13 21 11 15 7 20 28 17 18 10 9 22 19 30 12 23 31 32 29 42 33 49 45 44 14

3.4

UNEP-WCMC et al. (2018) area (km2) 34,746 20,385 14,052 9570 7417 5638 4033 3693 3452 3232 3101 2873 2752 2710 2688 2183 2176 2140 2137 2048 1962 1704 1541 1402 1294 1244 1118 1060 982 953 918 897 836 707

Place 2018 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34

Hurricane Frequency by Sectors

According to Oliveros (2021), corals distributed over the northwestern tip comprising Los Colorados Archipelago and up to the Hicacos Peninsula are located in the zone most prone to the occurrence of extreme hydrometeorological events with a frequency of 3–5 years. Also, the same could be asserted for the southern part of this region, home to coral reefs adjacent to the keys forming Los Canarreos Archipelago (Fig. 3.19). The country’s central region, on the north zone from Cárdenas to Puerto Padre (Sabana-Camagüey Archipelago)

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Fig. 3.19 Frequency of extreme hydrometeorological events (hurricanes) by sectors in Cuba. Hurricane trajectories were compiled from the National Hurricane Center Data Archive (2021) to produce the map in this figure

and in the southern zone from Zapata Peninsula to Guacanayabo Gulf, has a hurricane return frequency between 6 and 7 years, which allows classifying this region as medium impact probability. The least affected regions by these extreme hydrometeorological events are located east of the national territory both in the north and south coasts, with a frequency higher than 7 years, and the coral reefs north of Las Tunas province, the ones that may have the least effects with a frequency or return period higher than 10 years. Overall, according to Oliveros (2021), hurricane frequency decreases from west to east across Cuba, with a return frequency ranging from 3 to 5 years in the west to 10 years in the east.

3.5

Conclusions

• The Landsat 8 and Sentinel 2 images are excellent for submarine bottom cartography, due to their complementarity and relative medium/high spatial/spectral quality

and water penetration characteristics. Also, the use of high-resolution images and maps is a very useful complement for manual work and reviewing objects, classes, and errors. • The total reef area on the Cuban insular platform is estimated, based on Estrada Estrada et al. (2022), in 2197 km2 plus 142 km2 of seamounts, for a total 2645 km2 of reefs for Cuba (4.7% of the Cuban insular platform). This figure and the results of UNEP-WCMC et al. (2018) place Cuba in 11th in world coral reef area. • We agree with previous authors stating there are barrier reefs in Cuba, one in the western portion of Los Colorados Archipelago and another in a large part of the Cuban southwestern platform. • The fore reefs, reefs located on the edges and surfaces of submarine terraces, usually below 8 m depth, constitute approximately 40% of Cuban reefs, followed by coral structures that develop in the gulfs of Guacanayabo and Ana Maria (atoll-like reef structures, 21%); other surfaces, usually rocky, with coral cover (other coral habitats,

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15%); reef crests and patches in shallow waters (less than 8 m deep, 9%); and corals on seamounts (5%). • Cuba’s main reef systems are located on four large platforms upon which four archipelagos have developed, respectively, Los Colorados Archipelago, Los Canarreos Archipelago, Sabana-Camagüey Archipelago (Jardines del Rey), and Jardines de la Reina Archipelago. The four large platforms are connected by four narrow platforms. Additional coral formations are located on the six seamounts outside the platform areas. These reef systems, except for the seamounts, are traditionally included in the Cuban regional classifications. Regarding subdivisions and borders of the reef systems, there may be less agreement, above all, if the approach tries to prioritize multifactorial results that are more clearly expressed in coral formations. We present an additional subdivision in this work. • The largest coral formations in Cuba develop on the Cuban southeastern platform (45% of total), followed by the southwestern platform (21% of total). On the basis of its complex geomorphology and evolutionary history, we consider the southeastern platform to be the most diverse and interesting in Cuba. • All the Cuban seamounts have been mapped for the first time, and an interesting group of coral reef structures are reported in the Ana María Gulf, the Cazones Gulf, and the paleo-Cauto riverbed and a delta coral mouth, the latter being reported for the first time in scientific literature.

3.6

Recommendations

• To continue improving the base map (Estrada Estrada et al. 2022), including the cartographic and thematic review, the modelling of mesophotic reefs and error analysis based on high resolution images and field points • To make the base map publicly available (Estrada Estrada et al. 2022) and expand it to consider the complete range of benthic habitats, as a basic tool for their characterization in Cuba • To examine in more detail, the least known reef areas, mainly the Cuban southeastern platform outside Jardines de la Reina, the Cauto River paleo-bed and submarine mouth, the interior of the Ana Maria and Guacanayabo gulfs, The Great Bank of Buena Esperanza, the Cuban seamounts, and some of the less transparent water areas where visibility occasionally improves sufficiently to allow remote sensing Acknowledgments To the Antonio Núñez Jiménez Foundation for Nature and Humanity, FANJ, and Wildlife Conservation Society, WCS, without their technical and logistical support, we would not have produced the map and wrote this article.

To the many experts, institutions, and persons who have supported and will continue supporting field work, the map of the Cuban sea bottoms and the article for this series, especially Sergio Lorenzo Sánchez, Patricia González-Díaz, Juliett González Méndez, José Luis GerhartzMuro, Sergio González Ferrer, and Antonio Magaz, article collaborators. To the anonymous and acknowledged reviewers, Antonio Magaz, Dr. David Blakeway, Dr. Philip Kramer, Dr. Simon F. Mitchell, and Dr. Susana Perera, for their many recommendations that improved the study. To Pedro Alcolado, friend and teacher of many the persons involved in Cuban coral studies.

References Alcolado PM (2004) Manual de capacitación para el monitoreo voluntario de alerta temprana en arrecifes coralinos. Ministerio de Ciencia, Tecnología y Medio Ambiente, Proyecto PNUD/GEF Sabana-Camagüey, Instituto de Oceanología y MINTUR. Creaciones Gráficas. 80 p Alcolado PM, Claro-Madruga R, Menéndez-Macías G, GarcíaParrado P, Martínez-Daranas B, Sosa M (2003) The Cuban coral reefs. In: Cortés J (ed) Latin American coral reefs, vol 18406. Elsevier, pp 53–75. https://doi.org/10.1016/B978-044451388-5/ 50004-7 Allen Coral Atlas (2020) Imagery, maps and monitoring of the world’s tropical coral reefs. https://doi.org/10.5281/zenodo.3833242 Andréfouët S, Cabioch G (2011) Barrera de coral (Ribbon Reef). In: Hopley D (ed) Encyclopedia of Modern Coral Reefs. Serie Enciclopedia de Ciencias de la Tierra. Springer, Dordrecht. https:// doi.org/10.1007/978-90-481-2639-2_43 Bechtel B, Ringeler A, Boehner J (2008) Segmentation for object extraction of trees using MATLAB and SAGA. In: Boehner J, Blaschke T, Montanarella L (eds) SAGA – seconds out. Hamburger Beitraege zur Physischen Geographie und Landschaftsoekologie, vol 19, pp 59–70 Bing (2022). https://www.bing.com/maps/aerial Blaschke T (2010) Object based image analysis for remote sensing. ISPRS J Photogramm Remote Sens 65:2–16 Burke L, Maidens J (2005) Reefs at risk in the Caribbean. World Resources Institute. reef-satrisk.wri.org Burke L, Reytar K, Spalding M, Perry A (2011) Reefs at risk revisited CESAM-VC (2017) Estudio de Impacto Ambiental para los proyectos de recuperación de las playas La Salina y Caracol en Cayo Las Brujas. Informe técnico, Inédito Claro R (2006) La Biodiversidad Marina de Cuba. Instituto de Oceanología, Ministerio de Ciencias, Tecnología y Medio Ambiente, La Habana Congalton RG (1991) A review of assessing the accuracy of classifications of remotely sensed data. Remote Sens Environ 37: 35–46 Conrad O, Ringeler A, Olaya V, Wichmann V (2002) SAGA system for automated geoscientific analyses (3.7.0). http://www.saga-gis.org/ Cuba (1998) Sistema de terrazas marinas de Cabo Cruz y Maisí. Parque Nacional Desembarco del Granma. Reserva Ecológica Maisí. Elemento Natural Destacado Caleta. Expediente para la declaración a la Lista del Patrimonio Mundial. 42 p. https://whc.unesco.org/ uploads/nominations/889.pdf ESA (2020). https://scihub.copernicus.eu/dhus/#/home Estrada Estrada R, Martín Morales G, Acosta JO, Hernández-Albernas JH, Martínez IA, Betancourt YC, Sánchez SL, González-Díaz P, González Méndez J (2022) Mapa de arrecifes de coral de Cuba. Versión 2022 (Inédito)

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73 National Hurricane Center Data Archive (2021). https://www.nhc.noaa. gov/data/ Núñez Jiménez A (1984) Cuba jardín coralino, Edición Turística edn, Catey, p 176 Oliveros J (2021) Informe sobre incidencia de huracanes y temperaturas en las crestas arrecifales cubana. 23. Inédito Purdy EG (1974) Karst-determined facies patterns in British Honduras: Holocene carbonate sedimentation model. Am Assoc Pet Geol Bull 58:825–855. https://doi.org/10.1306/83D914A2-16C7-11D78645000102C1865D QGIS Development Team (2020) QGIS geographic information system. Open source geospatial foundation project. http://qgis.osgeo.org Reed JK, González-Díaz P, Busutil L, Farrington S, MartínezDaranas B, Cobián Rojas D, Voss J, Díaz C, David A, Dennis Hanisak M, González Méndez J, García Rodríguez A, GonzálezSánchez PM, Viamontes Fernández J, Estrada Pérez D, Studivan M, Drummond F, Jiang M, Pomponi SA (2018) Cuba’s mesophotic coral reefs and associated fish communities. Rev Investig Mar 38(1):56–125 Roelfsema C, Phinn S (2010) Integrating field data with high spatial resolution multi spectral satellite imagery for calibration and validation of coral reef benthic community maps. J Appl Remote Sens 4(1):043527-043527-043528 Schill SR, McNulty VP, Joseph Pollock F, Lüthje F, Li J, Knapp DE, Kington JD, McDonald T, Raber GT, Escovar-Fadul X, Asner GP (2021) Regional high-resolution Benthic habitat data from planet dove imagery for conservation decision-making and marine planning. Remote Sens 13(21):4215. https://doi.org/10.3390/ rs13214215 Spalding MD, Ravilious C, Green EP (2001) World atlas of coral reefs. Prepared at the UNEP world conservation monitoring centre. University of California Press, Berkeley, USA. http://coral.unep.ch/ atlaspr.htm Sprague PJ, Iturralde MA, Escobar E (2019). Posible Rift submarino activo al sur de Cuba Central. Convención Cubana de Ciencias de la Tierra UNEP-WCMC, WorldFish Centre, WRI, TNC (2018) Global distribution of coral reefs, compiled from multiple sources including the Millennium Coral Reef Mapping Project. Version 4.0, updated by UNEP-WCMC. Includes contributions from IMaRS- USF and IRD (2005), IMaRS-USF (2005) and Spalding et al. (2001). UNEP World Conservation Monitoring Centre, Cambridge (UK). wcmc.org/ datasets/1 United States Geological Survey (2020) Landsat collection, vol 2. https://www.usgs.gov/core-science-systems/nli/landsat, https:// earthexplorer.usgs.gov/ Zlatarski VN, González-Ferrer S (2017) Gran Banco de Buena Esperanza: unique Caribbean coral reef system. Reef Encounter 32(1):60–62 Zlatarski VN, Greenstein BJ (2020) El sistema reticulado de arrecifes de coral en el Golfo de Guacanayabo, SE Cuba. Coral Reefs 39:509– 513. https://doi.org/10.1007/s00338-020-01933-7 Zlatarski VN, Martínez Estalella N (1980) Scleractinians of Cuba, with data on associated organisms. Bulgarian Academy of Sciences Press Zlatarski VN, Martínez Estalella N (2018) Los escleractinios de Cuba con datos sobre sus organismos asociados. Harte Research Institute for Gulf of Mexico Studies at Texas A&M University-Corpus Christi

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf and Uplifted Marine Terraces of Cuba: Tectonic and Sea-Level Control of Present-Day Coral Reef Distribution Manuel A. Iturralde-Vinent and Albert C. Hine

Abstract

Cuba, similar to other islands in the Greater Antilles (Hispaniola, Puerto Rico, Jamaica), has an extensive and morphologically complex coastal, shelf and upper slope system that is interconnected and has supported coral reefs over geologic timescales. Additionally, the coastal topography of the island includes up to 14 elevated and a minimum of nine submerged terraces. Herein, we review the history of these carbonate systems since the Pliocene but concentrate on the late Quaternary shelf surrounding Cuba. Additionally, we address how this region has responded to tectonic movement and repeated sea-level change, leading to subaerial exposure or flooding influencing coral reef development. Extensive uplift since the Pliocene produced the fundamental topographic features of the Cuban archipelago, but only during the Holocene was the final shape and topography established. Thus, the present Cuban shallow marine coral reef ecosystem may not be older than ~7 ka. Keywords

Cuban shelf · Coral reef assemblage · Fore reef · Mesophotic · Benthic communities · Facies · Carbonate depositional system · Karst · Red soils · Marine terraces · Paleoshorelines · Sea-level change · Tectonic movement · Marine Isotopic Stage (MIS) · Last Glacial Maximum (LGM) · Antecedent topography · Stratigraphic unit · Escarpment

M. A. Iturralde-Vinent (✉) Cuban Academy of Sciences, La Habana, Cuba e-mail: [email protected] A. C. Hine College of Marine Science, University of South Florida, St Petersburg, FL, USA

4.1

Introduction

Cuba, similar to other islands in the Greater Antilles (Hispaniola, Puerto Rico, Jamaica), has an extensive and morphologically complex coastal, shelf and upper slope system that has supported coral reefs. This complexity is due to the extreme length (~6000 km) of the coastal system, its geological history including neotectonic activity, and the island’s exposure to a wide range of paleoceanographic processes. Our present understanding of Cuba’s coral reef history is preliminary. Sea floor mapping, rock drilling, and sample analysis, providing the known coral reef geologic history, are insufficient, which allow for only broad generalizations. This chapter reviews the late Quaternary morphology and geology of the slope, shelf, and coastal areas including the record of emerged and submarine marine terraces. We synthesize the data available in order to outline the late quaternary evolution of the coral ecosystems and the origin of present-day reefs responding to repeated sea-level changes and tectonic movements leading to periodic subaerial exposure and flooding influencing coral reef development. Particularly, we concentrate on the evolution after the MIS 6 (~135 ka) sea-level low stand and the coral reef development commencing during the MIS 5.5 (~125 ka) sea-level highstand up to present-day coral reef communities. For clarity, the names of glacial events historically used (e.g., Illinoian glacial, Sangamonian interglacial, Wisconsin glacial, and Flandrian interglacial), in this chapter, have been replaced by Marine Isotopic Stages (MIS), based on oxygen isotope ratios that can be converted to sea level. We adopt this system widely used by paleoceanographers and paleoclimatologists (Spratt and Lisiecki 2016; Muhs et al. 2017). When this chapter was in editorial process, a revision of the Cuban emerged marine terraces was published (PeñalverHernández et al. 2021), where the elevation, number of terraces, age, and their uplift rate were preliminarily

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_4

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investigated. The authors did not account for some basic research on the subject such as those published by Núñez Jiménez (1959), Shantzer et al. (1975), Ionin et al. (1977), Franco-Álvarez and De la Torre y Callejas (1980), Bresznyánszky et al. (1983), Díaz-Díaz et al. (1990), Hernández-Santana et al. (1991), and Magaz-García (2019) along with many other contributions. Therefore, several inferences and conclusions about rate of uplift may be subject to questioning. It is not our task to evaluate this paper, but here, we provide one example of the problem. Along the rocky keys and small islands of the Sabana-Camaguey Archipelago, Iturralde-Vinent and Cabrera-Castellanos (1998) reported MIS 5.5 Jaimanitas Formation to occur within the flat karstified surface developed up to 3–5 m above sea level, overlain by younger uplifted paleodunes of the Guillermo Formation. These paleodunes are partially eroded elongated hills of cross-laminated eolianites, whose height reach up to 15 ± 1 m, known as the highest paleodunes in the Cuban territory. But according to Peñalver-Hernández et al. (2021), in this area occur “a single terrace raised at elevations ranging from 8 ± 1 m to 15 ± 1 m,” which contradicts previous observations.

4.2

General Geology

Mainland Cuba has a complex geological structure with rock of Neoproterozoic to Holocene age, including components of the Caribbean and North American Plates (Fig. 4.1; IturraldeVinent et al. 2016; Iturralde-Vinent 2021). In the stratigraphic section, shallow marine carbonate rocks with isolated corals are known to be of late Jurassic, late Cretaceous, and middle to late Eocene and Oligocene age (Iturralde-Vinent 2021). In contrast, shallow marine coral reef assemblages appeared from late Oligocene to present, when widespread carbonate platforms developed (Iturralde-Vinent 1969; Ionin et al. 1977; Franco-Álvarez et al. 1981, 1983; GonzálezFerrer and Iturralde-Vinent MA 2004, Medina Batista 2007). These carbonate depositional systems are ultimately linked to the plate tectonic evolution of the area in an active geodynamic scenario with the Caribbean Plate moving north and eastward, overriding the North American Plate, to ultimately colliding with Bahamas platform around middle to late Eocene. As a consequence, Cuba became attached to the North American Plate, while the Cayman Trench developed as the Caribbean Plate independently drifted eastward (Iturralde-Vinent et al. 2016). Important tectonic activity is associated with this transform plate boundary along the Cayman Trench. The northern flank of the trench, in southeastern Cuba, is associated with the sinistral transcurrent Oriente Fault, in a geodynamic scenario where deformation and large vertical displacements took place on both sides of the fault (Calais et al. 1989; Calais

and Mercier de Lépinay 1990; Rojas-Agramonte et al. 2005). The more conspicuous topographic and bathymetric relief occurs associated with this boundary, having a maximum relief of ~9660 m extending from the highest peaks—the Turquino mountain—to the deepest sea floor, the Bartlett Trough (Fig. 4.1). The tectonic position of Cuba, northwestward of the main plate boundary, determined the neotectonic evolution of the whole Cuban territory (Iturralde-Vinent et al. 2016) when the fundamental topographic features of the Cuban archipelago were determined by a general uplift since the Pliocene (Liliemberg 1973), which combined with sea-level oscillations shaped the present morphology.

4.3

Marine and Coastal Geomorphology

The Cuban territory represents a NW-SE trending mountain range, which protrudes from the sea floor within the northwestern Caribbean. The Cuban archipelago includes the Isle of Youth and roughly 4400 other small islands and keys (Núñez Jiménez 1959, 1984). A large part of the area is occupied by the shallow water shelf (Fig. 4.1), which is underlain, with few local exceptions, primarily by limestone as identified by acoustic profiling, exploratory wells (Ionin et al. 1977; PeñalverHernández 1982; Peñalver-Hernández et al. 1997; Guerra et al. 1984; Cabrera-Castellanos 1997, Cabrera-Castellanos and Ugalde 2001), and video recordings of the shelf and slope (Reed et al. 2018). The shelf can be subdivided into several sectors according to its morphology and position (Fig. 4.1). North of Cuba, it is generally narrow (150 m), which are characterized below: Deep Fore Reef (30–50 m). This is a zone of shelf-edge reefs generally parallel to the shelf-edge break. At some sites, the deep fore-reef crest forms a ridge or sill-like feature (3–5 m relief). Generally, this ridge is not continuous but features a series of mounds (5–10 m diameter, 3–5 m high). The fore-reef slope of this zone is typically steep (45–60°) with rocky escarpments and spur and groove-like structures. Deep Fore-Reef Escarpment (the “Wall,” 50–125 m). This zone forms a near vertical wall (80–90° slope) at many sites. The base of the wall commonly occurs between 100 and 125 m depths although at many sites the wall continued to >150 m and locally >200 m depths at the NE coast. In some sites, the upper part of the wall (45–80 m) forms large rock buttresses which overhang the plane of the wall by 3–5 m. The buttresses are generally 10–20 m wide and are separated by sediment chutes. Deep Island Slope (>125). This is a steep (70°), relatively smooth rock pavement slope intersected with low relief, vertical sediment chutes (1 m deep, 1–3 m wide), although the slopes may vary between 45 and 90° at 150 m. The surface rock usually displays horizontal strata with scalloped morphology (10–30 cm diameter), suggesting erosion by deep currents. Slopes of 10–45° occur to depths of 500 m (Reed et al. 2018). The general geomorphology and mesophotic zonation of the deep fore-reef slope and escarpment along the Cuban

North shelf 2 6–8 8–10 14–15 18–20 25 30–32 40 60–65

coast are strikingly similar to those described for the Bahamas (Porter 1973; Reed 1985; Ginsburg et al. 1991), Jamaica (Goreau and Land 1974; Lang 1974), and the Belize barrier and atoll reefs (James and Ginsburg 1979). It is important to underline the lateral differences between the submarine features of the shelf and slope, as reported by Ionin et al. (1977) and Reed et al. (2018). These differences may be due to lateral variations of the slope morphology (Ionin et al. 1977) and/or the result of using different equipment and methods for data acquisition. Some authors have argued that within the Cuban archipelago, late Pleistocene sea level fell only to 70 m or perhaps as much as 90 m, considering the bottom depth of blue holes and some other features observed in the shelf and slope (Núñez Jiménez 1984, 2012). Additionally, Reed et al. (2018) reported karst-like topography including caves (1–3 m in diameter) and ledges (1–2 m wide) located between depths of 60–125 m (Fig. 4.3), which match with the Last Glacial Maximum (LGM; ~26 to~18 ka; forming MIS 2) ~125 m sea-level fall (Fairbanks 1989). During hydrogeologic investigations in the HabanaMatanzas southern karstified plain, karstic porosity was found in limestone as deep as 100 m. However, deeper rocks are less karstified (Skwaletski and Iturralde-Vinent 1971), indicating that circulation of subsurface waters and karstic processes were active mostly down to this depth. This observation and those of Reed et al. (2018) are consistent with the LMG ~125 m sea-level fall.

