Echinodermos Peru PDF

Echinoderm Research and Diversity
in Latin America
Juan José Alvarado
Francisco Alonso Solís-Marín
Editors
Echinoderm Research and

Diversity in Latin America
123
Editors
Juan José Alvarado Francisco Alonso Solís-Marín
Centro de Investigaciónes en Ciencias Instituto de Ciencias del Mar, y Limnologia
del Mar y Limnologia Universidad Nacional Autónoma de México
Universidad de Costa Rica México City
San José Mexico
Costa Rica
ISBN 978-3-642-20050-2 ISBN 978-3-642-20051-9 (eBook)

DOI 10.1007/978-3-642-20051-9
Springer Heidelberg New York Dordrecht London
Library of Congress Control Number: 2012941234
Ó Springer-Verlag Berlin Heidelberg 2013

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We would like to dedicate this book to the
pioneers of echinoderms research in Latin
American. To Marı´a Elena Caso, Irene
Bernasconi, Luis Roberto Tomassi and
Evelyn Zoppi de Roa.
Foreword
This is a remarkable book. First and foremost, it compiles an immense amount of

literature on all aspects of echinoderm biology and ecology over an extremely
broad geographical range. This will be extremely useful because much of the
literature is relatively unknown to the world at large. The description of the
oceanography and marine environments of each country in a comprehensive way
will be of considerable interest to marine biologists in general.
One of the most interesting aspects of this book is the section devoted to the
history of echinoderm studies in each country. The first reports about echinoderms
were isolated and sporadic, not systematic studies. These generally did not begin
until the 1850s with various foreign expeditions along the coasts of Latin America.
Studies of echinoderms by workers in Latin American countries themselves did
not begin to develop appreciably until the 1950s. This has changed in a dramatic
way. In the latter part of the twentieth century, interest in marine biology and
recognition of its importance in terms of basic science, the environment and
conservation, and economics increased greatly. This led to an increase in the
number of marine biologists in general and of echinoderm biologists in particular.
The amount of research activity that has developed since then is impressive.
Another thing that impressed me, apparent in the book, is the passion of Latin
American biologists for echinoderms, their commitment to conservation, their
desire to assist their countries through their efforts, and their collegiality.
Florida, USA, March 2012 John M. Lawrence

Department of Integrative Biology,
University of South Florida,
Tampa
vii
Foreword
Juan José Alvarado is one of those students who, from very early on, denoted a
clear interest in Marine Science and at some point defined his area of interest.
When he started looking at the echinoderms of Costa Rica and later, at those
reported and collected from Coco Island National Park, he found what he was
looking for. Since then, he has been actively carrying out research on echinoderms,
and most important, he has drawn attention to the activities regarding this group
that are taking place in Latin America. First, with the publication of Research on
Echinoderms in Latin America in 2005 (Revista de Biología Tropical, Volume 53,
Supplement 3: 387 p.). Then, with the creation of the Iberoamerican Echinoderm
Network Red Iberoamericana de Equinodermos (RIE) in 2006. And in 2008, with
the publication of another Special Issue on echinoderm research in Latin America:
Revista de Biología Tropical, 56 (Suppl. 3): 360 p. Now, together with Francisco
Solís-Marín from the Universidad Nacional Autónoma de México (UNAM), they
have put together an outstanding compilation of echinoderm research in Ibero-
america. The dedication of this book to the pioneering Latin American echinoderm
researchers is a tribute to their great work and an acknowledgment of the fruit their
early and many times solitary work has borne. It is a great honor for me to write
this Foreword, for I consider their work a significant step for biological research in
the region..
San José, Costa Rica, March 2012 Jorge Cortés

Centro de Investigación en Ciencias
del Mar y Limnología (CIMAR),
Universidad de Costa Rica
ix
Preface
The Iberoamerican Echinoderms Network (Red Iberoamericana de Equinodermos)