4.3.2

The Shelf Interior

The modern shelf interior is morphologically complex, as shown by the extensive development of coral framework, rocky exposures, sand shoals, channels, and depressions. Water depths range between 0 and 12 m, but in some internal depressions can reach depths of 20–30 m (Ionin et al. 1977). The north central shelf is represented by many shoals, keys, and small islands, some of them attached to the mainland as peninsulas or separated from the north shore of Cuba

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

Fig. 4.3 Morphological zones of the Cuban shelf’s slope. (Redrawn from Reed et al. 2018)

by a very shallow marine embayment. The keys and small islands often present a flat surface where the Jaimanitas Formation is exposed, covered by a series of eolian dunes, both lithified (paleodunes) and active, produced by NE winds (Iturralde-Vinent and Cabrera-Castellanos 1998). The acoustic basement of the shelf is represented by karstified limestone overlain by red soils and marine sediments of Holocene age (Ionin et al. 1977). The outer shelf has a barrier reef and a smooth slope with outcrops of reef limestone and depressions filled with sand. Also, within the external part of the shelf margin, there is relict coral framework overgrown by younger massive corals. The southwestern shelf is also underlain by a karstified, irregular limestone surface featuring pinnacles up to 12 m in height. There are depressions and sinkholes, as well, filled by yellowish-red soils, and Holocene marine sediments, which can reach a thickness of 24 m. The shelf is rimmed by a barrier reef and internally present patch of corals, several

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shallows, keys, and small islands. Along the south coasts of Guanahacabibes, some keys northeast of the Isle of Youth, and the Zapata Peninsula up to Cienfuegos Bay, limestone of the Jaimanitas Formation occurs locally covered by eolianites and calcarenites forming partially eroded paleodune-beach complexes (Ionin et al. 1977; Franco-Álvarez and De la Torre-Callejas 1980; Cabrera-Castellanos 1997; CabreraCastellanos and Ugalde 2001). In this area of the shelf, the seaward steep slope of 45° has several submarine terraces. At the base of the slope, between 440 and 480 m, there are abundant debris and large displaced blocks, possibly induced by gravitational slope processes (landslide and marginal collapse), possibly linked to extreme wave action during storms and/or earthquakes. The southeastern shelf sector at the Gulf of Ana María has a much thicker Holocene (50–70 m) sedimentary section over a karstified limestone surface which is covered by red soils. Present living coral reefs are locally growing on top of limestone highs. The modern coral reefs located within the outer margin of the shelf are less than 10 m in relief. In the Gulf of Guacanayabo, coral frameworks overlie a sandy layer 30 m thick and are partially buried in Holocene sediments at 45–50 m depth (Ionin et al. 1977). An example of rapid coral reef growth in response to sea-level rise is the Gran Banco de Buena Esperanza (Gulf de Guacanayabo, Fig. 4.1) where the top of the modern reef is at sea level, while the base is buried by ~20 m of marine sediments. In some locations, Sangamonian (MIS 5.5) fossil coral reef frameworks are buried beneath younger marine sediments (Ionin et al. 1977). In different locations, the shelf supports submerged paleofluvial valleys as deep as 65–70 m, presumably originating during the LGM sea-level low stand. Most modern bays have also been interpreted to be former river valleys flooded by the late Holocene (MIS 1) sea-level rise (Flandrian transgression sensu Núñez Jiménez 1959, 1984, 2012). All along the rocky coastal areas, subhorizontal caves occur at the present-day active notches within the tidal zone. Similar caves, but flooded, are developed at indentations within the seaward cliff of submarine terraces, probably originated at the same paleotidal position. The same features also are found at the base of the seaward cliff of many uplifted terraces. Other common features of the Cuban shelf are blue holes. Many morphologically similar sinkholes (casimbas) occur in the surface of uplifted terraces (Núñez-Jiménez 2012).

4.3.3

Acoustic Basement of the Shelf

As indicated previously, the acoustic basement beneath the whole Cuban shelf, with local exceptions, is represented by a

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Plate 4.1 Interpretation of seismic profiles in the northwestern and southwestern shelf interior. Adapted from Ionin et al. (1977)

karstified limestone surface overlain by yellowish-red soils and marine Holocene sediments (Plate 4.1). Commonly, the top of this limestone section is reduced to a layer of soil with fragmentary limestone inclusions, due to the long period of

exposure to subaerial (karstic) weathering. In some areas, the karst process produced an irregular surface with pinnacles and depressions, which resemble residual limestone pinnacles up to 8 m high, also located on uplifted marine

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

Plate 4.2 Examples of uplifted marine constructional terraces. 1, Punta Caleta, Maisí. Notice how they wedge out laterally and note the differences in the elevation of the seaward cliff of the terraces. Several local slope collapse and landslides are visible; 2, SE of Cabo Cruz where the 70 m deep Morlote sinkhole opens in the third terrace and the lateral

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variation in the numbers of terraces; and 3, Punta Jíjira, north coast of Mayabeque, where two terraces occur, with the second partially eroded. Remnants of the second terrace are now represented by several limestone pinnacles (as “Peñon del Fraile,” located on the same surface but few kms to the east)

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Fig. 4.4 Cuban coastal features (modified and updated from Iturralde-Vinent and Serrano Méndez 2016)

terraces (Plate 4.2, insert in 3). Contrary to Ionin et al. (1977) who tentatively assigned the limestone basement to be Miocene, we propose that it is Late Pleistocene given the extensive occurrence of the Late Pleistocene Jaimanitas Formation underlying local small islands, keys, and the coastal areas of the main island (Fig. 4.4). In the northwestern and southwestern shelf areas, shallow wells revealed Pleistocene limestone thought to be the Pleistocene Vedado Formation (probably MIS 7) and Late Pleistocene Jaimanitas Formation. The Vedado Formation is also weathered and overlain by yellowish-red soil with carbonate inclusions (PeñalverHernández 1982; Cabrera-Castellanos and Ugalde 2001; Guerra et al. 1984). Among these stratigraphic units, Jaimanitas Formation is the only one with aragonite. In older units, this mineral is dissolved or recrystallized to calcite. The Jaimanitas Formation was deposited during MIS 5 (Sangamonian age according to Ducloz 1963), which lasted from about 125 to ~80 ka, when sea level was about 6.6–8.3 m higher than today (in MIS 5.5) at its peak ~125 ka (Muhs et al. 2011, 2017; Spratt and Lisiecki 2016). Three U-series ages of corals from this formation near Havana date to ~120 ka (Toscano et al. 1999). Seventeen samples taken around the Bay of Matanzas yield ESR (electron spin resonance) and U-Th series ages of ~126 ka (Schielein et al. 2020). Sixty-five uranium-series analyses of nonrecrystallized, well-preserved aragonitic corals from Guantánamo Bay yielded ages ranging from ~133 to ~119 ka, corresponding to MIS 5.5 (Muhs et al. 2017). The Jaimanitas Formation is composed chiefly of lagoon calcarenites, with local elongated coral reefs forming a massive coral framework (Brönnimann and Rigassi 1963; Franco-Álvarez et al. 1981; Iturralde-Vinent and CabreraCastellanos 1998). Perera Montero and Rojas-Consuegra

(2005) described the Jaimanitas Formation east from Havana as having small Acropora palmata coral patches, surrounded by sedimentary lagoon facies indicating that Acropora palmata were not only at barrier reefs. Muhs et al. (2017) in a study of the Jaimanitas Formation in Guantánamo Bay recognized an inner, protected, lagoonal facies, with abundant corals in growth position and an exposed offshore Acropora palmata constructional reef built on what appears to be a wave-cut platform, as well as other species of corals in growth position. Nevertheless, an extant Acropora palmata reef tens of meters long and few meters wide occurs at Orihuela keys, within the southeastern lagoon interior (Iturralde-Vinent, personal observation), confirming that Acropora palmata facies is not restricted to outer reefs. Most likely, parts of the late Pleistocene barrier reef and frontal reef components now lie offshore, underwater, or buried by Holocene sediments. According to Ionin et al. (1977), some suspected Sangamonian (MIS 5) isolated reeflike structures, buried in Holocene sediments, were revealed by seismic reflection profiling on the southwestern and southeastern shelf. Other barrier and frontal reefs may have been eroded and/or weathered during the LGM sea-level low stand. Isochronous beach-dune, cross-laminated calcarenites and eolianites, developed landward of the Jaimanitas Formation, also have been described as the Santa Fe or Guillermo Formations (Brönnimann and Rigassi 1963; Shantzer et al. 1975; Iturralde-Vinent and Cabrera-Castellanos 1998).

4.3.4

Shorelines and Coastal Areas

Cuba’s shoreline has distinctly different characteristics allowing it to be categorized into three types (IturraldeVinent and Serrano-Méndez 2016), as illustrated in Fig. 4.4.

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

The first type of shoreline consists of a smooth transition between a shallow marine muddy bottom and the inundated coastal wetland underlain by sand, clay, and peat. This shoreline morphology is highly variable and frequently changes daily and seasonally due to the combined effects of tectonic movements, tidal and climatic sea-level fluctuations, as well as extreme climatic events. Consequently, this first type of shoreline consists of seasonally inundated coastal plains. A second type of shoreline consists of rocky shores where ancient elevated marine terraces occur, whose number and elevations vary spatially. Nuñez-Jiménez (2012) has described laterally persistent fossil notches (referred as isonotches) at 3, 4, 7, 9, 11, 21, and 37 m above sea level along this type of shoreline. Due to local vertical tectonic activity and gravity collapse is difficult to correlate these notches laterally, probably with the exception of the MIS 5.5 notches. A third type of shoreline is where pre-Quaternary rocks are exposed facing a large coastal escarpment, as developed in the central part of southeastern Cuba south of Sierra Maestra mountain range (Fig. 4.4). This huge submarine to subaerial escarpment has ~9660 m of relief extending from the bottom of the deep Bartlett Trough (part of Cayman Trench) to the highest (Turquino) point of the Sierra Maestra mountains (Fig. 4.1; Iturralde-Vinent 2003, 2021; Hernández-Santana et al. 1991). Along this coastline, emerged marine terraces are absent, possibly resulting from extensive landslides linked to active tectonics and earthquakes (Fig. 4.4; Hernández-Santana et al. 1991).

4.3.5

Emerged Marine Terraces

The second type of shoreline, described above, is associated with rocky coastal areas where uplifted marine terraces are present. Interestingly, the number of the terraces as well as their maximum elevation varies laterally. According to Liliemberg (1973), the age span of the Cuban uplifted marine terraces is Pliocene to Holocene. From Mariel to Havana along the northwestern shoreline, Pliocene shallow marine limestones of the Morro Formation occur, no higher than 10 m above sea level. Late Miocene to Pliocene marine deposits also occur along the northern coast of Matanzas (Canímar Formation) around 10–15 m above sea level, and at the southern Zapata Peninsula, the Pliocene outcrops a few meters above sea level (Peninsula de Zapata Formation; Fig. 4.4). On the other hand, the only place where Pliocene rocks reach a fairly high elevation is at Maisí and Cabo Cruz areas, in southeastern Cuba (Plate 4.2; IturraldeVinent 1969, 2003), suggesting that local tectonic uplift controlled by the Oriente Transcurrent Fault System (Calais et al. 1989; Calais and Mercier de Lépinay 1990; Rojas-

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Agramonte et al. 2005) elevated these rock units at a faster rate than elsewhere. In other areas of west and central Cuba in general, one to five elevated terraces are present along the shoreline (Fig. 4.4). The differences in number and elevation are a consequence of tectonic control, depending upon their location within large neotectonic blocks limited by active faults (Iturralde-Vinent 1978). For example, the north side of the Guanahacabibes Peninsula is a wetland, while the south shore is a rocky shoreline with two elevated terraces (Fig. 4.4; Iturralde-Vinent 1978, 2003). In north Matanzas, five terraces were described; the higher (25–51 m) surface resulting from marine erosion was named the Rayonera Terrace and dated Pliocene in age (Ducloz 1963). Recent studies by Schielein et al. (2020) describe sea-level indicators for the MIS 5.5 highstand in the Matanzas area ranging from 5.5 m above sea level (reef crest) to a maximum of 7 m (notches). Usually, the lowest uplifted terrace, also the least deformed, consists of the late Pleistocene coral limestone deposits of the Jaimanitas Formation, about 10–12 m thick (Plate 4.1). At the inland wall of this terrace, a MIS 5.5 notch is commonly well preserved, 2–7 m above sea level at different localities, which is probably linked to different uplift rate in different fault-bounded tectonic blocks. The coastal areas of southeastern Cuba south of Sierra Maestra mountains present a more complex situation, because within the same time interval (Pliocene to Holocene) as many as 11–14 terraces formed, many more than in west and central Cuba (Fig. 4.4, Plate 4.3). The width and elevation of each surface change laterally, so the number of terraces changes too. For example, in Cabo Cruz, 11 terraces have been distinguished, the highest at 240 m, which represents a wide erosional surface extending landward, constructed in marls of the middle to late Miocene La Cruz Formation. Below this level are terraces (210–20 m) built on coral limestone and marls of the Pliocene Maya Formation (Bresznyánszky et al. 1983; Hernández-Santana et al. 1991). Locally, as in the Guantánamo Bay area, only two uplifted terraces are present, the higher at 39–40 m above sea level (Muhs et al. 2017). But eastward in Punta de Maisí, as many as 14 terraces have been described. One of these is the highest in Cuba (460 m) coincident with an erosional surface extending landward, formed in rocks of the middle to late Miocene La Cruz Formation. Topographically below, there are several levels of marine terraces where the Pliocene Maya and Jaimanitas Formations are developed (Bresznyánszky et al. 1983; Hernández-Santana et al. 1991). Within this large series of depositional terraces expanding from late Miocene-Pliocene to Holocene, it is questionable that only three stratigraphic units have been distinguished (La Cruz, Maya and Jaimanitas Formations; Bresznyánszky et al. 1983). This suggests that more detailed dating and stratigraphic research are needed. This is a unique place to

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Plate 4.3 1–3: Position of the Jaimanitas Formation in three coastal areas (1, Havana’s ocean front (Malecón); 2, southeastern coast, near Buey Cabón, Santiago de Cuba; 3, Punta Jíjira, Boca de Jaruco, Mayabeque. 4–5: Jaimanitas Formation; 4, plan view of back-reef lagoon

M. A. Iturralde-Vinent and A. C. Hine

biodetrital facies with Strombus gigas at Mayabeque north coast; 5, vertical view of inner lagoon, massive coral reef in growth position, Playa Baracoa quarry, Artemisa; 6, plan view of lagoon with Acropora palmata reef facies, Mayabeque north coast

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

investigate the relationships between tectonic movements and sea-level oscillations. All along the southeastern shoreline sector (with the exception of the shoreline facing the Bartlett Deep, Fig. 4.1), the lowest terrace is eroded on top of the late Pleistocene Jaimanitas Formation, uplifted to 10 and 20 m. This formation rests unconformably on older Paleogene and Miocene rocks and wedges out, onlapping toward the wall of older terraces. As illustrated by Plate 4.2, in different parts of this multi-terrace shoreline, the number of terraces changes laterally, as some narrow and disappear. Also, the height of the surface of adjacent terraces varies significantly. These characteristics are possibly resulting from sea-level changes combined with active slope processes which tilted and partially collapsed the terraces downslope (Hernández-Santana et al. 1991; Magaz-García 2019). All along the northern wall of the Cayman Trench, the steep slope consists of many irregular features as terraces, ridges, and depressions (Calais et al. 1989; Calais and Mercier de Lépinay 1990).

4.3.6

Rates of Tectonic Uplift of Pleistocene Coral Limestone

In Cuba, there are insufficient dates of late Pleistocene coral deposits, so it is difficult to determine the rate of uplift along the seashore. Unique places to investigate the relationships between tectonic movements and sea-level oscillations are Cabo Cruz and Maisí (Plate 4.2), where large series of terraces built in late Miocene-Pliocene to Holocene rocks occur. Remarkably, only three stratigraphic units have been distinguished in these sections (La Cruz, Maya, and Jaimanitas Formations; Bresznyánszky et al. 1983). In regard to this question, Peñalver et al. (2021) concluded that late Pleistocene apparent coastal uplift rates are lower in the NW of Cuba with respect to the area in front of the Oriente Transform Fault System. This conclusion is acceptable, as rocky coastal areas of the Cuban archipelago (Fig. 4.4) do not reach the elevation of Maisí and Cabo Cruz (Plate 4.2), and is supported by dates of corals as discussed below. This statement is supported by Muhs et al. (2017), who collected samples for dating in a low terrace at the Guantánamo Bay, north of the Oriente Fault Zone, and calculated the uplift rate as 0.02–0.11 m/ka for a terrace that formed ~130 to ~119 ka. They considered these figures to be a low rate and indicated that on the coasts east and west of the sample locality, there are many more marine terraces at higher elevations that could record a higher rate of uplift, implying that Guantanamo Bay may be anomalously low. West of Bay of Matanzas, Schielein et al. (2020) obtained

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lower uplift rates for the MIS 5.5 between 0.003 and 0.036 m/ ka and inside the Bay of Matanzas, 0.027–0.060 m/ka. In Haiti, on the other side of the transform fault (within the Caribbean Plate), at least seven elevated terraces have been recorded, which rise to 200 and even to 600 m above sea level. Three of them are dated to 81 ka (16 m), 108 ka (28 m), and 130 ka (52 m). Using information from the terraces dated to 81 and 130 ka, the uplift rate was calculated to be 0.35 m/ ka (Dodge et al. 1983), much higher than Guantanamo and Matanzas. In any case, more data are needed before the cause of the remarkable elevation and large amount of the eastern Cuban terraces are fully understood.

4.4

Late Pleistocene to Holocene Evolution of Coral Reef Communities

Here, we analyze the overall evolution of the coral-bearing shallow marine carbonate deposits of Cuba, which are chronologically correlated with the sea-level curve dominated by the Milankovitch cycles, particularly after MIS 7 (Fig. 4.5). This figure confirms that glacial and interglacial events were an important factor controlling the evolution of the submarine and subaerial landscape since the Pliocene. These events, in combination with tectonic movement, explain the retreats and expansions of the shallow marine environments which gave rise to the distribution of the coral communities and morphology of the present shelf (Fig. 4.1).

4.4.1

Stage Correlated to MIS 7 and 6

During MIS 7 interglacial maxima, ~190 ka, the sea level rose, and extensive shallow carbonate deposition took place within the Cuban territory, represented by the Vedado and equivalent formations (Fig. 4.4). During the succeeding sea-level oscillations leading to the MIS 6 low stand, the extent of exposed terrains gave rise to erosion, weathering, karstification, and extensive terrestrial deposition. This glacial expansion and resulting sea-level fall subaerially exposed most of the Cuban shelf to ~120 m depth, forcing shallow marine benthic communities to become reestablished further seaward leaving their earlier remains behind as part of the sedimentary fossil record (Iturralde-Vinent 2003; GonzálezFerrer and Iturralde-Vinent 2004).

4.4.2

Coral Reef Expansion During MIS 5

The MIS 6 sea-level fall established the conditions for the subsequent evolution of the Cuban landscape, as sea level

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M. A. Iturralde-Vinent and A. C. Hine

Fig. 4.5 Oxygen isotope sea-level curve adapted from Chappell et al. (1996), with the MIS 7–1, and major events related to marine and uplifted landscape transformations and their consequences for the shallow marine communities, especially corals, in Cuba

rose between MIS 6 and MIS 5.5 climate optimum (~125 ka). Marine waters flooded the present Cuban shelf area, allowing for the reestablishment of extensive coral reefs and related ecosystems (Fig. 4.6). During MIS 5.5, sea level rose up to 6.6–8.3 m higher than today (Muhs et al. 2011), covering the area of present coastal plains, where the Jaimanitas Formation was deposited (Iturralde-Vinent 2003).

4.4.3

Coral Reef Demise During MIS 2

Following the MIS 5 expansion of Cuba’s coral reef ecosystems, a progressive sea-level fall extended from about 80 to 26 ka. During this time, there were several stages of ice sheet expansion and contraction (Fig. 4.5). Shallow marine ecosystems suffered additional major stress and

Fig. 4.6 Paleogeographic reconstruction during the MIS 5.5 (~125 ka) sea-level high and land minima. Compare with Fig. 4.4 where the distribution of the Jaimanitas Formation is illustrated

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

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Fig. 4.7 Paleogeographic map of Cuba during MIS 2 Late Glacial Maximum (26–18 ka), when sea level fell ~120 m below present-day level

partial annihilation as a result of these several sea-level oscillations which occur during MIS 4 and 3 (Chappell 2002). However, this overall sea-level fall may have had little influence in the evolution of shallow marine communities of the Cuban shelf. During MIS 2 at ~26 ka, the sea level drops to ~125 m below present-day sea level, thus beginning the LGM (Carter et al. 1986; Shinn 2001; Muhs et al. 2011; Joy 2019; Praxedes Gomes et al. 2020 and references contained therein). In Cuba, this sea-level fall can be correlated with several submerged marine terraces located from near the present shore and down to 100 m water depth (Ionin et al. 1977). As a consequence of this low-stand event, the marine biota, which previously populated the Pleistocene shallow water, shifted seaward, and new reefs were established further seaward as well. Today, these older sites are more deeply submerged (Ionin et al. 1977). From the paleogeographic standpoint, Iturralde-Vinent (2003) considered this a time of “land maxima,” when Cuba became a prominent topographic high within the western Caribbean (Fig. 4.7). During MIS 2, the Cuban shelf was completely exposed to subaerial weathering (karstification and soil formation) (Ionin et al. 1977; Iturralde-Vinent 2003; González-Ferrer and Iturralde-Vinent 2004). Earlier shallow water coral reefs became exposed, and according to Ionin et al. (1977), they were nearly completely eroded or reduced to small pillars a few meters high. Examples of these features are the remnant hill named “Peñon del Fraile” and some smaller erosional remnants, consisting of recrystallized limestone of the Pleistocene Vedado Formation. These features are common within an uplifted coastal plain located in the north coast west of Santa Cruz del Norte, indicating that a large sector of the second terrace was almost completely eroded away, as illustrated by image 3 of Plate 4.3.

4.4.4

Sea-Level Rise and Initiation of Modern Reef Development

As sea-level flooding increased right after the LGM (Fig. 4.5), coral reefs became reestablished, possibly reaching their present extent during the deceleration of the late Holocene sea-level rise (Flandrian transgression) beginning around ~7 ka (Ionin et al. 1977). Transgression may have formed areas of wetlands on the shelf prior to the full marine inundation as suggested by shallow water sediments recovered from drill holes (Ionin et al. 1977). Holocene clay layers with very small mollusks, characteristic of brackish water environments, are found in several localities of the shelf. Radiocarbon dates of peat layers provide insight into the timing of inundation. Samples from the Gulf of Batabanó yield 4.710 ± 0.07 ka; from Siguanea 4.2 ± 0.12, 5.55 ± 0.15, and 6.05 ± 0.2 ka; and from north of Hicacos 12.20 ± 0.1 and 13.7 ± 0.08 ka. Some organic mud beds yield dates of 5.98 ± 0.140 and 6.5 ± 0.15 ka (Puerto Padre Bay), 5.0 ± 0.11, 5.5 ± 0.115, and 5.4 ± 0.12 ka (offshore around Isle of Youth) (Ionin et al. 1977). In the Zapata Swamp, ages of ~5 ka were obtained from peat samples near the surface (NEDECO 1959). All these ages are consistent with the post-MIS 2 sea-level rise. One exception is an 18 ka radiocarbon date obtained from a peat sample at 7 m depth in the Zapata Swamp. Theoretically, this area would have to be dry, because ~18 ka the sea level must have been ~125 m lower than present (Joy 2019). This peat can represent an inland swamp, but this date must be revisited. The flooding and subsequent coral recolonization of the present shelf created several morphological features (crests, ridges, terraces, sediment accumulations, and coral growth). These features formed because sea-level rise did not progress as a linear trend but, instead, took place through a series of

88

stillstand events followed by rapid sea-level rise pulses (e.g., Carter et al. 1986; Blanchon and Shaw 1995; Blanchon et al. 2002). Sea-level stillstands created features such as paleoshorelines, terraces, and reefs on the shelf, while a subsequent rapid sea-level rise followed by another stillstand interval created a new generation of shallow water features to form further upward across the shelf (Hine and Mullins 1983; Locker et al. 1996). In South Florida, this step-by-step rise took place on the order of 5–9 m jumps, within 200–500 year time intervals (Locker et al. 1996). If sea-level rise continued in this manner, one may assume that shoreline features on the shelf and slope must get younger upward. Therefore, in order to estimate the age of the flooded terraces, here, we plotted the position by depth of the Cuban submerged terraces in the Late Pleistocene-Holocene Gulf of Mexico sea-level curve (Joy 2019). This method is only a rough approximation, but the results suggest the possibility that some of the submerged shoreline features off Cuba may be younger than 12 ka (Fig. 4.8). However, submerged terraces can pose problems when trying to correlate age, depth, and sea level, because they may not be direct indicators of sea-level position at all. For example, there are prominent terraces, such as the Pourtalès and Miami Terraces in South Florida, formed by erosion of the strong Florida Current, a component of the Gulf Stream western boundary current (Mullins and Neumann 1979). These marine erosional terraces are generally deep (>400 m) and wide (10’s of km). In shallower water, seaward of the Florida keys reef tract, there is a laterally discontinuous terrace that is ~1 km wide Fig. 4.8 Comparison between the late Pleistocene to Holocene Gulf of Mexico sea-level curve (adapted from Joy 2019) and the topographic position of some Cuban submerged marine terraces after Ionin et al. (1977). In the graphic N: 8-10 m and S: 10-12 m, identify north and south shelf terraces and their depth

M. A. Iturralde-Vinent and A. C. Hine

lying consistently at ~35 m depth extending for ~100 km (Lidz et al. 2008; Hine et al. 2009; Shinn and Lidz 2018). Sporadically, superimposed on this terrace are coral reefs structures, called “outlier reefs,” up to 25 m in relief, yielding ages from ~106.5 to 82 ka or MIS 5.3 to 5.1 (Ludwig et al. 1996; Toscano and Lundberg 1996). Rock drilling only penetrated a portion of these 25 m high outlier reefs so their date of origin is unknown. Additionally, the age and origin of the 35 m deep terrace upon which they rest are unknown, but this terrace could be much older that MIS 5.5. It may have been formed by a much earlier free-standing reef edifice (an older outlier reef?) that had been backfilled, thus forming a terrace underlain by coral reef rock and back-reef sediments (Lidz et al. 1991, 1997). Regardless, this terrace played a key role in the emplacement of these late MIS 5 coral reefs. Previous analysis suggests that an age progression for submerged terraces from deeper (older) to higher (younger) may not always be true, and a more complicated scenario is possible. The problem is that no date is available from the submarine terraces and related features in Cuba.

4.5

Conclusions

The focus of this chapter is on the late Quaternary shelf surrounding Cuba and how this region has responded to repeated sea-level changes and tectonic movements leading to periodic subaerial exposure and extensive flooding, influencing development of the benthic marine communities and ultimately defining the present distribution of coral reef.

4

Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

Thus, this contribution may serve as an introduction to other themes presented in this book, mostly dedicated to the living coral ecosystems. Here, we summarize a large amount of data which were published, some of them in hard-to-find publications, now available in the Cuban Digital Library of Geosciences (http:// www.redciencia.cu/geobiblio/inicioEN.html). During this exercise, we became aware that more research is required, including more dates of the submarine and subaerially exposed fossil corals, terraces, paleoshorelines, and other features, needed in order to understand how the landscape, paleoceanography, and biogeography evolved during the Pleistocene and Holocene and how they gave rise to the present marine ecosystems. Although sea floor acoustic imaging and high-resolution bathymetry are being done in some parcels of the shelf, a larger coverage is required to produce better maps. Also, more continuous track-line video to target here-to-fore poorly characterized living coral reefs is needed. Such new products of the sea floor would then lead to site-specific coral reef study to include rock drilling, thus revealing coral reef development bringing about opportunities for dating and ultimately understanding paleo-coral reef response to sea level and paleoceanographic changes. This knowledge will definitely provide more effective strategies to protect this delicate ecosystem, today jeopardized by increasing global warming and climate change. Acknowledgments The authors are deeply grateful to Daniel Muhs (USGS), Simon Mitchel (UWI), Paul Blanchon, and Ilsa B. Kuffner who critically read an early draft of this manuscript and provided many important suggestions. We also thank John Reed for his advice concerning the final draft of the manuscript. Idelfonso Díaz and Reynaldo Estrada kindly supplied some of the base maps and satellite images utilized.

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M. A. Iturralde-Vinent and A. C. Hine Medina Batista A (2007) Nuevos datos estratigráficos del desarrollo de arrecifes coralinos a partir del Eoceno superior y su extensión hasta el Mioceno inferior. In: Memorias, Trabajos y Resúmenes. II Convención Cubana de Ciencias de la Tierra (Geociencias’ 2007, CD ROOM). Centro Nacional de Información Geológica, Instituto de Geología y Paleontología de Cuba, La Habana. (unpublished) (*) Muhs DR, Simmons KR, Schumann RR, Halley RB (2011) Sea level history of the past two interglacial periods: new evidence from U-series dating of reef corals from South Florida. Quat Sci Rev 30: 570–590 Muhs DR, Schweig ES, Simmons KR, Halley RB (2017) Late quaternary uplift along the North America-Caribbean plate boundary: evidence from the sea level record of Guantanamo Bay, Cuba. Quat Sci Rev 178(2017):54–76 Mullins HT, Neumann AC (1979) Geology of the Miami terrace and its paleoceanographic implications. Mar Geol 30:205–232 NEDECO (1959) Reclamation of Cienaga de Zapata. Cuba. The Hague, Netherlands. Oficina Nacional de Recursos Minerales, Ministerio de Energía y Minas, La Habana. (unpublished) Núñez Jiménez A (1959) Geografía de Cuba. Editorial Lex, La Habana. 545 p Núñez Jiménez A (1984) Bojeo. Serie Cuba: La naturaleza y el hombre. Editorial Letras Cubanas, La Habana. 681 p Núñez-Jiménez A (2012) Litorales y mares. La Naturaleza y el Hombre. Editorial Ciencias Sociales, La Habana, Cuba. 255 p Peñalver-Hernández LL, Lavandero R, Barrientos A (1997) Sistema Cuaternario. In: Furrazola-Bermúdez G, Cambra KN (eds) Estudios sobre geología de Cuba. Instituto de Geología y Paleontología, La Habana, pp 165–178. (*) Peñalver-Hernández LL, Pedoja K, Martín-Izquierdo D, Authemayou C, Núñez A, Chauveau D, de Gelder G, Davilan P, Husson L (2021) The Cuban staircase sequences of coral reef and marine terraces: a forgotten masterpiece of the Caribbean geodynamical puzzle. Mar Geol 440:1–15 Peñalver-Hernández LL (1982) Correlación estratigráfica entre los depósitos cuaternarios de la plataforma noroccidental de Pinar del Río y las zonas emergidas próximas. Ciencias de la Tierra y el Espacio 5:63–84 Perera Montero Y, Rojas-Consuegra R (2005) Distribución facial de los corales de la Formación Jaimanitas en un área al oeste de Cojímar, Ciudad de la Habana. Memorias Geomin. (*) Porter JW (1973) Ecology and composition of deep reef communities off the tongue of the ocean, Bahama Islands. Discovery 9(1):3–12 Praxedes Gomes M, Vital H, Droxler A (2020) Terraces, reefs, and valleys along the Brazil northeast outer shelf: deglacial sea-level archives? July Geo-Marine Lett. https://doi.org/10.1007/s00367020-00666-4 Reed JK (1985) Deepest distribution of Atlantic hermatypic corals discovered in The Bahamas. In: Proceedings fifth international coral reef congress, vol 6, Papeete, Tahiti, pp 249–254 Reed JK, González-Díaz P, Busutil L, Farrington S, MartínezDaranas B, Cobián Rojas D, Voss J, Díaz C, David A, Dennis Hanisak M, González Mendez J, García Rodríguez A, GonzálezSánchez PM, Viamontes Fernández J, Estrada Pérez D, Studivan M, Drummond F, Jiang M, Pomponi SA (2018) Cuba’s mesophotic coral reefs and associated fish communities. Rev Investig Mar 38(1):56–125 Rojas-Agramonte Y, Handler R, Neubauer F, García-Delgado D, Friedl G, Delgado Damas R (2005) Variation of palaeostress patterns along the Oriente transform fault, Cuba: significance for Neogenequaternary tectonics of the Caribbean realm. Tectonophysics 396: 161–180 Schielein P, Burow C, Pajon J, Rojas Consuegra R, Zhao J, Schellmann G (2020) ESR and U-Th dating results for last interglacial coral reef terraces at the northern coast of Cuba. Quat Int. https://doi.org/10. 1016/j.quaint.2019.11.041

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Outline of the Geology, Geomorphology, and Evolution of the Late Quaternary Shelf. . .

Shantzer EV, Petrov OM, Franco GF (1975) Sobre las formaciones costeras del Holoceno en Cuba, las terrazas pleistocénicas de la región Habana-Matanzas y los sedimentos vinculados a ellas. Serie Geológica 21:1–26 Shinn EA (2001) Coral reefs and shoreline dipsticks. In: Gerhard LC, Harrison WE, Hanson BM (eds) Geological perspective of global climate change, pp 251–265 Shinn EA, Lidz BH (2018) Geology of the Florida keys. University Press of Florida, Gainesville, FL. 153 p Skwaletski E, Iturralde-Vinent MA (1971) Estudio ingeniero-geológico del carso cubano. Serie Espeleológica y Carsológica 31. 58 p (*)

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5

A Remote Sensing Appraisal of the Extent and Geomorphological Diversity of the Coral Reefs of Cuba Serge Andréfouët and Océane Bionaz

Abstract

The coral reefs and lagoons of Cuba are mapped using Landsat satellite images at 30 m spatial resolution following the Millennium Coral Reef Mapping Project (MCRMP) protocol. The mapped classes include five hierarchical levels of geomorphological description, which are consistent with other MCRMP products worldwide. Cuba is characterized by a high richness of entities, with barrier, fringing, patch, and shelf complexes, each with its own range of variations due to exposure to the ocean, open shelf, lagoon, and bay environments. At Level 5, the most detailed level, Cuba is the richest Caribbean country with the presence of 108 classes. At Level 2 and 3, Cuba has 8 and 23 entities, respectively. Among them, 58 classes covering 5807 km2 are reefal classes that harbor coral and hard-bottom communities or are dominated by these communities. Other classes represent 58,142 km2 of sedimentary areas. This chapter overviews Cuban coral reefs and lagoon within the Millennium terminology and provides areal coverage statistics and maps that can be compared with other Millennium products in the Caribbean and worldwide. Keywords

Remote sensing · Geomorphology · Cuban reefs · Mapping

S. Andréfouët (✉) · O. Bionaz Institut de Recherche pour le Développement, UMR9220 (IRD, Université de La Réunion, Centre National de la Recherche Scientifique, Ifremer, Université de la Nouvelle-Calédonie), Tahiti, French Polynesia e-mail: [email protected]

5.1

Introduction: The Millennium Coral Reef Mapping Project of Cuba’s Reefs

The Millennium Coral Reef Mapping Project (hereafter, MCRMP) aimed to map coral reefs worldwide using a specifically designed hierarchical coral reef geomorphological classification scheme (Andréfouët et al. 2006). MCRMP has enhanced our understanding and knowledge of reef distribution, coral reef area, and diversity at the geomorphological scale. Since 2004, MCRMP also turned out to be particularly useful for various applications in conservation planning, risk assessment, fishery stock assessment, connectivity modelling, and inventory of other ecosystems than coral reefs (Andréfouët and Bionaz 2021). Similar applications can be targeted with the Cuban MCRMP product that is presented in this chapter, although the focus here will be on the inventory of reef geomorphological units and its diversity. The project was initially funded in 2002 by the National Aeronautics and Space Administration (NASA) to the Institute for Marine Remote Sensing (IMaRS), University of South Florida (USF), and, from 2003, by the French National Research Institute for Sustainable Development (IRD). MCRMP first used Landsat 7 Enhanced Thematic Mapper Plus (ETM+) images. Landsat 7 is part of a series of multispectral optical sensors that started in 1972. Landsat 7 was launched in 1999 and its ETM+ sensor provided 180×180 km scenes, and for the first time, space mission has been specifically scheduled and acquired images for coral reef science. Landsat 7 images have a 30 m spatial resolution and seven spectral bands, but only four (blue, green, red, and near infrared) are useful for geomorphological description of coral reefs. In 2013, the Landsat 8 satellite continued the series, with the Operational Land Imager multispectral sensor onboard. For all sites worldwide, the MCRMP unique design provides a description of coral reef geomorphological units using a five-level hierarchy (Andréfouët et al. 2006). The first

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_5

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and simplest binary Level 1 differentiates between continental and oceanic reefs. Subsequent hierarchical levels provide more detailed geomorphological description. Level 2 identifies major complexes, such as atolls, banks, barrier reef, fringing reefs, islands, etc. Level 3 provides more detail within each Level 2 category in particular introducing the differences due to exposure, with several geomorphological reef entities, for instance, “outer barrier reef,” “coastal barrier reef,” “ocean-exposed fringing reef,” and “lagoon-exposed fringing reef.” Level 4 provides a separate catalogue of coral reef geomorphological units than can be present in each of the Level 3 structures. For instance, the Level 4 unit “forereef” can be found either in the Level 3 “outer barrier reef complex” class or in the Level 3 “ocean-exposed fringing reef” class. Finally, Level 5, the most complex level, combines the Levels 1 to 4. As such, the Level 5 class: “reef flat” (L4) of an “ocean-exposed fringing reef” (L3) of a “island” (L2) “continental” (L1) is different from the “reef flat” of an “lagoon-exposed fringing reef” of a “island” “continental.” Each Level 5 entity is defined by a specific code. All polygons are also described by several attributes that indicate if they are part of the land area or not (LAND = 1, 0) and if they are hard-bottom or sedimentary dominated areas (REEF = 1, 0) and provide a qualitative indication of depth. Overall, a total of 806 Level 5 classes are defined for all coral reefs worldwide, but not all classes can occur in a single location (Andréfouët et al. 2006). Several characteristics of the MCRMP products stand out from typical remote sensing-based mapping projects and their products (Andréfouët and Bionaz 2021): • First, MCRMP follows the user-oriented approach described in Andréfouët (2008) meaning that photointerpretation and therefore simple image processing toolboxes are used. No atmospheric, sun-glint, or water column correction are needed, although they could be applied as well as preprocessing. Production can start once an image of good environmental quality is available. • Second, criteria for identification rely on shapes and positions between objects (including Land) and not on spectral properties. • Third, because the MCRMP uses a geomorphological classification scheme, it is not necessary to perform field-based ground-truthing, for both training and control of the classification process. The products were global in scope and therefore specifically defined to avoid the need for a global ground-truth data, which would be an unrealistic costly task. Ground-truthing is unnecessary due to the use of coarse geomorphological thematic classes and considering their generally large extent directly visible on Landsat images. Comparing with satellite images at higher spatial resolution than Landsat is however useful and can be used to validate some MCRMP outlines (Andréfouët et al. 2022). It is also useful to compare MCRMP with

S. Andréfouët and O. Bionaz

maps at different spatial and thematical resolutions to assess if both products are in agreement in terms of areal statistics (as demonstrated by Hamylton et al. 2012). The MCRMP minimum discernable units are about 3–4 Landsat pixels (about 3000 m2), including for patch reefs when they are individually mapped. • Fourth, while classical field-based ground-truthing to compute accuracy statistics is not required for the broad MCRMP thematic classes, how to describe and interpret a reef complex can follow multiple views, because how to assign to mapped polygons the MCRMP classes require expert opinions. Hence, qualitative disagreements can occur, but this does not correspond to a statistical measurement of accuracy. • Last, importantly, a strong criterion when mapping a MCRMP reef complex is to highlight its differences compared to other complexes. Favoring descriptions to emphasize differences is a MCRMP paradigm, because it is useful in a conservation context to identify remarkable zones or assess conservation achievements (Andréfouët et al. 2006; Gairin and Andréfouët 2020; Andréfouët and Bionaz 2021). The Caribbean coral reefs were targeted by MCRMP, and prototype products were designed early in the project in particular for the San Blas Archipelago in Panama (Andréfouët and Guzman 2005) and for Cuba. For the latter, information collected during the CUBAGRRA II August 2001 cruise that investigated the Jardines de la Reina (Gardens of the Queen) Archipelago in southeast Cuba allowed defining a first typology of Cuban reef geomorphological units that could be mapped accurately with Landsat 7. These prototypes (both typologies and maps) were also critical to finalize the global design of the MCRMP products. This design was subsequently systematically used after 2004. The current version of MCRMP products uses a typology updated in 2012. However, the mapping of Cuba using the MCRMP final classification scheme was not updated using the 2012 typology until the announcement for the present book. The following sections describe the characteristics of the recently updated Cuba Millennium product.

5.2

Remote Sensing Image Database

To update the product, a number of Landsat 8 OLI images (NIR, Green and Blue bands 5, 3, 2) were preferentially used. Images were acquired between 2014 and 2020 and were available freely from the United States Geological Survey (USGS)’s Earth Explorer portal (https://earthexplorer.usgs. gov/). Level 2 radiometrically corrected data were downloaded.

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Table 5.1 Landsat 8 images used for Cuba mapping. Images can be downloaded from one of the USGS Landsat data portal. They were selected for their high environmental quality (no wind effects, clouds, or heavy turbidity) Path 10 11 11 12 12 13 13 13 14 14 15 15 16 16 17 17

Row 46 45 46 45 46 44 45 46 44 45 44 45 44 45 44 45

Date (YYYYMMDD) 20150203 20190120 20170215 20150217 20181124 20151006 20170402 20180216 20151029 20151216 20140102 20191202 20191123 20200126 20140217 20140217

Full image ID LC08_L1TP_010046_20150203_20170413 LC08_L1TP_011045_20190120_20190201 LC08_L1TP_011046_20170215_20170228 LC08_L1TP_012045_20150217_20170412 LC08_L1TP_012046_20181124_20181210 LC08_L1TP_013044_20151006_20170403 LC08_L1TP_013045_20170402_20170414 LC08_L1TP_013046_20180216_20180307 LC08_L1TP_014044_20151029_20170402 LC08_L1TP_014045_20151216_20170331 LC08_L1TP_015044_20140102_20170306 LC08_L1TP_015045_20191202_20191216 LC08_L1TP_016044_20191123_20191203 LC08_L1TP_016045_20200126_20200210 LC08_L1TP_017044_20140217_20170307 LC08_L1TP_017045_20140217_20170307

To be entirely mapped, Cuba needs to be covered by at least 16 Landsat images. Landsat images are available according to a predefined grid, called World Reference System 2. With Landsat 8, it is possible to have up to one image every 16 days for each cell of the WRS2 grid. However, cloud cover and environmental quality of the image constrain the choice of the image selected for the mapping. Turbid areas varied from one image to another depending on the local weather conditions the days before the acquisition (rain, wind, storms, etc.), but several areas were always turbid. Table 5.1 lists the 16 cloud-free images used for the mapping, according to the WRS2 paths and rows.

mainland. At Level 2, we used several entities relevant for Cuba (Fig. 5.1):

5.3

The distribution of these entities defines the main reef ensembles around Cuba. Then, differences exist among these Level 2 entities, which are developed using the finer Level 3 description. A quick overview of the various complexes around Cuba is developed hereafter. A main characteristic of Cuba lagoons is the presence of mangrove islands at the edge of the shelf, for instance, in the Jardines de la Reina area in the south, where they are very close to the shelf slope. These islands, broadly organized as wide barrier systems, mark the limit of vast shallow lagoons. These lagoons are thus closed and protected by these islands, in contrast to the open shelf without barrier that lack islands and have gradual slopping transition from lagoons to outer shelf forereefs and slopes. Associated with the mangrove islands are large seagrass areas, which are typical of fringing coastal areas in many places worldwide. Hence, these barrier systems with fringing zonations are described here as outer shelf barrier (Level 2)/barrier-fringing complex (Level 3).

Key Structural Mapping Choices

To map Cuba using the MCRMP classification scheme, a number of choices were made to highlight local differences and specificities. These choices are made also considering a broader context than just Cuba. Guidance in the distribution of coral ecosystems and communities was provided by Zlatarski and Martinez Estalella (1982, Chap. 5) and Zlatarski (2018) who describe community profiles from 0 to up to 90 m all around Cuba. These descriptions were made in 1970–1973 but are still relevant to indicate where coral communities can be found. Whereas these observations are not ground-truthing data per se, they allowed to frame precisely the extent of various REEF = 1 polygons especially on the deeper oceanic parts. Cuba, as a whole, is considered as a Level 1 = continental system, but Isla de la Juventud is separated and considered as a Level 2 = continental island, hence a subsystem from Cuba

1. 2. 3. 4. 5. 6. 7. 8. 9.

Continental island (Isla de la Juventud) Continental fringing reefs Continental patch complex Continental outer shelf barrier Continental intra-shelf barrier Shelf marginal structures (including the lagoons) Cross-shelf area Continental banks Aquatic land features, which are coastal ponds, closed basins in mangroves, and river mouths, hence not reefal.

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Fig. 5.1 The distribution and spatial extent of the 10 MCRMP Level 2 complexes of Cuba. Land is considered as a Level 2 here. Details for each location and each Level 2 types are provided in the text

This type has been defined in the MCRMP classification for various areas worldwide (Mayotte, Papua New Guinea, etc.), but it is not very frequent. Several mapping and labelling choices were constrained by the position of these offshore mangrove islands relative to the outer shelf limit and the mainland. Barrier-fringing complexes are present also in the north but further away from the shelf than in the south. Furthermore, mangrove islands are interspersed with large islands in the Camaguey area. As such, in this area, we observe a transition from the continental fringing reefs (Level 2)/ocean-exposed fringing (Level 3) typical of the east to these barrier-fringing complex about west of Cayo Romano National Park. East of El Baga island, the barrier-fringing complex continues to Varadero where the narrow ocean-exposed fringing is again present. Between the barrier-fringing complexes of the north side and the mainland, there are also archipelagoes of mangrove islands, but these lagoonal entities are classified as intralagoon patch-reef complexes (Level 3). They have in fact little reefal area (attribute REEF = 0) and are dominated by sedimentary areas and seagrass on vast shallow terraces. These are very turbid areas, and it is not always clear in the Landsat images where the shallow seagrass terraces fringing the mainland end relative to the terraces of the patch island,

and some limits were arbitrarily defined based on dense seagrass bed limits. The north shore presents a number of embayments and lagoons that are closed by deep narrow passes and channels. Fringing and patch reefs, some reticulated, are found in these closed water bodies. Northwest Cuba is characterized by an outer barrier reef complex (Level 3), without mangrove islands, which bounds the Archipelago de los Colorados, which is itself a series of lagoonal mangrove islands connected to the mainland by fringing terraces. The outer area is bounded on its oceanic edge by numerous, fragmented coral crest sections. The southern part of this area, which forms the western tip of Cuba, is an open slopping shelf with gradient from lagoonal to shelf areas. One of the three small Cuban drowned banks is just offshore. In the northeast, another outer barrier reef complex with continuous outer reef flats and deep passages is present on the northeast in front of the mining complex of Punta Gorda. A small barrier system also exists in the northwest in Guanabo, which defines a narrow shallow lagoon. The latter is thus a continental outer shelf barrier (Level 2)/coastal barrier complex (Level 3) since these barrier reef types are used to label intermediate situation between a fringing reef without lagoon and a barrier reef separated from the mainland by a deep

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A Remote Sensing Appraisal of the Extent and Geomorphological Diversity of the. . .

lagoon. Cuba has only one occurrence of the coastal barrier complex, but they are common in many countries. South Cuba presents two vast lagoonal domains that are markedly different. In the southwest, the Batabano Gulf and the continental island of Isla de la Juventud present in particular an intra-shelf linear system of mangrove islands that we have continued on the shelf east of Isla de la Juventud and corresponding to the Archipelago de los Canarreos. This complex is parallel and distant to the outer shelf. As such, a cross-shelf area (Level 2) is also defined here, to highlight the wide channel between the Canarreos mangrove islands and the shelf. This continental intra-shelf barrier (Level 2) could be represented differently and is not reefal, but it is also unique. It also separates in the north of Isla de la Juventud the Batabano Gulf in two areas that differ by their average depth. The shelf that borders the Batabano lagoon presents different sections, from open to almost closed by mangrove islands. The tiny lagoonal patch reefs found in Batabano are very different in size and structure than the large patch reefs of the turbid Ana Maria Gulf, Caballones Channel, and Guacanayabo Gulf on the southeast of the Cuban shores. The gradient of protection offered by mangrove islands bordering the shelf in Jardines de la Reina (represented as barrier-fringing complexes) and the open shelf to the east in Guacanayabo justifies the gradual separation of the patch-reef systems from lagoonal exposed (Level 3) in Ana Maria Gulf to intra-seas exposed (Level 3). This representation in particular singles out the Gran Banco de Buena Esperanza reticulated system (Zlatarski and Greenstein 2020). Finally, to complete this overview of the Cuban main complexes, the narrow Cazones Gulf in the central south Cuban coast presents in close succession a variety of reef configurations and a unique combination of exposed and protected shelf reefs (Caballero Aragon et al. 2016).

5.4

Cuba Geomorphological Representation (Level 3 and Level 5)

Following the overview of the previous section, we detail hereafter several remarkable and representative areas and include MCRMP maps at Level 3 and 5, which are best to highlight the extent and positions of individual reef complexes, unlike Level 4 which would merge units found in different complexes (e.g., all “reef flats,” from fringing, patch, or barrier reefs would be mapped as one entity). The caption of the maps is self-explicit for Level 3, but Level 5 legends are provided through Table 5.2 below, where all Level 5 codes found in Cuba are described.

5.4.1

97

Jardines de la Reina, Ana Maria Gulf, and Guacanayabo Gulf

Figure 5.2 highlights at Level 3 the richness of southeast Cuba, with a variety of barrier, patch, fringing, and shelf systems. The map highlights the strong longitudinal variations in the distribution of the type of reef complexes. Figure 5.3 details at Level 5 the central part of the Jardines de la Reina characterized with a barrier-fringing complex and its central wide terrace, dominated here by seagrass, and with mangrove islands close to the shelf. The reefal area is really only the outer shelf and falls steeply outward. This section also shows the gradient in patch structures in the lagoon. Patch reefs in the southeast, unprotected by an open gradually slopping shelf, show more developed coral constructions than westward. Figure 5.4 details at Level 5 the reticulated patch-reef system of Gran Banco de Buena Esperanza. All cloud-free Landsat images were acquired in very turbid conditions, some much worse than others, but this is typical of the area (Zlatarski and Greenstein 2020). These turbid images highlight the numerous eddies around the reefs (not shown). Nevertheless, the outline of the subtidal and intertidal reef flats organized in ridges and the vertical deep basins that have formed in this area can be characterized.

5.4.2

Batabano Gulf and Isla de la Juventud

Figure 5.5 presents the distribution of the Level 3 complexes in the Batabano Gulf, Isla de la Juventud, and Cazones Gulf area. The area east of Isla de la Juventud is a zone where different Level 2 complexes merge, resulting in a high diversity of Level 3 structures as well. The eastern point of Batabano dives gently toward deeper water, while the Cazones Gulf, with its narrow stretch, is bordered by shelf subtidal and intertidal reefs with a gradient of exposure to the wind and waves. Inside the lagoon, there are very few reefal structures, although a huge number of tiny patch reefs are present in the east of the lagoon (not visible in Fig. 5.5). Their benthic composition is unknown but could be Halimeda macroalgae growing on old eroded framework.

5.4.3

Continental Barrier Reef in the Moa Region

A final example is provided with the barrier reef present in the Moa-Punta Gorda mining region in northeast Cuba (Fig. 5.6). The Landsat image in Fig. 5.6 shows a sediment plume exiting the lagoon. Its source is from the mining complex also visible on the image. This reef complex is a typical barrier reef, with a deep lagoon bordered by fringing

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Table 5.2 Definition of the Level 5 codes and their surface areas. All codes are also Level 1 = continental (not shown). Surface areas for each Level 5 classes are provided. C. continental, Constr. constructions, Fr. fringing L5 307

L2_ATT C. bank

L3_ATT Drowned bank

L4_ATT Drowned bank

REEF 1

km2 169.70

L5 672

L2_ATT C. outer shelf barrier

L3_ATT Outer barrier reef

487

C. island

Deep terrace

0

15.31

673

489

C. island

Oceanexposed Fr. Oceanexposed Fr.

Forereef

1

38.56

707

C. outer shelf barrier C. outer shelf barrier

Outer barrier reef Coastal barrier reef

491

C. island

Reef flat

1

4.09

710

493

C. island

43.59

713

C. island

0

26.35

714

511

C. island

Shallow terrace Shallow terrace Diffuse Fr.

0

509

Oceanexposed Fr. Oceanexposed Fr. Lagoonexposed Fr. Diffuse Fr.

0

46.74

718

525

C. patch

C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. island

528

C. patch

531

C. patch

532

C. patch

535

C. patch

536

C. patch

C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier

Coastal barrier reef Coastal barrier reef Coastal barrier reef Coastal barrier reef Oceaexposed Fr. Barrier-Fr. reef Barrier-Fr. reef Barrier-Fr. reef Barrier-Fr. reef Barrier-Fr. reef

538

C. patch

539

C. patch

540

C. patch

541

C. patch

545

C. patch

549

C. patch

550

C. patch

551

C. patch

552

C. patch

Coastal/Fr. patch Coastal/Fr. patch Coastal/Fr. patch Coastal/Fr. patch Patch land Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef Intralagoon patch reef

REEF 1

km2 777.23

1

174.96

1

2.81

1

2.58

Pass

0

0.57

Pass reef flat

1

0.38

Subtidal reef flat Subtidal reef flat Channel

1

5.24

1

7.51

0

21.27

Enclosed lagoon Forereef

0

17.15

1

80.17

Pass

0

38.41

Reef flat

1

16.47

L4_ATT Shallow terrace w/constr. Subtidal reef flat enclosed lagoon w/constr. Forereef

Enclosed basin Forereef

0

4.37

799

1

1.20

719

Reef flat

1

68.09

722

Shallow terrace Land on reef

0

44.07

723

0

455.27

724

Channel

0

1.24

725

Deep terrace w/ constr.

1

382.73

726

C. outer shelf barrier

Barrier-Fr. reef

Shallow terrace

0

3078

Enclosed basin

0

16.59

727

C. outer shelf barrier

Barrier-Fr. reef

1

109.68

Enclosed lagoon

0

37.37

796

C. outer shelf barrier

Barrier-Fr. reef

Shallow terrace w/ constr. Deep terrace

0

19.30

Enclosed lagoon w/ constr. Forereef

1

40.55

806

C. outer shelf barrier

Barrier-Fr. reef

Subtidal reef flat

1

3.63

1

0.52

740

C. Fr.

Oceanexposed Fr.

Deep terrace

0

80.54

Pinnacle

1

16.52

741

C. Fr.

Oceanexposed Fr.

0

0.87

Reef flat

1

37.03

742

C. Fr.

Oceanexposed Fr.

Enclosed lagoon or basin Forereef

1

769.75

Shallow terrace

0

1123

743

C. Fr.

Oceanexposed Fr.

Pass

0

35.75

Shallow terrace w/ constr.

1

62.85

744

C. Fr.

Oceanexposed Fr.

Reef flat

1

156.22

(continued)

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Table 5.2 (continued) L3_ATT Intralagoon patch reef Intra-seas patch reef Intra-seas patch reef Intra-seas patch reef

L4_ATT Subtidal reef flat

REEF 1

km2 0.06

L5 745

L2_ATT C. Fr.

L3_ATT Oceanexposed Fr.

L4_ATT Reticulated Fr.

REEF 1

km2 8.88

Deep terrace w/ constr. Enclosed basin Enclosed lagoon

1

44.86

746

C. Fr.

193.33

11.36

748

C. Fr.

Shallow terrace Deep terrace

0

0

0

8.04

0

519.36

749

C. Fr.

Oceanexposed Fr. Intra-seasexposed Fr. Intra-seasexposed Fr.

0

12.66

C. patch

Intra-seas patch reef

1

11.90

750

C. Fr.

Intra-seasexposed Fr.

1

7.60

563

C. patch

1

37.07

752

C. Fr.

1

8.33

C. patch

Pinnacle

1

0.79

754

C. Fr.

69.00

C. patch

Reef flat

1

197.40

760

C. Fr.

Shallow terrace Reef flat

0

568

1

8.00

569

C. patch

0

58.20

761

C. Fr.

Reticulated Fr.

1

55.70

570

C. patch

1

20.15

762

C. Fr.

Shallow terrace

0

1332

571

C. patch

1

291.04

763

C. Fr.

1

7.61

764

C. Fr.

Bay-exposed Fr. Diffuse Fr.

30.94

C. patch

Bayexposed Fr. Diffuse Fr.

1

574

0

684.41

586

C. patch

Shallow terrace Shallow terrace w/ constr. Subtidal reef flat Deep terrace w/ constr. Reef flat

Intra-seasexposed Fr. Intra-seasexposed Fr. Lagoonexposed Fr. Lagoonexposed Fr. Lagoonexposed Fr.

Reef flat

567

Intra-seas patch reef Intra-seas patch reef Intra-seas patch reef Intra-seas patch reef Intra-seas patch reef

Enclosed lagoon w/ constr. Forereef

Enclosed lagoon or basin Forereef

1

11.50

769

C. Fr.

Channel

0

2.43

589

C. patch

Subtidal reef flat

1

5.46

776

Forereef

1

567.83

590

C. intrashelf barrier C. intrashelf barrier C. intrashelf barrier C. intrashelf barrier C. intrashelf barrier C. intrashelf barrier C. intrashelf barrier C. outer shelf barrier

Barrier land

Land on reef

0

214.76

777

Exposed shelf reef

Reef flat

1

23.87

Outer barrier reef

Channel

0

0.71

779

Sheltered margin reef

Forereef

1

34.97

Outer barrier reef

Pass

0

13.72

780

Sheltered margin reef

Reef flat

1

5.44

Outer barrier reef

Reef flat

1

3.53

781

Shelf terrace

Shelf terrace

0

3817

Outer barrier reef

Shallow terrace

0

322.15

782

Shelf terrace

Shelf terrace w/ constr.

1

141.17

Outer barrier reef

1

22.24

783

Shelf structure

Pass

0

26.44

Outer barrier reef

Shallow terrace w/ constr. Subtidal reef flat

1

16.23

784

Shelf structure

Shelf hardground

1

566.37

Barrier land

Land on reef

0

1551

785

Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures

Fr. of barrier-Fr. Exposed shelf reef

Shelf structure

Subtidal reef flat

1

189.82

L5 553

L2_ATT C. patch

556

C. patch

557

C. patch

558

C. patch

559

592

601

603

604

605

606

657

Intra-seas patch reef Shelf patch reef Shelf patch reef Shelf patch reef

(continued)

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S. Andréfouët and O. Bionaz

Table 5.2 (continued) L5 658

659

661

662

663

664

666

668

670

671

672

L2_ATT C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier C. outer shelf barrier

L3_ATT Outer barrier reef

L4_ATT Barrier reef pinnacle/patch

REEF 1

km2 6.88

L5 786

L3_ATT Shelf slope

L4_ATT Shelf slope

REEF 0

km2 1737

Shelf slope

Undetermined envelop

0

6.79

C. lagoon

Deep lagoon

0

14,700

C. lagoon

Shallow lagoon

0

29,451

C. lagoon

64.54

Cross shelf

Shallow lagoon w/ constr. Cross shelf

1

793

L2_ATT Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Shelf marginal structures Cross shelf

Outer barrier reef

Channel

0

2.95

787

Outer barrier reef

Deep terrace

0

167.27

788

Outer barrier reef

Deep terrace w/ constr.

1

168.05

790

Outer barrier reef

Enclosed basin

0

27.59

791

Outer barrier reef

Enclosed lagoon

0

2.09

0

60.98

Outer barrier reef

Forereef

1

234.44

797

C. Fr.

Oceanexposed Fr.

Subtidal reef flat

1

39.11

Outer barrier reef

Pass

0

66.54

798

C. outer shelf barrier

Coastal barrier reef

Deep terrace

0

0.72

Outer barrier reef

Reef flat

1

47.29

800

Shelf structure

Deep terrace

0

83.11

Outer barrier reef

Shallow terrace

0

113.89

1000

Shelf marginal structures Mainland

Mainland

Mainland

0

124,333

Outer barrier reef

Shallow terrace w/ constr.

1

777.23

1001

Aquatic land features

Aquatic land features

Aquatic land features

0

732.36

lagoon-protected reef flats. It is however different than the other barrier reef complexes found all around Cuba because of its narrow lagoon (Fig. 5.1).

5.5

5.6

Discussion

5.6.1

Geomorphological Diversity of the Coral Reefs of Cuba

Areal Coverage Statistics

Since its design in ~2000, the primary goal of MCRMP was to provide accurate statistics on coral reef coverage, which was a major priority for national managers at the dawn of the new millennium as this information was seldom available for most countries (Andréfouët and Bionaz 2021). Surface areas are provided hereafter in km2, computed using the WGS 1984 Word Mercator projection and ArcGIS 10.8 software. The geographical information system digital product allows to query interactively any polygon or specific groups of polygons. Here, the main ensembles at Level 2 (Table 5.3) and Level 3 (Table 5.4) are summarized. Statistics for all Level 5 classes are provided in Table 5.2.

In the Caribbean, Cuba appears as the richest complex with 108 Level 5 classes (not including the mainland and the aquatic land features). In contrast, the Honduras-BelizeMexico (southeast Yucatan coast and Chinchorro Bank) ensemble represents 99 Level 5 classes. Bahamas and Turk and Caicos together had 49 classes; the Dominican Republic provides 85 classes. Finally, the Lesser Antilles Arc, from Anguilla to Grenada, reaches 69 classes. It is worth pointing out that, globally, Fiji holds the highest richness with 282 classes. In Cuba, 58 classes are reefal classes (attribute REEF = 1) meaning they are likely to harbor coral and hard-bottom communities (such as gorgonian plains) or are dominated

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Fig. 5.2 Map of the geomorphological units mapped at Level 3 in the Ana Maria Gulf, Jardines de la Reina, and Guacanayabo Gulf region

by these communities. Altogether, they cover 5807 km2. Other classes are sedimentary areas, in particular the vast gulfs representing 58,142 km2. These numbers can be compared with the previous figures of 3020 km2 of “reef area” by Spalding et al. (2001). Although the topics addressed here are coral reefs and not mangroves and seagrass, both constrain the interpretation of many coral reef features, in particular, as explained above, the mangrove islands at the edge of the shelf (Claro et al. 2001). The present inventory has not discriminated mangroves and seagrass because these entities are not part of the MCRMP classification scheme, but this would be a logical extension of this work. Mangroves along the mainland are not identified here, but all mangroves found in the lagoonal and barrier reef areas are. They could closely match the patch land (L5 = 535) and barrier land (L5 = 590) classes (Table 5.2). Similarly, seagrass beds are not mapped here, but they are mostly present on the diffuse fringing and terrace Level 4 classes of the lagoon-exposed patch and fringing Level 3 classes. They are also extensive in the shallow lagoon class. Wabnitz et al. (2008) have already used MCRMP data and this contextual information to enhance seagrass mapping in various Caribbean locations, and Cuba could be added to add spatial knowledge to the existing body of work on Cuban seagrass (Martínez-Daranas and Suárez 2017).

5.6.2

Potential for Coral Reef Management Applications

The recent review on how MCRMP products were used for coral reef science and management worldwide (Andréfouët and Bionaz 2021) provides examples (and caveat reminders) on how to use the products. It is beyond the scope of this chapter to provide such examples for Cuba, and we do not have necessarily on hand the relevant ancillary data, but some common applications can be cited. First, MCRMP data have been useful for the stratification of field data collection and data gap analysis, for instance, to define representative monitoring and sampling sites for UNESCO World Heritage Areas (Andréfouët and Wantiez 2010). This application is relevant for areas of the size of the Level 2 complexes, for instance, by selecting stations in each of the Level 5 classes, possibly combined by a filter on REEF = 0 or 1. Second, it is likely that the MCRMP product will be useful to assess how well conservation efforts capture the whole diversity of Cuban marine area (Perera Valderrama et al. 2017). Assessment of the representation of different habitats in network of MPAs were common at the time to assess the fulfillment of the 2020 Aichi Targets. New targets are now issued nationally and internationally, and similar assessments

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Fig. 5.3 Map of the geomorphological units mapped at Level 5 in the central Jardines de la Reina region. The Landsat image used for the mapping is shown above as a true-color composite

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Fig. 5.4 Map of the geomorphological units mapped at Level 5 in the Gran Banco de Buena Esperanza region. The Landsat image used for the mapping is shown above as a true-color composite. Note the high turbidity of this region

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Fig. 5.5 Map of the geomorphological units mapped at Level 3 in the Batabano Gulf region

of protection scores per habitat types will likely occur in the near future. MCRMP products can be used for this purpose, at each of the different hierarchical levels (Gairin and Andréfouët 2020). Third, the diversity and richness of reef structures quantified by MCRMP at different levels can be used as a surrogate of biological diversity, under the rationale that higher biodiversity will be found in areas of high structural complexity (or higher number of MCRMP classes). While this assumption is difficult to prove, as it would require extensive biological ancillary datasets, examples of such study exist (Andréfouët and Guzman 2005). The three above examples of conservation and monitoring applications are inherently closely related to the geomorphological richness and surface area metrics provided here (Tables 5.3 and 5.4). Fourth, MCRMP data have been used for coral reef fishery stock assessment for selected commercial species (Andréfouët and Bionaz 2021). For Cuba, knowledge of the extent of lagoon areas, possibly refined by knowledge of specific benthic habitats (e.g., seagrass), can be used to infer national stock of, for instance, queen conch resources. This, however, requires field data on this species density throughout the country in selected places, stratified using the MCRMP product (first application above).

Fifth and last example is represented by the enhancement of connectivity modelling using habitat data, such as in Van Wynsberge et al. (2017) for giant clams in the Pacific Ocean. Previous larval dispersal and connectivity modelling around the Cuban region, for instance, for snapper and lobster populations (Paris et al. 2005; Garavelli et al. 2018), could take benefits of the full thematic resolution of the MCRMP data to refine source and sink locations.

5.6.3

Future Coral Reef Mapping Efforts

Future detailed coral reef mapping work should serve targeted research and management questions, and there are myriads of opportunities for Cuba, with the possibility to combine with in situ surveys like the frequently visited Jardines de la Reina (Pina-Amargós et al. 2008; FigueredoMartin et al. 2010; Hernández-Fernández et al. 2011, 2019; Ferrer et al. 2016; Gonzales-Diaz et al. 2018). Higherresolution benthic mapping could target the protected areas (Perera Valderrama et al. 2017; Navarro-Martínez et al. 2020) and assess changes in these areas, as well as impacted areas due to development for mining, aquaculture, and tourism. Finally, because the mapping of Cuban reefs was completed in two steps, a preliminary one using Landsat

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Fig. 5.6 Map of the geomorphological units mapped at Level 5 in the Moa region of northeast Cuba

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Table 5.3 Surface areas for the Level 2 classes present in Cuba

Level 2 Continental bank Continental fringing Continental intra-shelf barrier Continental island Continental outer shelf barrier Continental patch complex Cross shelf Shelf marginal structures

km2 170 3503 593 182 6737 3508 61 51,415

Table 5.4 Surface areas for the Level 3 classes present in Cuba

Level 3 Barrier land Barrier-fringing reef complex Bay-exposed fringing Coastal barrier reef complex Coastal/fringing patch Continental lagoon Cross shelf Diffuse fringing Drowned bank Exposed shelf reef Fringing of barrier-fringing complex Intra-lagoon patch-reef complex Intra-seas-exposed fringing Intra-seas patch-reef complex Lagoon-exposed fringing Ocean-exposed fringing Outer barrier reef complex Patch land Shelf patch-reef complex Shelf slope Shelf structure Shelf terrace Sheltered margin reef

km2 1766 3384 31 12 118 44,215 61 731 170 592 2 1719 106 1192 1422 1394 2168 455 25 1744 866 3958 40

7 images collected between 1999 and 2003 and the present effort based on Landsat 8 images from 2014 to 2020, a comparison using the images available in the MCRMP archive suggests significant changes along the barrier reefs of the south coast between the two periods. These are immediately visible for mangrove islands, which have severely shrunk (hence, it impacts the land classes and not the reefs or lagoons per se), and more subtly on the shelf areas but not to the point that the geomorphology has changed. These changes are probably explained by hurricane damage, but they warrant further investigations using the history of hurricane tracks in each area. MCRMP products cannot be used for change detection applications, and different products will be needed to assess changes at decadal or yearly time scales. Acknowledgments The Millennium Coral Reef Mapping Project was initially funded by NASA grants NAG5-10908 to SA and Frank Muller-

Karger (University of South Florida) and grant CARBON-0000-0257 to Julie Robinson (NASA). The project is now funded by Institut de Recherche pour le Développement for specific application and projects. We are indebted to the PIs and to Christine Kranenburg, Damaris Torres-Pulliza, Alan Spraggins, Brock Murch, and Chuanmin Hu at USF for their time and hard work during the initial phase of project (2002–2007). The initial geomorphological classification of Cuba took advantage of the information collected during the CUBAGRRA II 2001 cruise organized by Robert Ginsburg, Pedro Alcolado, and Philip Kramer in the frame of the AGRRA (Atlantic and Gulf Reef Rapid Assessment) program. We warmly thank Vassil Zlatarski for the invitation to contribute to this book.

References Andréfouët S (2008) Coral reef habitat mapping using remote sensing: a user vs producer perspective. Implications for research, management and capacity building. J Spat Sci 53:113–129

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Andréfouët S, Bionaz O (2021) Lessons from a global remote sensing mapping project. A review of the impact of the Millennium Coral Reef Mapping Project for science and management. Sci Total Environ. https://doi.org/10.1016/j.scitotenv.2021.145987 Andréfouët S, Guzman HM (2005) Coral reef distribution, status and geomorphology/biodiversity relationship in Kuna Yala (San Blas) archipelago, Caribbean Panama. Coral Reefs 24:31–42 Andréfouët S, Wantiez L (2010) Characterizing the diversity of coral reef habitats and fish communities found in a UNESCO World Heritage Site: the strategy developed for Lagoons of New Caledonia. Mar Pollut Bull 61:612–620 Andréfouët S, Muller-Karger FE, Robinson JA, Kranenburg CJ, TorresPulliza D, Spraggins SA, Murch B (2006) Global assessment of modern coral reef extent and diversity for regional science and management applications: a view from space. In: Proceeding of 10th international coral reef symposium, pp 1732–1745 Andréfouët S, Paul M, Farhan AR (2022) Indonesia’s 13558 islands: a new census from space and a first step towards a One Map for Small Islands Policy. Mar Policy 135:104848 Caballero Aragon H, Alcolado PM, Rey-Villiers N, Perera Valderrama S, González Méndez J (2016) Coral communities condition in varying wave exposure: the gulf of Cazones, Cuba. Rev Biol Trop 64:79 Claro R, Reshetnikov YS, Alcolado PM (2001) Physical attributes of Coastal Cuba. In: Claro R, Lindeman KC, Parenti LR (eds) Ecology of the marine fishes of Cuba. Smithsonian Institution Press, Washington, DC, pp 1–20 Ferrer VM, González-Díaz SP, Hernández-Fernández L, Siciliano D, Bretos F, Appril A, Hughes K, Santoro A (2016) Salud de las comunidades de corales en arrecifes de Jardines de la Reina Golfo de Ana María, región surcentral de Cuba. Rev Investig Mar 36:34–53 Figueredo-Martin T, Pina-Amargós F, Angulo-Valdés J, GómezFernández R (2010) Buceo contemplativo en Jardines de la Reina, Cuba: caracterización y percepción del estado de conservación del área. Rev Investig Mar 31:23–32 Gairin E, Andréfouët S (2020) Role of habitat definition on Aichi Target 11: examples from New Caledonian coral reefs. Mar Policy 116: 103951 Garavelli L, White JW, Chollett I, Chérubin LM (2018) Population models reveal unexpected patterns of local persistence despite widespread larval dispersal in a highly exploited species. Conserv Lett 11:e12567 Gonzales-Diaz P, Gonzales-Sanson G, Betancourt CA, Fernandez SA, Perez OP, Fernandez LH, Rodriguez VMF, Caballero YC, Almanza MA, de la Giardoa LE (2018) Status of Cuban coral reefs. Bull Mar Sci 94:229–247

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Hamylton S, Andréfouët S, Spencer T (2012) Comparing the information content of coral reef geomorphological and biological habitat maps, Amirantes Archipelago (Seychelles), Western Indian Ocean. Estuar Coast Shelf Sci 111:151–156 Hernández-Fernández L, Guimarais M, Arias R, Clero L (2011) Composición de las comunidades de octocorales y corales pétreos y la incidencia del blanqueamiento del 2005 en Jardines de la Reina, Cuba. Rev Mar Cost 3:77–90 Hernández-Fernández L, González de Zayas R, Weber L, Apprill A, Armenteros M (2019) Small-scale variability dominates benthic coverage and diversity across the Jardines de La Reina, Cuba coral reef system. Front Mar Sci 6:747 Martínez-Daranas B, Suárez AM (2017) An overview of Cuban seagrasses. Bull Mar Sci 94:269–282 Navarro-Martínez ZM, Crespo CM, Hernández-Fernández L, FerroAzcona H, González-Díaz SP, McLaughlin RJ (2020) Using SWOT analysis to support biodiversity and sustainable tourism in Caguanes National Park, Cuba. Ocean Coast Manag 193:105188 Paris C, Cowen R, Claro R, Lindeman K (2005) Larval transport pathways from Cuban snapper (Lutjanidae) spawning aggregations based on biophysical modeling. Mar Ecol Prog Ser 296:93–106 Perera Valderrama S, Hernández Ávila A, González Méndez J, Moreno Martínez O, Cobián Rojas D, Ferro Azcona H, Milián Hernández E, Caballero Aragón H, Alcolado PM, Pina Amargós F, Hernández González Z, Espinosa Pantoja L, Rodríguez Farrat LF (2017) Marine protected areas in Cuba. Bull Mar Sci 94:423–442 Pina-Amargos F, Hernández L, Clero L, González-Sanson G (2008) Características de los hábitats coralinos en Jardines de la Reina, Cuba. Rev Investig Mar 29:225–237 Spalding MD, Ravilious C, Green EP (2001) World atlas of coral reefs. Prepared at the UNEP World Conservation Monitoring Centre. University of California Press, Berkeley, USA Van Wynsberge S, Andréfouët S, Gaertner-Mazouni N, Tiavouane J, Grulois D, Lefèvre J, Pinsky ML, Fauvelot C (2017) Considering reefscape configuration and composition in biophysical models advance seascape genetics. PLoS One 12:e0178239 Wabnitz CC, Andréfouët S, Torres-Pulliza D, Müller-Karger FE, Kramer PA (2008) Regional-scale seagrass habitat mapping in the Wider Caribbean region using Landsat sensors: applications to conservation and ecology. Remote Sens Environ 112:3455–3467 Zlatarski V (2018) Investigations on mesophotic coral ecosystems in Cuba (1970–1973) and Mexico (1983–1984). CICIMAR Océanides 33:27–43 Zlatarski VN, Greenstein BJ (2020) The reticulate coral reef system in Golfo de Guacanayabo, SE Cuba. Coral Reefs 3:509–513 Zlatarski VN, Martinez Estalella N (1982) Les Scléractiniaires de Cuba avec des données sur les organismes associés. Académie bulgare des Sciences, Sofia. 472 p

Part IV Biota

6

Macrophytes Associated with Cuban Coral Reefs Ana M. Suárez and Beatriz Martínez-Daranas

Abstract

Macrophytes constitute an important component of coral reef communities. This chapter will review the knowledge of macroalgae and marine angiosperms associated with Cuban coral reefs over the last 50 years. Most frequent macrophytes associated with Cuban coral reefs are mentioned, including those from Cuba’s mesophotic zone. Five hundred seventy-four infrageneric taxa of macrophytes have been found on Cuban coral reefs: five species of marine angiosperms (Phylum Tracheophyta), 192 Chlorophyta, 301 Rhodophyta, and 76 Phaeophyceae. Several methodologies have been used to study macrophytes on Cuban coral reefs which make it difficult to perform comparisons. Despite this variety of methodologies, it is evident that macroalgae abundance is high in many Cuban coral reefs, mainly in zones with high anthropogenic impacts, as in La Habana. The algal species are distributed from the intertidal to the mesophotic depths. In shallow protected areas with soft bottoms such as reef lagoons, marine angiosperms and Chlorophyta species dominate. Higher species richness of Rhodophyta and Phaeophyceae occurs on rocky substrates. Macroalgae abundance tends to be higher at shallow fore reefs than at crests and deeper fore reefs. Most investigations have been descriptive, and many gaps need should be included in future research. Keywords

Macroalgae · Seagrasses · Diversity · Abundance

A. M. Suárez · B. Martínez-Daranas (✉) Centro de Investigaciones Marinas/Universidad de La Habana, La Habana, Cuba e-mail: [email protected]; [email protected]

6.1

Introduction

Macrophytes constitute an important component of coral reef communities. Macroalgae of different taxa can be found from the rocky intertidal to mesophotic depths (Suárez et al. 2015). They can colonize rocky substrates (i.e., epilithon), rooting in sediments with rhizoids (rhizobenthos) or growing on sand grains (epipsammon) (Round 1981). Many macroalgae can live attached to other organisms such as plants (epiphyton) (Jover et al. 2020) or animals (epizoon) (Reyes-de Armas 2016; Alfonso Sánchez et al. 2020) and are capable of boring into rocks (endolithon) (Lukas 1974; Round 1981). Seagrasses (marine angiosperms) dominate shallow waters in zones with relatively stable sediments as a result of their rhizome-roots system. These plants are important components in reef lagoons and patch reefs surrounding ecosystems. Macroalgae have a great diversity of morphologies and life strategies, regardless of their phylogenetic affinities. Their thalli vary from simpler forms (delicately filamentous and foliose) to intermediate complexity (corticated fleshy and thick leathery) and higher complexity (articulated calcareous and encrusting forms). Their physiological strategies also vary, from higher net primary production rates in foliose and filamentous forms to lower productivity in calcareous algae and crustose forms, due to the allocation of resources for defenses against herbivores and other physical impacts (Littler et al. 1983; Steneck and Dethier 1994). As a result of these strategies, macroalgae can compete for resources (space, light, nutrients) among them and with corals and other organisms of the reef. Highly competitive and opportunistic species can dominate over corals under high nutrients and low herbivory levels (Littler and Littler 2005). Macroalgae play important roles in coral reefs as a consequence of their contribution to the reef productivity, along with boring algae and symbiotic unicellular algae within hermatypic corals, providing the base for trophic webs.

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_6

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From the structural point of view, calcareous crustose macroalgae are important contributors to both the bulk and frame structures of most limestone structures of coral reefs (Ionin et al. 1977; Littler and Littler 1994). Articulated calcareous green and red algae largely contribute to the reef and lagoon sediments (Littler 1976; Ionin et al. 1977; HillisColinvaux 1980; Harney et al. 2000). On the contrary, boring algae play an important role in reef bioerosion (Littler and Littler 1994). Crustose coralline algae are among the first colonizers of uncolonized hard substrata, along with algal turf (Adey and Vassar 1975; McClanahan 1997), and can induce the coral’s larval settlement and metamorphosis (González-Ferrer et al. 2004b; Martínez-Estalella 2018; Ramos-Romero et al. 2019). Cuban macroalgae have been studied on coral reefs, seagrass meadows, and other habitats for decades. Several authors studied the taxonomy of marine flora, including reef areas. The first paper concerning macroalgal ecology in reef zones is based on samplings in La Habana between 1975 and 1976, where the species richness and similarities of the community were studied between 0 and 15 m depth (Suárez and Cortés 1983). In this study of marine macrophytobenthos, the main taxa on Cuban marine biotopes were the coralline algae with the genus Halimeda (Bryopsidales, Chlorophyta) dominating in coral reefs (Suárez 1989). Three main associations were highlighted on coral reefs: Caulerpa ambigua + Laurencia intricata + Jania rubens; Caulerpa ambigua + Bryopsis pennata + B. plumosa; and Dictyota bartayresiana + Halimeda opuntia. Seagrasses are frequent in Cuban reef lagoons, and they provide essential habitat for mobile coral reef organisms such as fishes and crustaceans that use them as feeding and refuge areas, supporting high biodiversity (Unsworth et al. 2018). Extensive meadows, composed of marine angiosperms, can be found in shallow coastal waters on almost all continents. Cuba possesses a relatively broad extent of seagrass spatial area (McKenzie et al. 2020). These meadows play essential roles in the coastal ecosystem processes. They have high primary production rates contributing to the base of trophic webs and yielding organic matter that exports to other areas and ecosystems. As a result, they support fisheries worldwide (Duarte et al. 2013; Nordlund et al. 2016; Baisre 2018). Seagrass meadows may enhance coral reef resilience to climate change due to several ecological functions (Nordlund et al. 2016). The canopies of seagrasses are efficient at filtering particles, including plastics (Sanchez-Vidal et al. 2021) and bacterial pathogens, out of the water column (Lamb et al. 2017). They also prevent sediment erosion and deposition on coral reefs with their rooting system and canopies. Recent evidence indicate that they may also enhance coral reef resilience to future ocean acidification by increasing pH and the aragonite saturation state of seawater (Unsworth et al. 2012).

A. M. Suárez and B. Martínez-Daranas

Studies of macrophytes associated with coral reefs have been undertaken in several zones of the Cuban shelf. Cuban marine shelf has been divided into nine districts, according to their ecological characteristics (Atlas Nacional de Cuba 2019) (Fig. 6.1). The most studied zones are the Jardines de la Reina Archipelago (District II), Los Canarreos (IV), the south of the Guanahacabibes Peninsula (V), Los Colorados (VI), the north coast between Artemisa and Matanzas provinces (VII), Sabana-Camagüey (VIII), and some small areas of north oriental zone of Cuba shelf (IX) (Fig. 6.1). More recently, the region south of Guamuhaya Mountain mass (III) has received more attention on macroalgae inventory in coral reefs (Suárez et al. 2015). While in several papers macrophytes were the research target (e.g., Suárez and Cortés 1983; Valdivia and de la Guardia 2004; ZúñigaRíos et al. 2012; Semidey Ravelo 2013; Zúñiga-Ríos 2016; Ramos-Romero et al. 2019; Alfonso Sánchez et al. 2020), in others, they are included only as indicators of the monitoring of coral reefs’ condition (e.g., Alcolado et al. 2001, 2003, 2010, 2013; González-Díaz et al. 2003; Caballero and de la Guardia 2003; Caballero et al. 2007, 2009). This chapter provides an overview of macroalgae and marine angiosperms associated with Cuban coral reefs from the scientific literature published over the past 50 years. The rocky intertidal and the rocky terrace zones are included in the coral reef zones, following the criteria of González-Ferrer et al. (2004a) and Zlatarski (2018a). The number of species of macroalgae by the substrata they colonize and by ecological districts of Cuban marine shelf is detailed (Atlas Nacional de Cuba 2019). The most frequent species reported at different zones are listed. The ecological role of macroalgae and seagrasses for coral reefs is also discussed, with special emphasis on the knowledge gaps and recommendations of future research.

6.2

Species Richness and Distribution

In a literature review by Suarez and Martínez-Daranas (2020), a comparison of the marine phycoflora in zones of the western Tropical Atlantic and the subtropical Atlantic ranged from 36 to 76% similarity. The phycoflora of Cuba was 73% similar to that of Mexico’s coast along the Gulf of Mexico and the Caribbean Sea. However, this study did not distinguish species from the different marine habitats. Five hundred and seventy-four infrageneric taxa of macrophytes have been found associated with Cuban coral reefs. These include 192 green macroalgae (Chlorophyta), 301 red (Rhodophyta), 76 brown (Ochrophyta, Phaeophyceae), and five species of marine angiosperms (Phylum Tracheophyta). They have been found from the rocky intertidal to 169 m on the deep island slope (Table 6.1), although depth data is not available for every species.

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Fig. 6.1 Map of Cuba with the nine ecological districts of Cuban marine shelf. I, southeastern; II, south-central (Jardines de la Reina Archipelago); III, south-central (Guamuhaya Mountains); IV, southwestern (Los Canarreos Archipelago); V, southwestern (Corrientes inlet); VI, northwestern (Los Colorados Archipelago); VII, northwestern

(Habana-Matanzas); VIII, north-central (Sabana-Camagüey Archipelago); and IX, northeastern. BSA, Banco de San Antonio; GBBE, Gran Banco de Buena Esperanza. Modified from Atlas Nacional de Cuba (2019)

The total number of macroalgae species reported from Cuban coral reefs represents 91.7% of the infrageneric taxa reported for the Cuban shelf, due to the high affinity of most seaweeds to hard substrata and high frequency of epiphytism. Macroalgae have the capability of colonizing different kinds of substrata in Cuban coral reefs besides limestone. Two species of the genus Ostreobium (Chlorophyta) have been found boring hard substrates (Lukas 1974); 298 species are epiphytic, 87 epizoic, 403 epilithic, and 74 rhizophytic (Chlorophyta only), and 271 have appeared on more than one substrate, showing their ability for colonizing different organisms and objects (Fig. 6.2). Many epiphytes and epizoic macroalgae have small thallus, so they are usually

disregarded in coral reef monitoring efforts, despite their importance in primary productivity and trophic webs, and coral reef functioning (Connell et al. 2014; Cetz-Navarro et al. 2015; Short et al. 2015b; Speare et al. 2019). Regarding the zonation of macroalgae on the Cuban coral reefs, 159 species have been documented in rocky intertidal, 140 in reef lagoons, 78 species in the back reef, and only seven at the crest of coral reefs. Hernández-Fernández et al. (2013) reported 35 macroalgae and two seagrass species associated with fragile, branching colonies of sedimentresistant scleractinians, with predominance of the hybrid Acropora prolifera along with A. cervicornis at the Gran Banco de Buena Esperanza (Appendix). These coral reefs have a singular reticular structure with vertical ridges with planar tops rising 20–25 m from the soft bottom (Zlatarski and Greenstein 2020) (Fig. 6.1). A recent survey of the mesophotic coral reefs around Cuba added so far 42 macroalgae species to Cuban deep waters, with high coverage of macroalgae at the Banco de San Antonio (MartínezDaranas et al. 2018; Reed et al. 2018; Chap. 15, Reed et al.). The number of species reported for coral reefs is greatest at the Habana-Matanzas and Sabana-Camagüey districts (VII and VIII) and lowest at the south of the Guamuhaya Mountains (III) and the south of Guanahacabibes Peninsula

Table 6.1 Number of infrageneric taxa of macrophytes by phylum and depth on Cuban coral reefs. Shallow = 0–15 m; moderate = 15–30 m; deep = 30–170 m Phylum/depths Rhodophyta Ochrophyta (Phaeophyceae) Chlorophyta Tracheophyta Total

Intertidal 86 29

Shallow 277 75

Moderate 84 25

Deep 50 13

47 0 162

178 5 535

83 0 192

59 0 122

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Fig. 6.2 Macroalgae species richness (S) by phylum and the substrata they colonize on Cuban coral reefs. Boring algae species are not included

(V) (Figs. 6.1 and 6.3). The first two areas have received more sampling effort than others have, which could explain these differences. Five species of marine angiosperms are reported in Cuban reef lagoons with Thalassia testudinum (Fig. 6.4a) dominating (Zayas et al. 2006; Martínez-Daranas et al. 2007, 2014, 2016; Gómez González et al. 2019). Although seagrasses are more light limited than many algae (Duarte 1991), four of them have been found at shallow to moderate depths in sandy plains of the fore reefs: T. testudinum (15 m), Halophila engelmannii (15 m), Halophila decipiens (25 m), and Syringodium filiforme (20 m) (Buesa 1975; de la Guardia et al. 2001). The most frequent macrophytes recorded in different zones of Cuban coral reefs are listed in Appendix. Some species can be found from the intertidal and shallow waters to the mesophotic zone. Halimeda is one of the most important genera for sediment production (Martínez-Daranas et al. 2013) and appears with a high frequency from the reef lagoon to the mesophotic coral reefs (Fig. 6.4b, c). In areas of soft bottom in reef lagoons, marine angiosperms and rhizophytic Chlorophyta species proliferate, creating seagrass meadows. Simultaneously, higher species richness of Rhodophyta is found on rocky substrates (Fig. 6.4d). Among reds, many filamentous Ceramiales are frequent as epiphytes and epizoic (Alfonso Sánchez et al. 2020; Jover et al. 2020). Crustose coralline algae (mostly of the subclass Corallinophycidae) and the order Peyssonneliales (Rhodophyta) are frequent in all coral reef zones (Fig. 6.5a, b), but their identification in situ is exceptionally challenging. As a result, they remain little studied. The same situation is for the order Ralfsiales (Phaeophyceae) present at the rocky intertidal zone. The class Phaeophyceae have fewer species than the other two phyla in Cuba and the Greater Caribbean

zone in general (Suárez et al. 2015). Still, species of the order Dictyotales (Dictyota, Lobophora, Stypopodium) (Fig. 6.5c) dominate in hard substrates, whereas the order Fucales (Sargassum, Turbinaria) (Fig. 6.5d) is widespread (Appendix).

6.3

Abundance of Macroalgae on Coral Reefs

Van den Hoek (1969) early hypothesized that intensive grazing by herbivorous fishes and echinoids is necessary to maintain the health of coral reefs. This author states that under-grazing might cause degradation of the coral reef since unhampered growth of fleshy algae would cause the death of crustose corallines by covering them and the death of corals by entrapping sediments. Before 1980, Caribbean coral reefs usually presented dominance of corals and coralline algae, while filamentous and fleshy benthic algae were inconspicuous, appearing only as low patchy turfs and scattered sprigs (Bula-Meyer, pers. comm. 1996; van den Hoek 1969; Wanders 1977; Van den Hoek et al. 1978). Later, coral reefs worldwide have endured phase shifts to alternate, degraded assemblages from coral to macroalgae dominance due to the combined effects of overfishing, declining water quality, and the direct and indirect impacts of climate change (Hughes et al. 2007; Bruno et al. 2009; Martínez-Rendis et al. 2016). Increases in nutrients from land-derived eutrophication promote the fast growing of opportunistic filamentous and leafy algae over the slowergrowing corals, so the latter become endangered by competition for space and light. Additionally, the herbivores’ activity has been diminishing due to overfishing and to the massive die-off of the black sea urchin, Diadema antillarum (Philippi 1845) in

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Fig. 6.3 Macroalgae species richness at Cuban coral reefs (S) by phylum and by the ecological districts where they were reported. I, southeastern; II, south-central (Jardines de la Reina); III, south-central (Guamuhaya Mountains); IV, southwestern (Los Canarreos Archipelago); V, southwestern (Corrientes inlet); VI, northwestern (Los Colorados Archipelago); VII, northwestern (Habana-Matanzas); VIII, northcentral (Sabana-Camagüey Archipelago); and IX, northeast

1983, resulting in the collapse of the top-down control over algae (Hughes et al. 1987; for more information on this topic, see Chap. 11, Duran et al.). The demise of corals creates open space for macroalgae colonization and hinders coral recruitment (Littler and Littler 1988; Szmant 2002; Hughes et al. 2007). For those reasons, the abundance of macroalgae has been frequently used in coral reef monitoring, and Cuba is no exception. A variety of studies have assessed the abundance of macroalgae and their possible impacts on Cuban coral reefs Fig. 6.4 Examples of common Tracheophyta, Chlorophyta, and Rhodophyta on Cuban coral reefs. (a) Thalassia testudinum mixed with Syringodium filiforme in a shallow reef lagoon; (b) Halimeda incrassata with T. testudinum in a reef lagoon; (c) Halimeda spp. on a mesophotic coral reef, 80 m deep (distance between the two lasers = 10 cm); and (d) Amphiroa beauvoisii mixed with A. tribulus and Anadyomene sp. as epiphyte at 15 m deep. (Photo credits Angel Fernández Medina except for c courtesy of John Reed)

(>40 scientific publications). Unfortunately, few studies on Cuban coral reef ecology were done before 1980. Macroalgal ecology studies of Cuban coral reefs began in the 1980s using the relative abundance based on Braun-Blanquet’s phytosociological methods (Prado Díaz and Suárez 1997). This method detects differences in the structure of macrophyte communities. Another method assessed species biomass by scraping all vegetation from a determined area. This method allows detecting all species’ abundance regardless of their size,

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Fig. 6.5 Examples of common Rhodophyta and Phaeophyceae of Cuban coral reefs. (a) Crustose coralline algae at 0.5 m deep; (b) a rock with a crustose coralline and an unknown green alga at 169 m deep (distance between the two lasers = 10 cm); (c) Lobophora sp. at 15–18 m deep; and (d) Sargassum hystrix on a slope at 18 m deep. (Photo credits Angel Fernández Medina except for b courtesy of John Reed)

except for most crustose algae, and overestimates calcareous algae’s abundance. As a comparison, total macroalgal biomass in Sabana-Camagüey’s coral reefs in 1994 (ZúñigaRíos et al. 2012) was several orders higher than those obtained in La Habana in 1975–1976 (Suárez and Cortés 1983) and in the SW of Cuba in 1987–1988 (MartínezDaranas et al. 2016). In the 1990s, several monitoring coral reef methods, such as Reef Check and Atlantic and Gulf Rapid Reef Assessment (AGRRA), were implemented in Cuba. These monitoring methods assess the cover of total macroalgae by morphofunctional groups (Littler et al. 1983) or lower categories (Alcolado et al. 2013; Caballero et al. 2013; PereraValderrama et al. 2016). These monitoring methods allow rapid assessments but lose detailed information on macroalgae. Other authors assess the abundance of macroalgae from lineal transects or the point count transect methods that permit more detailed data (de la Guardia et al. 2003; Semidey Ravelo 2013; Ferrer Rodríguez et al. 2016; González Sánchez 2016). Those investigations also differ in different coral reef zones and depths where the surveys were conducted and the way the sample units were located. These

varieties of methodologies make difficult comparisons among zones and dates. Despite these differences in methodology, it is evident that macroalgae abundance is high in many Cuban coral reefs. Fleshy algae can be abundant in zones with high anthropogenic impacts as near contaminated rivers and bays, sites with tourism development, or with uncontrolled fisheries (Table 6.2). But high cover of seaweeds can be also found in areas far away from anthropic development (Table 6.2). Fleshy macroalgae abundance tends to be higher at shallow fore reefs (around 5 m deep) than at crests and deeper fore reefs (de la Guardia et al. 2006; Caballero et al. 2009; Alcolado et al. 2010). The dominant macroalgal cover on many of the Cuban coral reefs are of the order Dictyotales (Canistrocarpus, Dictyota, Lobophora, Stypopodium) and the genera Halimeda and Sargassum. Opportunistic algae such as Cladophora, Microdyction, and Ulva (Chlorophyta) dominate in some sites of Los Canarreos and Sabana-Camagüey archipelagos (Alcolado et al. 2001; Zúñiga-Ríos et al. 2012). These communities are quite different from the results obtained in other areas of the Caribbean during the 1960s

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Table 6.2 Macroalgae cover in several Cuban coral reefs. In parenthesis, the ecological districts (see Fig. 6.1) Locality Guantanamo Bay (I) Jardines de la Reina Archipelago and Golfo de Ana María (II) Jardines de la Reina Archipelago (II) Los Canarreos Archipelago (IV)

Date July–Aug/1996 Feb/2015

Mean macroalgae cover (%) Macroalgae: 50–78 Macroalgae: 6–70

Nov/2017 May–Sept/1999

Macroalgae: 11–42 Macroalgae: 17–86

Los Canarreos Archipelago (IV)

May/1999

Los Canarreos Archipelago (IV)

March/2001

Los Canarreos Archipelago (IV) Guanahacabibes Peninsula (V) Guanahacabibes Peninsula (V) La Habana (VII)

Artemisa to Matanzas provinces (VII)

June/2012 July/1999 Sept/2004 Oct/2000–Apr/ 2001 May–July/2002 Nov/2000–Apr/ 2002 June– Aug/2004

Fleshy macroalgae: 52–87 Fleshy macroalgae: 18–33 Fleshy macroalgae: 0–22 Macroalgae: 46–60 Fleshy macroalgae: 7–65 Macroalgae: 47–95 Fleshy macroalgae: 82 Macroalgae: 70 Fleshy macroalgae: 5–39

Artemisa to Matanzas provinces (VII) Camagüey, Santa Lucía (IX)

May/2016 Apr–Oct/2008

La Habana (VII) Rincón de Guanabo, La Habana (VII)

a

Macroalgae: 97–99 Fleshy: 28–40 Algaea: >60 Macroalgae: 55–65

Source Chiappone et al. (2001) Ferrer Rodríguez et al. (2016) Hernández-Fernández et al. (2019) De la Guardia and González Díaz (2002) Alcolado et al. (2001) Alcolado et al. (2010) Alcolado et al. (2013) Alcolado et al. (2003) Caballero et al. (2007) Caballero and de la Guardia (2003) González-Díaz et al. (2003) Castellanos et al. (2004) Caballero et al. (2009) Duran et al. (2018) Busutil et al. (2011)

Including Cyanobacteria

and 1970s, when filamentous and coralline algae dominated (van den Hoek 1969; Wanders 1977; Van den Hoek et al. 1978). Nevertheless, as baseline data on the natural state of reefs is not available, “great care should be taken when using information about the baseline-range in one location to make inferences about the degree of degradation in another” (sic. Bruno et al. 2013). The abundance and community structure of macroalgae depend on many abiotic factors and biotic interactions. Although the term “turf algae” is controversial, it is frequently used in coral reef ecological studies. Macroalgae of the three phyla, Cyanobacteria, and diatoms have been included in turfs (Connell et al. 2014; Cetz-Navarro et al. 2015). Turfs have been defined by Connell et al. (2014) as short or “low-lying” epilithic algae with different heights (0.5–10 cm) and may refer to filamentous (corticated and uncorticated), foliose, and calcareous articulated. In Cuba, several studies on coral reefs use this term, but few define it. De la Guardia et al. (2006) include filamentous, corticated, and calcareous articulated macroalgae lower than 1 cm. AGRRA methodology has been followed in Cuba, where turf is defined as “multispecific assemblages of red, green, and brown algae and cyanobacteria, usually short (75% of corals bleached (Durán et al. 2018).

15.5.1.2 Temporal Trends of Environmental Factors The physical environment related to Cuban coral reefs has changed in recent years when compared to two decades ago: seawater is warmer (X2 = 190.18, p < 0.001) and coral reefs have been influenced by fewer hurricanes (X2 = 223.65, p < 0.001) (Fig. 15.4). Habitats around the coral reef, rugosity, and chronic wave exposure have not changed significantly through time. This is the first analysis ever published about sea surface temperature and wave exposure (chronic and acute) on coral reef sites nationwide. Our results on acute wave exposure and chronic wave exposure match those of Cuba documented in a Caribbean-wide study (Chollett et al. 2012). Sea surface temperature was warmer, almost 1 °C, in 2019 when compared to 1988. It is ecologically relevant and higher than in other studies (Chollett et al. 2012). On the other hand, hurricane frequency has been lower in recent years when compared 2010s with previous decades, despite the increase of sea surface temperature. However, lower hurricane frequency does not necessarily mean less impact on coral reefs: there is substantial spatial variation in damage following

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Fig. 15.2 Temporal trends in the density of selected components of coral reef biota in Jardines de la Reina Horizontal lines indicate medians. Boxes represent interquartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

hurricanes (Hughes and Connell 1999; Lugo et al. 2000) related to several factors including hurricane intensity and time elapsed since the previous disturbance (Highsmith et al. 1980; Woodley et al. 1981).

15.5.1.3 Temporal Trends of Anthropogenic Factors Although many of the anthropogenic factors studied here have not changed through time (population, tourism, pollution, protection, integrate impact), fishing pressure decreased Fig. 15.3 Temporal trends of selected components of coral reef biota in Sabana-Camagüey Horizontal lines indicate medians. Boxes represent inter-quartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

(rank scale, X2 = 43.55, p < 0.001; quantitative, X2 = 167.66, p < 0.001) and enforcement increased in Cuba (X2 = 110.68, p < 0.001) (Fig. 15.5). Analysis by zones revealed that enforcement kept the same trend in Jardines de la Reina (X2 = 75.23, p < 0.001) and SabanaCamagüey (X2 = 41.75, p < 0.001) as in Cuba nationwide, but fishing pressure has decreased in Jardines de la Reina (quantitative, X2 = 108.21, p < 0.001) and has not changed significantly in Sabana-Camagüey (quantitative, X2 = 10.21, p = 0.050) (Fig. 15.6).

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291

Fig. 15.4 Temporal trends of selected physical environmental factors in Cuban coral reefs. SST, sea surface temperature; Wt, wave period; Wh, wave height. Horizontal lines indicate medians. Boxes represent inter-quartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

15.5.2 Environmental and Anthropogenic Factors Related to the Status of Cuban Coral Reefs 15.5.2.1 Environmental Factors Related to the Status of Cuban Coral Reefs Of the 180 correlations between environmental factors and the Cuban coral reefs biota, 9 (5%) were significant and of high absolute value. All were with benthos (7, 78%) and small fish (2, 22%), and most of them (7, 78%) were related to water movement (acute and chronic wave exposure). Habitat is a main factor determining the characteristics of benthic and fish communities in Cuba (González-Sansón et al. 2009a, b; González-Díaz et al. 2010) and elsewhere (Chittaro et al. 2005; Dorenbosch et al. 2007). While D. antillarum is consistently more abundant in reef crests (X2 = 77.91, p < 0.001) as other studies have found (see Chap. 12 Durán et al.), coral density differentiation is not consistent between shallow reefs (reef crest) and deep reefs (drop off and spur and groove). Coral density is higher on deep reef in this chapter (X2 = 53.89, p < 0.001), in Jardines de la Reina (Pina-Amargós et al. 2008a), Havana (GonzálezSansón et al. 2009b), and the Northwestern zone (GonzálezDíaz et al. 2010). Whereas coral density is similar at both depths in Sabana-Camagüey (González-Ferrer et al. 2007a) and in the southern archipelagos (Caballero-Aragón and Perera-Valderrama 2014).

Density and biomass of small fish were higher in deeper Cuban coral reefs but not consistently different in Jardines de la Reina and Sabana-Camagüey (Table 15.2). Herbivorous fish have shown higher biomass not only in reef crests in Canarreos and Sabana-Camagüey (Sierra et al. 2001) but also nationwide (see Chevalier Monteagudo et al., Chap. 12). Density and biomass of commercial fish and herbivorous fish tend to be higher in reef crests (10 comparisons out of 12). However, commercial fish showed higher biomass in fore reefs decades ago in Jardines de la Reina, Canarreos, and Sabana-Camagüey (Sierra et al. 2001). In the case of commercial fish, differences between our results and the previous ones might be related to the fact that we found higher fishing pressure and weaker enforcement in spur and groove habitats (fishing pressure, X2 = 83.50, p < 0.001, enforcement, X2 = 114.14, p < 0.001) when compared to reef crests nationwide. The opposite pattern found in small fish (more in deeper reefs) suggested control by predation, since many of the commercial fish are predators of the small fish (Sierra et al. 2001; Hernández-Hernández et al. 2008). Acute wave exposure did not differ between deeper habitats and reef crests (X2 = 0.08, p = 0.779) but chronic wave exposure did (X2 = 8.05, p = 0.004) (Fig. 15.7). However, small data range distribution suggested no ecological relevance between habitats or zones in the Cuban coral reef system scale. Several studies have addressed the potential impact of hurricanes on coral reefs all around Cuba: Sabana-Camagüey (Alcolado et al. 2007), Guanahacabibes

292 Fig. 15.5 Temporal trends of coral reef anthropogenic factors in Cuba. Horizontal lines indicate medians. Boxes represent interquartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

Fig. 15.6 Temporal trends on enforcement of regulations and fishing pressure in Jardines de la Reina (JR) and Sabana-Camagüey (SC). Horizontal lines indicate medians. Boxes represent interquartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

F. Pina-Amargós et al.

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Table 15.2 Biomass and density of fish among several coral reef habitats in Cuba Variable Commercial fish biomass

JR Habitat comparison Cre > Dof

Commercial fish density

Cre = Dof

Herbivorous fish biomass

Cre > Dof

Herbivorous fish density

Cre > Dof

Small fish biomass

Cre = Dof

Small fish density

Cre = Dof

Significance X2 = 4.53 p = 0.033 X2 = 0.35 p = 0.580 X2 = 78.74 p < 0.001 X2 = 88.75 p < 0.001 X2 = 0.60 p = 0.806 X2 = 3.26 p = 0.071

SC Habitat comparison Cre = Dof ≥ S&G Cre = Dof > S&G Cre = Dof > S&G Cre = Dof = S&G Cre = Dof = S&G S&G > Dof = Cre

Significance X2 = 12.30 p = 0.002 X2 = 53.69 p < 0.001 X2 = 22.06 p < 0.001 X2 = 2.53 p = 0.282 X2 = 3.65 p = 0.162 X2 = 24.42 p < 0.001

Cuba Habitat comparison Cre = Dof ≥ S&G Cre > Dof > S&G Cre > Dof Cre > Dof Dof = S&G > Cre Dof = S&G > Cre

Significance X2 = 12.40 p = 0.002 X2 = 53.69 p < 0.001 X2 = 70.86 p < 0.001 X2 = 72.64 p < 0.001 X2 = 23.50 p < 0.001 X2 = 21.60 p < 0.001

Legend: JR, Jardines de la Reina; SC, Sabana-Camagüey; Cre, reef crest; Dof, drop off; S&G, spur and groove

(González-Ferrer et al. 2007b; Perera-Valderrama et al. 2013), Northwestern (see Chap. 18, González-Díaz et al.), Los Colorados (Caballero-Aragón et al. 2019), Southern archipelagos (Caballero-Aragón and Perera-Valderrama 2014), Jardines de la Reina (Pina-Amargós et al. 2008b), Havana (Durán et al. 2018), and condition of A. palmata in Cuba (Caballero-Aragón et al. 2020). In Guanahacabibes, the condition of coral reefs is related to wave exposure (chronic and acute) (Perera-Valderrama et al. 2013). However, quantitative data on wave exposure are missing in all those studies. Some studies used quantitative data to discuss their findings but were limited to sites on a scale of a few square kilometers (Alcolado et al. 2013; Caballero-Aragón et al. 2016). Exploratory analysis revealed a similar trend of ecological indicators among habitats. Therefore, we grouped data from all the habitats for most analysis and clarified when a specific habitat was selected. Chronic wave exposure correlated positively with small fish (biomass, rs = 0.477, p < 0.001, n = 412), density of juvenile corals (rs = 0.432, p = 0.046, n = 66), and density and species richness of sponges (rs = 0.439, p = 0.002, n = 9; rs = 0.568, p = 0.001, n = 54, respectively). The waves might be suspending preys and sediments (nutrients), benefiting directly and indirectly small fish through enhanced feeding opportunities (Hernández-Hernández et al. 2008). Studies have shown that several species of crustose coralline algae usually dominate habitats exposed to waves, and some could promote coral recruitment (Gouezo et al. 2020). High density of juvenile corals and sponges has been observed in Havana and its surroundings (Durán et al. 2018; see Chap. 17, González-Díaz et al.). Those authors emphasized the role of pollution on that pattern, but the role of waves on enhancing autotrophy and heterotrophy processes on juvenile corals should not be excluded. Waves might facilitate density and species richness of sponges by providing nutrients for

bacteria since sponges feed predominantly by filtration, targeting mostly ultra-plankton (Rützler 2004). Coral cover correlated negatively (but low absolute value) with chronic wave exposure (rs = –0.219, p < 0.001, n = 597), suggesting potential deleterious effect. Acute wave exposure (hurricane frequency) correlated negatively with species richness of gorgonians (rs = – 0.705, p < 0.001, n = 43) and sponges (rs = –0.566, p = 0.001, n = 54). Acute waves might heavily impact gorgonian and sponge communities by detaching colonies from the substrate, as found in other studies (Yoshioka and Yoshioka 1991). Our results suggest that rugosity is correlated with the diameter of M. cavernosa (rs = 0.367, p < 0.001, n = 260) and relative abundance of A. palmata (rs = 0.375, p = 0.003, n = 131) likely due to their condition as coral reef engineer species (defined by Jones et al. 1997) as observed previously (Caballero-Aragón et al. 2019, 2020). Correlations between rugosity and fish are significant but have a low absolute value. Small fish might be benefiting from the shelter and food availability with increased rugosity (density, rs = 0.269, p < 0.001, n = 413), as broadly found elsewhere (Bohnsack and Bannerot 1986; Aguilar-Betancourt and GonzálezSansón 2002; Friedlander et al. 2003a). Biomass of commercial and herbivorous fish might be enhanced by increased rugosity (rs = 0.235, p < 0.001, n = 421, rs = 0.305, p < 0.001, n = 412, respectively), as found previously (Cobián-Rojas et al. 2011). Coral cover correlated positively with rugosity (rs = 0.250, p < 0.001, n = 597) as has been found in other Cuban studies (Caballero-Aragón et al. 2019, 2020). Other studies have not found statistical differences of rugosity among zones in Havana using quantitative data (Durán et al. 2018) and among zones in Jardines de la Reina using qualitative estimates (Pina-Amargós et al. 2008a). However, a countrywide comparison using quantitative data showed difference among zones and among sites

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Fig. 15.7 Acute and chronic wave exposures on Cuban coral reefs. Deep, drop off, and spur and groove habitats; shallow, reef crest; Hurr, hurricane frequency; Wh, wave height; Wt, wave period. Horizontal lines indicate medians. Boxes represent interquartile range (IQR). Whiskers tips correspond to quartile values ± 1.5 × IQR. Dots identify outliers

inside the zones (Caballero-Aragón et al. 2019). In reef crests, Canarreos, Cazones, and Southcentral had the highest complexity, while the lowest and most variable complexity occurred in Sabana-Camagüey. In the fore reefs, the highest complexity occurred in Girón and Southcentral with Canarreos, Cazones, and Guanahacabibes having the highest variability, followed by Los Colorados (Northwestern zone), and the lowest complexity appeared in Jardines de la Reina, Southeastern, San Felipe, Havana (Northwestern zone), Sabana-Camagüey, and Northeastern (Caballero-Aragón et al. 2019). On Guanahacabibes, a study using quantitative and qualitative variables found differences in rugosity among sites for both methods (Cobián-Rojas et al. 2011). Those studies suggest that rugosity is as highly variable inside zones as it is among them. Thus, we considered rugosity is not explaining the patterns observed at the scale of our study. Other environmental factors such as water clarity and salinity were not analyzed in our study. However, results related to Cuba from a Caribbean-wide study (Chollett et al. 2012) showed water clarity and salinity have a narrow range to influence differences among the zones of the Cuban coral reefs.

15.5.2.2 Anthropogenic Factors Related to the Status of Cuban Coral Reefs Of the 238 correlations between anthropogenic factors and Cuban coral reefs biota, 27 (11%) were significant and of high absolute value. Most of the significant and of high absolute value correlations (20, 74%) were related to commercial and herbivorous fish, and some others to fishing pressure (7, 26%). Fishing pressure is widely recognized as the most important anthropogenic factor impacting fish in the world ocean (Barange et al. 2018; FAO 2018; Bennett et al. 2018). Our results showed consistently lower density and biomass of fish, particularly commercial species, in coral reefs with a

higher level of fishing pressure except in Sabana-Camagüey. Fishing pressure correlated negatively with biomass of commercial fish in Cuba (rank scale, rs = –0.550, p < 0.001, n = 421; quantitative, rs = –0.432, p < 0.001, n = 413), biomass of commercial fish (low absolute value, rs = –0.303, p < 0.001, n = 160), and density and biomass of herbivorous fish in Jardines de la Reina (low absolute value on both, density, rs = –0.340, p < 0.001, n = 160; biomass, rs = – 0.289, p < 0.001, n = 160) but positively with density and biomass of herbivorous fish (density, rs = 0.405, p = 0.002, n = 78; low absolute value, biomass; rs = 0.341, p < 0.001, n = 78) and with density and biomass of commercial fish in Sabana-Camagüey (low absolute value on both, density, rs = 0.259, p = 0.004, n = 78; biomass, rs = 0.125, p = 0.013, n = 78). Several studies reported high fishing pressure and poor status of fisheries resources in Cuba: in Sabana-Camagüey (Claro et al. 2004, 2009); in Golfo de Batabanó (Claro et al. 2001); in Jardines de la Reina (Claro et al. 2009); Golfo de Ana María and Guacanayabo (Alzugaray and Puga 2010); and nationwide (Baisre 2018). Several studies in Cuba have related density and biomass of fish to qualitative fishing effort: in Sabana-Camagüey (Claro et al. 2007; GonzálezFerrer et al. 2007a) and in Havana (Aguilar-Betancourt et al. 2007; Durán et al. 2018). Some studies on Northwestern Cuba (from Havana to the westernmost part of Cuba) used a rank scale and fisheries data for exploring the relationship between fish density and biomass and fishing (GonzálezSansón et al. 2009a, b). They found a strong negative relation between fishing and fish communities’ status. In Jardines de la Reina, density of commercial fish was examined taking into account quantitative assessments of fishing pressure, and it was found that fishing reduction due to the MPA was the main cause of the high level of fish density (Pina-Amargós et al. 2009). In Sabana-Camagüey, the highest values of density and biomass of fish were reported in the 1980s,

15

Status of Cuban Coral Reefs

despite the high levels of fishing. Since that fishing pressure has not been significantly reduced, fish assemblages have been diminished through time, currently showing the lowest levels (Fig. 15.6). No significant correlations were detected between fishing pressure and sessile benthic variables, suggesting no direct influence. Enforcement of regulations (fisheries, biological diversity, and MPA) is critical for conservation (Jackson et al. 2014; Pina-Amargós et al. 2014a). It correlated positively with biomass of commercial (rs = 0.380, p < 0.001, n = 421) and herbivorous fish (low absolute value, rs = 0.259, p < 0.001, n = 412) in Cuba. It also correlated positively with biomass of commercial (low absolute value, rs = 0.173, p < 0.001, n = 160) and herbivorous fish (low absolute value, rs = 0.214, p < 0.001, n = 160) and density of D. antillarum (rs = 0.312, p < 0.001, n = 153) in Jardines de la Reina. Implementation of regulations correlated negatively with density of commercial fish (rs = –0.386, p < 0.001, n = 78) and biomass of herbivorous fish (rs = – 0.412, p < 0.001, n = 78) on Sabana-Camagüey (Fig. 15.6) and negatively with fishing pressure in Cuba (rank scale, rs = –0.704, p < 0.001, n = 752; quantitative, rs = –0.607, p < 0.001, n = 741), Jardines de la Reina (quantitative, rs = – 0.838, p < 0.001, n = 195), and Sabana-Camagüey (low absolute value, quantitative, rs = –0.364, p < 0.001, n = 243). It correlated positively with protection in Cuba (low absolute value, rs = 0.400, p < 0.001, n = 752). Our results suggest a direct effect of enforcement on reducing fishing pressure in Cuba, Jardines de la Reina, and Sabana-Camagüey, and that the enforcement is stronger in MPAs. Few sources reported enforcement of regulations in Cuba’s marine environment, but this seems to be increasing in the last few years (CBD 2019). In our study, most of the correlations of enforcement with density and biomass of fish were positive. However, absolute value was low, which supports our assumption that enforcement directly reduces fishing pressure and indirectly increases fish density and biomass. Previous studies linked the high density and biomass of fish to strong enforcement (Pina-Amargós et al. 2014a) but without estimating it. On the other hand, enforcement correlated negatively with commercial and herbivorous fish in Sabana-Camagüey. In this zone, the highest values of density and biomass of fish were documented in the 1980s with low levels of enforcement (X2 = 41.75, p < 0.001). Law on Fisheries and its enforcing body, the fisheries inspectors, were not operating until 1994 (permits, quotas, gear restrictions, protected species, etc.). From that time on, enforcement increased to a weekly frequency (X2 = 41.75, p < 0.001), but fishing pressure did not decrease significantly, and consequently it has been impacting fish assemblages in Sabana-Camagüey until now. This fact contrasts with findings in the Jardines de

295

la Reina coral reefs, where enforcement has increased up to a daily frequency (X2 = 72.99, p < 0.001), effectively reducing fishing effort (X2 = 107.25, p < 0.001) with a positive impact on fish assemblages (Fig. 15.6). The enforcement level between Jardines de la Reina and Sabana-Camagüey is different (W = 40,202, p < 0.001). These findings suggested that enforcement levels must be very high to reduce fishing pressure and positively impact target species. No significant and high absolute value correlation was detected between enforcement and benthic variables, suggesting no direct influence. The MPAs are a worldwide conservation tool and have been strongly promoted in Cuba in the last 20 years (see Chap. 20, Perera-Valderrama et al.). Protection (through MPAs) correlated positively with biomass of commercial (rs = 0.415, p < 0.001, n = 421) and herbivorous fish (low absolute value, rs = 0.382, p < 0.001, n = 412) and negatively with species richness of gorgonians (rs = 0.541, p = 0.029, n = 43) in Cuba. Analysis of species richness of gorgonians in MPAs in Guanahacabibes, Girón, and Cazones showed an opposite pattern (see Chap. 9, Rey-Villiers et al.) likely due to the smaller sample size of our data. Our results suggest a successful role of the Cuban MPAs in reducing fishing pressure and increasing biomass of commercial and herbivorous fish when compared to coral reefs outside MPAs. However, these findings might not be related to the performance of the MPAs but to the selection of coral reefs with a higher biomass of fish to establish MPAs (see Chap. 20, Perera-Valderrama et al.). Only in Jardines de la Reina studies have proved the effectiveness of MPAs on fish assemblages’ conservation in Cuba. Most of the other studies have not introduced a BACI design (Before-After, ControlImpact) for assessing their effectiveness, neither at the time of implementing monitoring and research programs (PinaAmargós et al. 2014a, b). Studies from Jardines de la Reina have ruled out other alternative explanations such as habitat quality, differential recruitment, and historical fishing, leaving protection as the most plausible cause of higher abundance of commercial fish inside this MPA. No significant and high absolute value correlations were detected between protection and other benthic variables, suggesting no direct influence on them. Coral reefs near populated cities are in poor conditions due to pollution and overfishing (González-Díaz et al. 2003, 2018; González-Sansón et al. 2009a; Castellanos-Iglesias et al. 2018; Durán et al. 2018; see Chap. 17, González-Díaz et al.). Population, pollution, and fishing pressure are positively and highly correlated, suggesting the synergic action of multiple stressors (Table 15.3). Population correlated negatively with density (low absolute value, rank scale, rs = – 0.331, p < 0.001, n = 422; quantitative, rs = –0.458, p < 0.001, n = 422) and biomass (rank scale, rs = –0.465, p < 0.001, n = 421; low absolute value, quantitative, rs = –

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0.344, p < 0.001, n = 421) of commercial fish and positively with corals recruits’ density (quantitative, rs = 0.415, p < 0.001, n = 492) in Cuba. Our results support findings of other studies showing lower density and biomass of commercial fish near large human settlements, suggesting heavy fishing for self-consumption and to supply private restaurant (Durán et al. 2018). No significant and high absolute value correlation was detected between population and most benthic variables, suggesting no direct influence. The potential positive influence of population (through pollution) on the density of coral recruits was discussed above (chronic wave exposure paragraph). The negative influence of pollution on the biomass of commercial (rs = –0.523, p < 0.001, n = 421) and herbivorous fish (rs = –0.466, p < 0.001, n = 412) might result from the positive correlation between pollution and fishing pressure (Table 15.3), but sub-lethal effects of pollution might be affecting those fish as well (Aguilar-Betancourt et al. 2007, 2008). No significant and high absolute value correlation was detected between pollution and benthic variables, suggesting no direct influence. This is presumably due to the qualitative nature of our pollution estimates, since other studies have associated coral reef deterioration with pollution from populated urban areas such as Havana (González-Sansón and Aguilar-Betancourt 2010; Castellanos-Iglesias et al. 2018; Durán et al. 2018; Rey-Villiers et al. 2020a, b; see Chap. 17, González-Díaz et al.) and Cienfuegos (Rancho Luna and Guajimico coral reefs) (Caballero-Aragón and Perera-Valderrama 2014). Even smaller cities such as Cárdenas, Varadero, Isabela de Sagua, and Nuevitas in Sabana-Camagüey impact adjacent marine waters through pollution (Montalvo-Estévez et al. 2007). In Cuba, marine pollution far from large cities is strongly related to the sugarcane industry and river discharge (Alcolado et al. 2003, 2007). Studies from SabanaCamagüey provide data on both sources (Penié and García 1998; Perigó et al. 2004; Montalvo-Estévez et al. 2007). In addition, there is evidence of pollution reaching coral reefs in Villa Clara and Camagüey, in Sabana-Camagüey zone (Montalvo-Estévez et al. 2007). However, studies suggested a 3–4-fold decrease of pollution (but highly variable) from the 1990s to the 2000s in Sabana-Camagüey (MontalvoEstévez et al. 2007) due primarily to the closure of half of the sugar cane factories (SC 2008). Sugarcane factories were reduced 45% nationwide, and the most recent report available (CBD 2019) suggests a reduction in pollution reaching the Cuban marine environment in the timeframe covered by our data. However, cumulative effects of point and longtime pollution and their synergistic effects with other stressors might be occurring (Aguilar-Betancourt et al. 2007, 2008).

F. Pina-Amargós et al.

Isolation is one of the key elements usually associated with coral reef conservation worldwide (Brewer et al. 2013; Cinner et al. 2016; Juhel et al. 2017). Distance from the coast correlated positively with density of commercial fish (rs = 0.440, p < 0.001, n = 422) and negatively with biomass of small fish (rs = –0.471, p < 0.001, n = 387) and density of corals (total) (rs = –0.427, p < 0.001, n = 581). Distance from the coast is considered a proxy for several anthropogenic factors such as population, land-based pollution, and fishing pressure (Brewer et al. 2013; Cinner et al. 2016; Juhel et al. 2017). Thus, our results are not surprising, and most of those previously discussed apply here. No significant and high absolute value correlation was detected between distance from the coast and other benthic variables, suggesting no direct influence. The social and economic benefits of tourism are usually coming together with environmental degradation of coral reefs and adjacent interconnected ecosystems, as shown by several studies (Barker and Roberts 2004 (impact of diving); Brownscombe et al. 2015 (impact of recreational fishing)). In our study, tourism did not correlate significantly with any Cuban coral reef biota variable. However, since tourism correlated positively with population and pollution, its impacts might be masked (Table 15.3). For example, Havana is the most populated and polluted city of the country and also the one with the highest demand of seafood by tourists dining in private restaurants (Durán et al. 2018). Generally speaking, Cuban coral reefs receive fewer visitors than similar destinations such as the Bahamas (Carwardine and Watterson 2002), Moorea Island (Clua et al. 2011), Belize (Graham 2004), and Playa del Carmen in México (Whitcraft 2010). There are relatively few diving shops (54 in the entire country, Tamara Figueredo Martín and Fabián Pina Amargós, unpublished data), and 35% of coral reef sites of this study receive a few hundred of mainly experienced visitors per year. Other nautical activities are also based on low infrastructure development and numbers of visitors: 17 marinas (1800 mooring sites, 80% concentrated in Havana and Varadero), 20 charter boats for coral reef and offshore fishing (800 clients per year) (Havana, Varadero, Villa Clara, northern Ciego de Avila, and Holguin), nine live aboard boats (diving and snorkeling in Cazones and Jardines de la Reina and fly fishing in Isla de la Juventud, Cazones, Jardines de la Reina, northern Camaguey), and 72 skiffs for fly fishing (3000 anglers per year) (Tamara Figueredo Martín and Fabián Pina Amargós, unpublished data). Several studies have attempted to assess tourism impact on coral reefs, but most of them do not support it with visitor impact data: in Sabana-Camagüey (García et al. 2007; SC 2008; Lorenzo et al. 2013), in Guajimico (Cienfuegos) (de la Guardia 2006), and in Southern archipelagos (CaballeroAragón and Perera-Valderrama 2014). Hernández-Fernández

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Table 15.3 Correlations between pairs of anthropogenic factors estimated using a rank scale (Table S15.1). All values shown are significant ( p < 0.001). Sample size for every correlation is 752 Tourism Fishing Pollution Enforcement Protection

Population 0.415 0.528 0.684 –0.295 –0.305

Tourism

Fishing

Pollution

Enforcement

NS 0.616 NS –0.283

0.276 –0.704 –0.314

–0.232 –0.411

0.400

NS, not significant

et al. (2008, 2016) assessed scuba diving impact in Cayo Coco (Sabana-Camagüey) and Jardines de la Reina, respectively, and found no evidence of coral reef deterioration related to scuba diving. Research and monitoring of tourism impact on coral reefs biota, particularly scuba diving, snorkeling, and recreational fishing, should be continued and expanded. Despite the potential impact of tourism on coral reefs, sound management is possible, as shown in Jardines de la Reina (Figueredo-Martín et al. 2010a, b) and Guanahacabibes (Cobián-Rojas and Chevalier-Monteagudo 2009), where tourism activities are profitable, socially responsible, and environmentally sensitive. Integrated impact correlated negatively with density (rs = –0.477, p < 0.001, n = 422) and biomass (rs = – 0.582, p < 0.001, n = 422) of commercial fish and biomass of herbivorous fish (rs = –0.450, p < 0.001, n = 412) nationwide. This result suggests the highest relevance of fishing pressure among the anthropogenic factors grouped by integrated impact. The NMDS for anthropogenic factors (discontinuous variables) split Cuban coral reef sites into two groups: one group associated with the highest pollution, fishing, tourism, population, and the lowest level of enforcement and protection and another group characterized by the lowest values of pollution, fishing, tourism, population, and the highest level of enforcement and protection (Fig. 15.8). PERMANOVA results confirmed the patterns observed in NMDS plots. Differences among levels of each factor were significant for enforcement (F3,175 = 11.64, p = 0.001), pollution (F2,176 = 7.53, p = 0.001), fishing (F5,173 = 12.76, p = 0.001), population (F3,175 = 5.59, p = 0.001), and protection (F1,177 = 22.77, p = 0.001). Site groups by level of impact using NMDS have been obtained in other studies in Cuba (see Chap. 17, González-Díaz et al.). According to BIOENV analysis for environmental factors (sea surface temperature, habitats around coral reef, chronic wave exposure, and acute wave exposure) and anthropogenic factors (continuous) (population, distance to coast, fishing, and tourism), the ones that better explain multidimensional variation are sea surface temperature, distance to coast, and fishing (rs = 0.379). The last two factors showed similar correlation than the former three (rs = 0.364), suggesting predominance of anthropogenic factors among those modulating Cuban

coral reefs. However, it is difficult to separate distance to coast and fishing given their high correlation (Table 15.3).

15.5.3 Potential Ecological Relationships on Cuban Coral Reefs Of the 180 correlations among ecological factors on Cuban coral reefs biota, 73 (41%) were significant and high absolute value. We discussed 24 (35%), the ones most deeply studied: D. antillarum with corals and macroalgae, lionfish and its potential predators and preys, among fish groups by potential predator–prey relationships.

15.5.3.1 Potential Ecological Relationships of D. antillarum Density of D. antillarum correlated negatively with macroalgal cover in Cuba (low absolute value, rs = –0.244, p < 0.001, n = 400), in Jardines de la Reina (rs = –0.321, p < 0.001, n = 153), and in Sabana-Camagüey (rs = –0.314, p < 0.001, n = 153), and with density of coral recruits in Jardines de la Reina (low absolute value, rs = –0.259, p < 0.001, n = 153). Our findings suggest that in Cuban coral reefs, particularly on reef crests, D. antillarum controls macroalgal cover. A similar result is found in Durán et al. (see Chap. 11). This is an encouraging finding regarding the role of D. antillarum in the recovery of the Cuban coral reefs dominated by corals taking into account the still low density of this species (see Chap. 11, Durán et al.). In the case of coral recruits, D. antillarum might consume them in Jardines de la Reina coral reefs, as reported by other studies in the Caribbean (Solandt and Campbell 2001; Herrera-López et al. 2004). 15.5.3.2 Potential Ecological Relationships of Corals Coral cover correlated negatively with macroalgal cover in Sabana-Camagüey (rs = –0.539, p < 0.001, n = 160) and Cuban coral reefs (rs = –0.224, p < 0.001, n = 535). This suggested an already established phase shift from coraldominated to macroalgae-dominated coral reefs in Cuba and Sabana-Camagüey. Other studies have documented this phase shift in Cuba (i.e., Martín-Blanco et al. 2011; Durán

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F. Pina-Amargós et al.

Fig. 15.8 Non-metric multidimensional scaling (NMDS) of the sites based on six selected variables. Points of the same ordination diagram (stress = 0.15) have been identified after different factors for comparative purposes. 1 means the lowest value of the factors and 3 (pollution), 4 (enforcement, tourism), and 6 (fishing) the highest values of the factors

et al. 2018; Caballero-Aragón et al. 2019) and the Caribbean (i.e., Jackson et al. 2014). However, in Jardines de la Reina, that pattern is not consistently established (significant but low absolute value), suggesting three alternatives: phase shift is underway, reversing, or never happened. The most likely scenario is the reversing one, according to other studies (Pina-Amargós et al. 2021). Since coral-dominated reefs can still have a negative correlation between coral and macroalgae, an alternative explanation is that this relation might also imply competition between corals and macroalgae (see Chap. 11, Durán et al.).

15.5.3.3 Potential Ecological Relationships of Fish Relationships among lionfish, potential prey, and predators are still controversial topics related to the invasion of that species in the Caribbean (i.e., Mumby et al. 2011; Valdivia et al. 2014). Lionfish density and biomass correlated negatively with density of commercial fish (low absolute value, density, rs = –0.251, p < 0.001, n = 403; biomass, rs = – 0.350, p < 0.001, n = 377) in Cuba and in Sabana-Camagüey (density, rs = –0.479, p < 0.001, n = 78). This might be an indication of lionfish control by predators, as found by other studies (i.e., Mumby et al. 2011). However, other studies found no control of the lionfish by predation in the Caribbean (Hackerott et al. 2013; Valdivia et al. 2014) and Cuba

(Cobián-Rojas et al. 2018). The relation of lionfish with small fish as potential preys is positive in Sabana-Camagüey (density, rs = 0.335, p = 0.002, n = 78) and in Cuba (low absolute value, density, rs = 0.201, p = 0.002, n = 368), suggesting no predation control by lionfish also found by other studies in the Caribbean (Elise et al. 2014; see Chap. 12, Chevalier Monteagudo et al.). An alternative explanation regarding the relationship between prey abundance and lionfish abundance suggests the latter is related to prey availability (Chevalier-Monteagudo et al. 2013). According to Hammerschlag-Peyer and Layman (2010), the lionfish selects its habitat following prey density, so it shows greater fidelity to sites that offer a variety of prey at small scales. However, another study of lionfish–prey analysis to species level suggests an impact by lionfish, confirmed by stomach content analysis for certain species (Cobián-Rojas et al. 2018). Other studies reported control of prey by lionfish predation in the Caribbean (Green et al. 2012; Albins 2015) and Cuba (Cobián-Rojas et al. 2018). These apparent contradictions likely reflect that local conditions, including environmental factors, might modulate predation of and from lionfish (see Chap. 13, Pina-Amargós et al.). Human control of lionfish in Cuban coral reefs is an alternative explanation to the above finding. Still, correlations are of low absolute

15

Status of Cuban Coral Reefs

value nationwide to be considered relevant, but they might be locally important. Commercial fish (biomass) correlated positively with herbivorous fish (density and biomass) in Jardines de la Reina (low absolute value on both, density, rs = 0.262, p < 0.001, n = 160; biomass, rs = 0.261, p < 0.001, n = 160) and Sabana-Camagüey (density, rs = 0.360, p < 0.001, n = 78; biomass, rs = 0.538, p < 0.001, n = 78) (including density of commercial fish with density of herbivorous fish) (rs = 0.373, p < 0.001, n = 78). Biomass of commercial fish correlated negatively with biomass of small fish in Jardines de la Reina (rs = –0.313, p < 0.001, n = 160). Our results suggest that commercial fish (predators) are not controlling herbivorous fish (prey) in coral reefs, but they do in the case of small fish. It is well established that most of the commercial fish of this chapter (i.e., snappers, groupers, and jacks) prey on herbivorous fish (Sierra et al. 2001; Hernández-Hernández et al. 2008). Since commercial and herbivorous fish increase in Jardines de la Reina while fishing pressure decreases and both groups decrease in Sabana-Camagüey while fishing pressure remains similar through time (Fig. 15.2, 15.3, and 15.6), the relationship between the two groups does not seem to be modulated by fishing. Similar biomass of herbivorous fish has been reported related to a wide range of their predators’ biomass in Cuba such as in Jardines de la Reina, Canarreos, and Sabana-Camagüey (Sierra et al. 2001) and in Martinica, Guadalupe Florida (Sierra et al. 2001), and the Bahamas (Mumby et al. 2006), supporting our findings. On the other hand, predator–small fish relationships through temporal and habitat trends and correlations suggest control by predation (Sierra et al. 2001; HernándezHernández et al. 2008). The abundance of damselfish (one of the two families included as “small fish”) has been increasing on Caribbean reefs (Ceccarelli et al. 2001), possibly due to the low abundance of predators, including groupers and snappers (Robertson 1996; Mumby et al. 2012; Vermeij et al. 2015; Rivera-Sosa et al. 2018). Since damselfish may play a role in maintaining feedback toward macroalgae-stressed states through algal gardening in wider Caribbean coral reefs (Randazzo Eisemann et al. 2019), specific research to assess that process should be designed and carried out in Cuban coral reefs. The density of small fish seems to be benefiting from the density of D. antillarum in Sabana-Camagüey (rs = 0.450, p = 0.004, n = 136). D. antillarum generalist feeding (Solandt and Campbell 2001; Herrera-López et al. 2004) might be creating turf-dominated spaces favoring turf gardening by damselfish (Randazzo Eisemann et al. 2019) and hunting grounds for wrasses. Herbivorous fish seem to be impacting macroalgae negatively in Jardines de la Reina (density, rs = –0.655,

299

p < 0.001, n = 160; biomass, rs = –0.502, p < 0.001, n = 160) and in Cuba (density, rs = –0.562, p < 0.001, n = 258; biomass, rs = –0.471, p < 0.001, n = 257). Herbivorous fish also seem to be impacting negatively corals (density of recruits (rs = –0.511, p < 0.001, n = 153), density of adults (rs = –0.507, p < 0.001, n = 195), and total density of corals (rs = –0.556, p < 0.001, n = 195)) and positively D. antillarum (rs = 0.464, p < 0.001, n = 153) in Jardines de la Reina. These results suggest herbivorous fish control macroalgae in Cuba and Jardines de la Reina. A more detailed analysis found the same pattern nationwide (see Chap. 11, Durán et al.), but others have not observed this pattern in Northwestern (see Chap. 17, González-Díaz et al.). The trends of parrotfish biomass recovery and decreasing macroalgal cover nationwide suggested previously (Jackson et al. 2014) match the results of this study. One study in Havana found high macroalgal cover and low biomass of herbivorous, stressing the importance of herbivory in keeping the Cuban coral reefs dominated by corals (Durán et al. 2018). The negative correlation between herbivorous fish and corals densities (recruits, adults, and total) in Jardines de la Reina has at least two potential explanations. First, herbivorous fish might be feeding on corals in Jardines de la Reina. It is known that several species of parrotfish frequently bite living corals, potentially influencing coral abundance (Rotjan et al. 2006; Burkepile 2012), but this is not supported by field observations in Jardines de la Reina (Tamara Figueredo Martín and Fabián Pina Amargós, unpublished data). Second, spurious relation: Jardines de la Reina protection from fishing has allowed large herbivorous assemblages with relatively low coral densities. Both hypotheses need to be tested in the field. Durán et al. (see Chap. 11) did not find a significant correlation between herbivorous fish and coral cover, but González-Díaz et al. (see Chap. 17) found in low impact sites more corals and less herbivorous fish. González-Díaz et al.’s (see Chap. 17) explanation is that sites with low impact are not rich in nutrients which favors the development of corals, creates unfavorable conditions for algae, and limits bottom-up control (algae-herbivores fish). A positive correlation between herbivorous fish and the density of D. antillarum in Jardines de la Reina has at least two potential explanations. First, it might be a facilitation process between them. Lessios (2016) hypothesized that macroalgae limit D. antillarum settlement, while the abundance of herbivorous fish controlling macroalgae facilitates it. Second, spurious relation: both are more abundant in Jardines de la Reina reef crests for reasons independent of their interaction.

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15.6

F. Pina-Amargós et al.

Integration of Results

Our analysis of several environmental, climate, anthropogenic factors and ecological relationships on Cuban coral reef benthos and fish in the last 20 years allows us to conclude that observed impacts on studied communities are driven by anthropogenic factors and not by environmental ones. The same conclusion was reached by González-Díaz et al. (see Chap. 17). Environmental factors do not seem to drive temporal and spatial trends or most of the ecological relationships observed in Cuban coral reef system. Despite a significant increase in sea surface temperature nationwide, no clear temporal or spatial trends on bleaching and no relationship between bleaching events and mortality and coral diseases were detected. Acute wave exposure (frequency of hurricanes) has decreased significantly in Cuban coral reefs, but the data range is too narrow to give clear explanations of the observed patterns. Chronic wave exposure, habitats around coral reefs, and rugosity remained stable nationwide along the timeframe covered by our data. Previous studies found rugosity is as highly variable inside zones as it is among them. Thus, rugosity does not explain the patterns seen at the scale of our research. Different coral reef habitats support different benthos and fish communities. Still, temporal trends of environmental, climate, anthropogenic factors and ecological relationships are primarily similar among the most representative ones (reef crest, drop off, and spur and groove). Environmental and climate factor correlations with biota were primarily with sponges, gorgonians, and small fish, components with less contribution to coral reef ecology in terms of substrate cover and rugosity (sponges, gorgonians) and biomass (small fish). Most of those relations were found with chronic and acute wave exposures, temporal trends of which were not significant statistically or ecologically (narrow range), respectively. Other studies show that water clarity and salinity data have narrow ranges to be relevant in detected patterns. Anthropogenic factors seem to drive temporal and spatial trends and most of the ecological relationships observed in the Cuban coral reef system. Most of the correlations among anthropogenic factors and biota involve commercial and herbivorous fish, and more than half are related to fishing pressure and enforcement. During the timeframe of this study, fishing pressure and enforcement have changed in Jardines de la Reina, Sabana-Camagüey, and nationwide. Fish targeted directly by fishing (commercial and herbivorous) respond to those changes: their abundance decreases when fishing increases, and are less abundant in zones and habitats where fishing pressure is higher, and vice versa. Commercial and herbivorous fish respond the same way to integrated impact than to fishing pressure, adding more

evidence to fishing pressure as the primary factor driving coral reef status and trends in Cuba. The other anthropogenic factors (population, tourism, pollution, protection) did not change along our data timeframe, but since many of them are correlated with fishing pressure and enforcement, their effects seem to be masked. We considered these correlations as evidence of multiple anthropogenic factors impacting Cuban coral reefs rather than a lack of independence of estimates of those variables. Our assessment methods prevented such a bias as multivariate analyses show. Multivariate analysis of anthropogenic factors (NMDS) splits the Cuban coral reefs into two groups: one group displaying the highest values of most factors (and the lowest enforcement and protection) and the other displaying the lowest values (and the highest enforcement and protection) (Fig. 15.8). Multivariate analysis of the correlation of anthropogenic, environmental, and climate factors (BIOENV) revealed the high relevance of distance to coast and fishing pressure, suggesting predominance of anthropogenic factors among those modulating Cuban coral reefs without excluding sea surface temperature as an important one. Recent studies demonstrated that while environmental factors such as sea surface temperature and wave energy have strong power in predicting benthic assemblages at remote reefs, this predictive power is lost or the relationship is fundamentally altered at reefs closer to human populations (Williams et al. 2015; Ford et al. 2020). They conclude that factors associated with local anthropogenic impacts may have overtaken biophysical drivers in structuring altered reefs. A comprehensive analysis of the biota status, trends, and ecological relationships in Jardines de la Reina, SabanaCamagüey, and nationwide allows us to reach some generalizations. Most Cuban coral reefs are under high fishing pressure and few are highly protected. Macroalgae dominates most of the Cuban coral reefs. However, herbivory from fish and D. antillarum seems to be happening, making restoration to coral-dominated reefs possible taking into account the low incidence of coral diseases nationwide. The invasion of lionfish is presumably not impacting small fish (potential prey) likely due to a combined control of environmental and human factors and native predators. The Jardines de la Reina coral reefs receive light fishing pressure due to a well-enforced MPA and fisheries regulations, likely responsible for the high abundance of commercial and herbivorous fish. Small fish are likely controlled by commercial fish in Jardines de la Reina coral reefs. The Jardines de la Reina coral reefs are dominated by macroalgae, but coral cover is often relatively high, presumably due to the stronger effect of herbivorous fish and D. antillarum on macroalgae. On the other hand, the Sabana-Camagüey coral reefs receive high fishing pressure. They are dominated by macroalgae as well, but herbivory seems to be weaker than nationwide and in

15

Status of Cuban Coral Reefs

Jardines de la Reina, compromising phase shift reversal to a coral-dominated reef. These trends and patterns are explained by a higher and stable fishing pressure in Sabana-Camagüey and a lower and decreasing fishing pressure in Jardines de la Reina. In the case of other Cuban zones, there might be peculiarities such as in Northwestern Cuba where differences in benthos and fish among low, intermediate, and high impact regions are consistently found (see Chap. 17, González-Díaz et al.), making understanding of local conditions a prerequisite for sound management, designing, and implementation of appropriate conservation measures. In summary, Cuban coral reefs are surrounded by progressively warmer waters and impacted by hurricanes and storms. They are accessible from populated coastlines, receive high fishing pressure from commercial and subsistence fishes, are exposed to land-based pollution, and affected by tourism. Cuban coral reefs are not receiving enough protection. Compliance and enforcement of fisheries, environmental, and MPA regulations are poor. There is not enough research to understand their status and trends in most of the zones. These factors limit the design and implementation of sound management and protection initiatives. However, these facts are likely reversible if management actions are implemented.

15.7

Recommendations for Management and Research

Cuban coral reef status has been studied extensively, but not all coral reef zones have received the same level of scientific research. Such studies should continue in a way that allows us to compare coral reefs at multiple spatial scales, have a better understanding of the factors that impact coral reefs, and assess their status through time. Several actions should be implemented, including the establishment of standardized research and monitoring methodologies, proper database storage, and share system and strong partnerships among scientists and scientific institutions to make knowledge more available for decision making. Collection of environmental, climate, anthropogenic, and ecological data of most of the Cuban coral reefs should be done at least yearly and should include biodiversity assessments of corals, macroalgae, D. antillarum, and fish (small, herbivorous, and commercial species and lionfish) based on regional efforts such as ReefCheck, Atlantic and Gulf Rapid Reef Assessment (AGRRA) initiative, and the Caribbean Global Coral Reef Monitoring Network. Our findings suggest several research topics that require the development of hypotheses for future studies: (1) role of chronic wave exposure on ecological processes related to density and biomass of small fish, coral cover, density of juvenile corals, and sponge density, (2) stages of phase shift in coral reefs and ecological processes involved including

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“traditional groups” (corals, macroalgae, herbivorous fish, D. antillarum) and “new groups” such as damselfish and wrasses, and (3) predation of and by lionfish on a combination of gradients of environmental variables, preys, potential predators, and human control. Fisheries regulations and management changes, MPA establishment, and tourism activities should include implementation of assessment of their performance following the BACI design (see Chap. 23, Figueredo-Martín and PinaAmargós). In the last 20 years, there have been significant changes on fisheries, MPA, and tourism management without scientific assessment, such as the approval of all Cuban MPAs, the banning of highly impacting gear (set nets in 2008 and finfish trawls in 2012), an increase of seafood demand due to the expansion of private business (from 2010 on) and of tourism on public and private sectors (from 2015 on), the recently passed new fisheries law, and presumably the increase of illegal fishing which has been estimated in few cases. However, illegal fishing data seem to be underestimated in Sabana-Camagüey (Claro et al. 2009), in Jardines de la Reina (Pina-Amargós et al. 2009), in Golfo de Ana María (Figueredo-Martín et al. 2014), and in the Southeastern fishing zone (Alzugaray et al. 2018). Research and monitoring with the focus on the impact of tourism on coral reefs, particularly scuba diving, snorkeling, and recreational fishing, should be continued and expanded. Enforcement needs to be systematic (daily) to reduce fishing pressure and positively impact fish. Currently, systematic enforcement is only accomplished in few marine areas, mostly MPAs. The high levels of enforcement in Jardines de la Reina are related to the multiple and robust partnership among stakeholders (fisheries inspection, tourism operator, protected areas managers, coast guard, environmental and scientific institutions). This is a model worthy of expanding nationwide if conditions allow. Besides, other alternatives should be considered to promote compliance with fisheries, biodiversity protection, and protected areas regulations in Cuba. Compliance based on strong enforcement is socially complex and economically prohibitive even for the most robust economies; alternatives that promote selfcompliance should be implemented (Salm et al. 2000; Friedlander et al. 2003b; Mora et al. 2006; Guidetti et al. 2008). High compliance with environmental and fisheries regulations is particularly critical for herbivorous fish because of their role in the health of coral reefs. These species are bycatch of commercial fisheries nationwide (Tamara Figueredo Martín and Fabián Pina Amargós, unpublished data) and targeted by subsistence fisheries, particularly in populated cities such as Havana (Durán et al. 2018). This fact potentially compromises the resilience of the coral reefs of Cuba. There are alternatives to the top-down enforcement options. Community-based coral reefs management models

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have been successfully implemented in the Caribbean and elsewhere. They are increasingly emulated by governments, as well as supported and assisted by non-governmental groups, fishermen associations, tourism operators, and international programs, working together to achieve sustainable use of resources in particular areas (Jupiter et al. 2014; Camacho and Steneck 2017; Fujita et al. 2017; Ayer et al. 2018; Domondon et al. 2021). Our results suggest a reduction in pollution reaching the Cuban marine environment in recent years. Still, cumulative effects of point and longtime pollution and the synergistic effect with other stressors might be occurring. Pollution should be monitored and research designed to test its influence on the coral reef biota, particularly in coral reefs near Havana, Cardenas, Varadero, Isabela de Sagua, Caibarién, Nuevitas, Guardalavaca, Trinidad, Cienfuegos, Girón, and Cayo Largo del Sur. Other anthropogenic factors such as river dams (for human and agricultural use and flood control) and causeway construction through the shallow waters of the Cuban shelf for tourism purposes precede most of our data. Still, they might continue to affect Cuban coral reefs through cumulative and synergistic actions. Although several studies have assessed their impact on the marine environment (Alcolado et al. 1999; SC 2008; CBD 2019), their scope is not enough to understand their effects on coral reefs. On the other hand, several factors such as mass sargassum arrival and plastic pollution as well as sedimentation increase in the coral reefs from beach nourishment have been occurring or increasing after our data timeframe. Their potential impact on Cuban coral reefs should be assessed. Other issues affecting Cuban reefs that were not directly considered in our study are the influence of economic, social, and cultural conditions that influence the environmental performance of the Cuban people and institutions. However, various anthropogenic factors such as population, fishing pressure, pollution, tourism, and enforcement indirectly include some of those social components. Acknowledgments The authors would like to dedicate this chapter to Dr. Pedro M. Alcolado Menéndez for his gigantic contribution to coral reef research in Cuba. The authors want to thank the many institutions and their personnel that through cooperative projects have recently advanced coral reef research and conservation in Cuba: Project “Atlantic and Gulf Rapid Reef Assessment,” Project “Sabana-Camaguey,” Project “Archipiélagos del Sur,” Instituto de Oceanología/Instituto de Ciencias del Mar, Centro de Investigaciones Marinas de la Universidad de la Habana, Centro de Investigaciones de Ecosistemas Costeros, Centro Nacional de Áreas Protegidas, Ecovida, Empresa para la Protección de la Flora y la Fauna, Centro de Estudios y Servicios Ambientales de Matanzas, Centro de Estudios y Servicios Ambientales de Villa Clara, Centro de Estudios y Servicios Ambientales de Cienfuegos, Centro de Investigaciones Medioambientales de Camagüey, Centro de Investigaciones y Servicios Ambientales de Holguín, BIOECO, and Centro de Investigaciones Pesqueras. Special thanks to Avalon-Marlin for logistical support in Jardines de la Reina and Wildlife Conservation

F. Pina-Amargós et al. Society Cuba Program for supporting this contribution. We appreciate the work of the editors and anonymous reviewers for their critical comments that have improved this chapter.

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Population Genetics of Cuba’s Scleractinian Corals

16

Gabriela Ulmo-Díaz, Jessy Castellanos Gell, Didier Casane, Alexis Sturm, Joshua Voss, and Erik García-Machado

Abstract

Cuba’s coasts support a large area of diverse and relatively healthy coral communities that are centrally located at the interface of the northern Caribbean Sea, the Gulf of Mexico, and the Atlantic Ocean, thus physically connected to coral reefs in the Gulf of Mexico and the Atlantic via oceanographic currents. However, we are only beginning to quantify the genetic diversity, structure, and connectivity among coral populations in Cuba. There are limited coral population genetic data available in the country, yet the studies that have been already conducted provide critical information for coral reef managers to increase the efficacy of their marine conservation approaches. In this chapter, we review the available population genetic data for scleractinian corals on the Cuban shelf and discuss the potential implications of these patterns on coral conservation actions in Cuba. Furthermore, we identify persisting gaps in our understanding of

G. Ulmo-Díaz (✉) · E. García-Machado Institute of integrative biology and systems, Université Laval, Québec, QC, Canada Centro de Investigaciones Marinas, Universidad de La Habana, Miramar, Playa, Ciudad Habana, Cuba e-mail: [email protected]; [email protected] J. Castellanos Gell Centro de Investigaciones Marinas, Universidad de La Habana, Miramar, Playa, Ciudad Habana, Cuba Gainesville, FL, USA e-mail: jessycastellanos@ufl.edu D. Casane Université Paris-Saclay, CNRS, IRD, UMR Évolution, Génomes, Comportement et Écologie, Gif-sur-Yvette, France e-mail: [email protected] A. Sturm · J. Voss Harbor Branch Oceanographic Institute, Florida Atlantic University, Fort Pierce, FL, USA e-mail: [email protected]

Cuban coral population genetics and recommend potential future studies to address these gaps. Finally, we propose novel and feasible methods for implementing conservation genetics to improve conservation management in Cuban waters. Keywords

Cuban reefs · Corals · Scleractinia · Population genetics

16.1

Introduction

Cuba’s coastal shelves are critical habitats for economically valuable and biodiverse scleractinian coral communities. Spalding et al. (2017) calculated that the tourism-related value of each km2 of reef on the Cuban shelf was US $57,585, not including the potential coastal protection value these reefs may provide to the archipelago (Beck et al. 2018). Additionally, Cuban coral populations are centrally located in the Tropical Western Atlantic and are heavily influenced by major regional oceanic current systems. Therefore, Cuban coral populations may act as a critical connectivity link among coral populations upstream in the Caribbean and Gulf and downstream in Florida, the western Bahamas, and perhaps even Bermuda (Schill et al. 2015). Marine genetic connectivity of coral reef-associated species, including fishes, across the Cuban archipelago, has been explored in depth in a previous review (García-Machado et al. 2018). However, due to the limited genetic data available for scleractinian coral species, no comparative analysis or review of genetic diversity and structure among coral species across the Cuban archipelago has been conducted until the present. In this chapter, we review genetic diversity and structure data from Cuban coral populations with an emphasis on the implications for conservation management. Additionally, we explore novel molecular approaches that could be implemented to further study these ecosystems. Of the 161 scleractinian corals species inhabiting Cuban waters

# The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 V. N. Zlatarski et al. (eds.), Coral Reefs of Cuba, Coral Reefs of the World 18, https://doi.org/10.1007/978-3-031-36719-9_16

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G. Ulmo-Díaz et al.

Table 16.1 Published research of population genetics in coral species in Cuba Species Montastraea cavernosa Orbicella palmata Orbicella annularis

Citation Sturm et al. (2020) Ulmo-Díaz et al. (2018) Foster et al. (2012)

(Chap. 8, Ferrer et al.), only a few have been included in genetic studies of Cuban coral populations across different spatial scales (Table 16.1; Fig. 16.1). Two of them focused on Orbicella (=Montastraea) faveolata and Montastraea cavernosa populations across the Cuban archipelago (Ulmo-Díaz et al. 2018; Sturm et al. 2020) and a third one sampled Orbicella (= Montastraea) annularis at three Cuban localities as part of a regional, Caribbean-wide study (Foster et al. 2012). All three of these studies utilized microsatellite markers, which are commonly used in population genetic studies, in conjunction with other approaches and genetic marker types including a larval dispersal model (Foster et al. 2012), the mitochondrial control region in Ulmo-Díaz et al. (2018), and single-nucleotide polymorphisms (SNPs) in Sturm et al. (2020, Table 16.1). Comparisons among these different genetic markers were presented in both Ulmo-Díaz et al. (2018) and Sturm et al. (2020) and have been reviewed extensively in the literature (Jeffries et al. 2016; Lemopoulos et al. 2019; D’Aloia et al. 2020; Sunde et al. 2020); thus, these should be referred to for a discussion of different marker classes and molecular approaches in coral population genetic studies.

Molecular markers Microsatellites (9) and SNP (9720) Mitochondrial control region, microsatellites (6) Microsatellites (6)

All three published Cuban coral population genetic studies focus on massive reef-building species (Orbicella faveolata, Orbicella annularis, and Montastraea cavernosa), and a fourth unpublished study (Ulmo-Díaz et al. in preparation) has generated genetic data for a branching coral species: Acropora palmata. All species studied so far are broadcast spawners, and there is no available genetic information for any brooding species in Cuba. This is an important caveat as different life history characteristics (i.e., reproductive mode, growth rate, and fecundity), ecological niches, and responses to stressors and abiotic factors may influence the observed patterns in population genetic structure and may have varying implications for informed conservation management decisions (Álvarez-Noriega et al. 2016; Razak et al. 2020).

16.2

Genetic Diversity

To compare estimates of genetic diversity using microsatellite markers, we downloaded the raw genetic data available in Foster et al. (2012), Ulmo-Díaz et al. (2018), and Sturm et al. (2020) and analyzed them in the population genetic program,

Fig. 16.1 Sampling sites in Cuba of coral species in published research of population genetics. Map made with QGIS and Natural Earth

16

Population Genetics of Cuba’s Scleractinian Corals

311

Table 16.2 Across localities averaged heterozygosity of three coral species in Cuba (see Table 16.1), using microsatellite markers. Observed heterozygosity (Ho), expected heterozygosity (He), and unbiased expected heterozygosity (uHe) Species Orbicella faveolata Orbicella annularis Montastraea cavernosa

Ho 0.736 0.781 0.658

He 0.779 0.729 0.64

GenAlEx v6.502 (Peakall and Smouse 2006, 2012), to calculate observed, expected, and unbiased expected heterozygosity values, common measures of genetic variation (Table 16.2). We should note that mean heterozygosity estimates averaged across loci are sensitive to sample size, number of loci, and “evenness of allele frequency” of each locus (Hale et al. 2012). Additionally, each of the studies in this comparison used different microsatellite loci developed for that specific species, which may vary in mutation rate and allelic diversity. Therefore, the following quantitative comparisons of genetic diversity across these coral species in Cuba were made as generalized assessments of geographic patterns in population genetic diversity and should be taken with caution. Mean observed and expected heterozygosity (Ho and He, respectively) across loci and localities are similar among both Orbicella faveolata and O. annularis species (Foster et al. 2012; Ulmo-Díaz et al. 2018), indicating similar levels of regional biodiversity within these two species. Mean heterozygosity estimates were lower for Montastraea cavernosa, but these values were highly influenced by the extremely low heterozygosity values observed in Banco de San Antonio, which only had a sample size of n = 2 (Sturm et al. 2020). Heterozygosity values were more similar to those reported for the other species in all other localities with larger sample sizes. Foster et al. (2012) and Ulmo-Díaz et al. (2018) sampled O. annularis and O. faveolata in Baracoa Beach, close to Havana City, and heterozygosity values were similar for both species in the site (Ho = 0.782 uHe = 0.759 in O. annularis; Ho = 0.729 uHe = 0.812 in O. faveolata). Both Sturm et al. (2020) and Ulmo-Díaz et al. (2018) sampled across areas in the Banco de San Antonio-Los Colorados Archipelago and Cayo Anclitas-Jardines de la Reina, respectively. In Cayo Anclitas-Jardines de la Reina, Ho is very similar in both species (Ho = 0.733 in M. cavernosa; Ho = 0.759 in O. faveolata), but unbiased expected heterozygosity is marginally higher in O. faveolata (uHe = 0.724 in M. cavernosa; uHe = 0.798 in O. faveolata). From a conservation genetics standpoint, it is a positive sign that the genetic diversity levels of this O. faveolata population remain relatively high despite recent severe population declines across the archipelago, especially when genetic diversity levels are similar to M. cavernosa, which remains one of the most dominant coral species in Cuba (González-Díaz et al. 2018).

uHe 0.812 0.748 0.693

# samples 82 67 78

#localities 5 3 8

Montastraea cavernosa samples from Banco de San Antonio were collected from the mesophotic zone (30–150 m; see mesophotic Chap. 14, Reed et al.), while O. faveolata samples from these locations were collected from shallow reefs. The estimated heterozygosity values of the mesophotic Banco de San Antonio M. cavernosa were much lower than the heterozygosity values for the shallow O. faveolata (Ho = 0.389, uHe = 0.519 in M. cavernosa; Ho = 0.739, uHe = 0.807 in O. faveolata). It is unknown if M. cavernosa from shallow depth zones exhibits similar heterozygosity values. Also, only two colonies of M. cavernosa from the mesophotic coral ecosystem (MCE) were collected at this location, which likely causes bias in the heterozygosity estimates for this population. Further sample collection in the MCE in Cuba is needed to determine if these mesophotic populations exhibit lower genetic diversity than their shallow counterparts. When contrasted, genetic diversity values of massive corals in Cuba are similar to those found elsewhere throughout the broader Caribbean. For example, heterozygosity values of M. cavernosa in Cuba are in the range of those values reported across the Belize Barrier Reef (Eckert et al. 2019) and across both shallow and mesophotic reefs in the Gulf of Mexico (Studivan and Voss 2018). Heterozygosity values reported for Cuban O. faveolata populations are within the range of values reported along the Colombian coast (Alegría-Ortega et al. 2021) and slightly higher than those found across the Caribbean and the Gulf of Mexico by Porto-Hannes et al. (2015) and Rippe et al. (2017). It should be noted that sets of microsatellite markers utilized in each study do not overlap completely; therefore, the differences in heterozygosity values could be due to this. Overall, these comparisons suggest that the genetic diversity levels among coral populations on Cuban reefs are relatively similar to those found elsewhere in the Caribbean. High levels of genetic diversity are critical to supporting coral population persistence and resilience in the face of unavoidable stressors associated with anthropogenic impacts and climate change.

16.3

Genetic Differentiation and Connectivity

Patterns of genetic differentiation between eastern and western sites in Cuba were observed for all three species of broadcast-spawning corals despite varying study approaches

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and marker types. We should note that Foster et al. (2012) found this East-West structure using a computational model, but could not validate it empirically, perhaps due to a lack of resolution among the six microsatellite markers. Additionally, Foster et al. (2012) found differentiation in a non-overlapped geographical area where the other two studies did not sample, which means that, to date, there is no way to empirically corroborate the differentiation observed between Siboney (CU3 in Foster et al. 2012) and the rest of the archipelago. In both M. cavernosa and O. faveolata, colonies from the locality Banco de San Antonio-Los Colorados Archipelago (the most northwestern locality) were genetically differentiated from the rest of the Cuban Archipelago, and especially sites in eastern Cuba. Montastraea cavernosa colonies sampled from the western Banco de San Antonio site were from the MCE, but O. faveolata from the same location were collected at shallow depths, suggesting that these northwestern populations exhibit limited gene flow with the rest of the archipelago regardless of depth. This would suggest two hypothetical scenarios: that this area is isolated and possibly self-seeding, or that it is more connected to reef populations upstream of Cuba via the Loop Current/Florida Current System. Both scenarios would require specialized management for this area, and possibly concerted regional and international management efforts. In recognition of the physical and likely larval connectivity (Holstein et al. 2014; Schill et al. 2015), among western Cuba and other reefs in the Tropical Western Atlantic, a Memorandum of Understanding agreement was drafted among the United States National Oceanic and Atmospheric Administration, the United States National Park Service, and the National Center of Protected Areas of the Ministry of Science, Technology, and Environment of the Republic of Cuba to form a sister-sanctuary management relationship between Banco de San Antonio and Guanahacabibes marine protected areas in western Cuba, the Flower Garden Banks National Marine Sanctuary in the Gulf of Mexico, and the Florida Keys National Marine Sanctuary (November 18th, 2015). Sturm et al. (2020) suggest that M. cavernosa populations in western Cuba may also represent a link among populations of this species in Mexico, Belize, and the Dry Tortugas. Network connectivity models suggest that western Cuba could be a hotspot of broad Caribbean connectivity (particularly as source) for populations of O. annularis and Porites astreoides, as well as some fish species (Holstein et al. 2014). Regional studies are needed to more accurately quantify the connectivity between western Cuban coral populations and other reefs within the Tropical Western Atlantic region. More research is also needed to characterize the connectivity patterns among reefs across both shallow (