(RIE) (http://zicatela.umar.mx/*redequinos/2/) was created in August 2006 at the
12th International Echinoderms Congress in New Hampshire, United States, as an
initiative to strengthen academic connections between Iberoamerican echinoderm
researchers. The RIE seeks to fortify the links of researchers in the different
countries of the region, being a bridge of communication and exchange of ideas. It
seeks to be the platform for discussion and communication of priority projects for
Latin America echinoderms. It seeks to offer the opportunity of collaborative
work, to place the region among the principal research and conservation regions of
the world. Likewise, the RIE intends to support students interested in echinoderm
research. To enrich their formation, it will offer information about researchers who
could help in their projects, courses, seminars, or symposia, as well as providing
literature. The goal of the RIE is to develop workshops, congresses, special
journals supplements and books, and to support the development of the knowledge
of the echinoderms in Latin America.
The first effort of the RIE was the preparation and publication of a Special Issue
of the Revista de Biología Tropical (International Journal of Tropical Biology and
Conservation) (Alvarado, J. J. and Cortés, J. (Editors). 2005. Research on Echi-
noderms in Latin America, Volume 53, Supplement 3: 387 p.) (http://www.ots.ac.
cr/tropiweb/intpages/suppl/sup53-3.html). This initiative was born during the 11th
International Echinoderms Congress in Münich, Germany, in 2003. This Special
Issue compiled 28 papers from 12 countries by 62 authors, touching topics as diverse
as paleontology in the Uruguayan region and fisheries models of sea cucumbers in
Baja California. The second output of the RIE was a second supplement of the
Revista de Biología Tropical (Alvarado, J.J. and Cortés, J. (Editors). 2008. Research
on Echinoderms in Latin America II, Volume 56, Supplement 3: 360 p.) (http://www.
ots.ac.cr/tropiweb/intpages/suppl/sup56-3.html). This issue compiled 21 papers
from eight countries by 46 authors in Latin America (including an invited paper from
the Canary Islands). Those papers demonstrated the high degree of knowledge of
some countries in topics such as diversity, ecology, aquaculture, and fisheries. It also
identified countries where more efforts are needed.
xi
xii Preface
The third effort was the establishment of the first Latin American Echinoderm
Congress (Congreso Latinoamericano de Equinodermos-CLE) held in Puerto
Madryn, Argentina, between 13 and 18 November 2011. This congress was
organized by the Centro Nacional Patagónico (CENPAT), the Museo Argentino de
Ciencias Naturales ‘‘Bernardino Rivadavia’’ (MACN), and the Universidad Nac-
ional de la Patagonia San Juan Bosco. A total of 112 people participated in this
event with 76 oral presentations and 54 Poster, covering the following themes:
biogeography, biochemistry, ecology, physiology, paleontology, fisheries and
aquaculture, reproduction and development, and taxonomy, systematics, and
evolution.
This book represents the fourth output and the confirmation of the commitment
of the RIE with the international scientific community. The book represents an
extensive recompilation of information about all the research done in the region
from theses, reports, books, and scientific journals. We were able to identify the
strengths and weaknesses of each country and also allow the international scien-
tific community to be able to access information that was held in reports, theses,
and local papers that were written in Spanish, allowing a greater diffusion of the
results and conclusions obtained. Our goal is to improve the echinoderm research
on the region and with this road map establish future research in collaboration with
the different laboratories and researchers that participated in this book.
Juan José Alvarado

Francisco Solís-Marín
Acknowledgments
First of all, we thank each of the authors of the chapters that have made this
compilation about Echinoderm Research and Diversity in Latin America possible,
to share their knowledge and passion for such a star group. Muchas gracias!
We also would like to acknowledge the help and support of all the reviewers who
with their comments and suggestions improved this book: Chris Pomory, Cynthia
Lara de Castro, David Pawson, Harilaos Lessios, John Pearse, Manuel Rey Méndez,
Marc Eléaume, Martin Brogger, Nieves Elvira, Pablo González, Philip Lambert,
Rosa del Valle, Tim O’hara, Tom Hopkins. We wish to grant special
credit to John Lawrence and Jorge Cortés who with all the patience, interest,
and care reviewed each of the chapters, leaving their wisdom on every page.
We wish to grant special credit to all the students from the Laboratorio de
Sistemática y Ecología de Equinodermos from the Instituto de Ciencias del Mar y
Limnología, Universidad Nacional Autónoma de Mexico, for their help with the
final elaboration of the appendix: Mauricio Valdés De Anda, Julio Adrián Arriaga
Ochoa, Andrea Alejandra Caballero Ochoa, Tania Pineda Enríquez, Carolina
Martín Cao-Romero, Yoalli Quetzalli Hernández Díaz, Alejandra Martínez Melo,
Viridiana Tapia Ramírez, Guadalupe Bribiesca Contreras, Pedro Josué Garcés
Solchaga, Nancy Escandón Flores, Lucia Alejandra Hernández Herrejón and
Carlos García Linares.
We appreciate the help of Monica Chavez in preparing all the figures in this
book and Cindy Fernandez who always believed in this project and was a keystone
for the final elaboration. JJ Alvarado is grateful to CONACYT (México) and
CONICIT-MICIT (Costa Rica). Finally, we thank Marion Schnider and
SPRINGER for their interest in this book.
xiii
Contents
1 Echinoderm Research and Diversity in Latin America. . . . . . . . . 1

Juan José Alvarado and Francisco Alonso Solís-Marín
2 The Echinoderms of Mexico: Biodiversity, Distribution

and Current State of Knowledge. . . . . . . . . . . . . . . . . . . . . . . . . 11
Francisco A. Solís-Marín, Magali B. I. Honey-Escandón,
M. D. Herrero-Perezrul, Francisco Benitez-Villalobos,
Julia P. Díaz-Martínez, Blanca E. Buitrón-Sánchez,
Julio S. Palleiro-Nayar and Alicia Durán-González
3 Central America Echinoderms: Diversity, Ecology

and Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Juan José Alvarado, Enrique Barraza and Tito Isaac Sancho-Mejías
4 Echinoderm Diversity in Panama: 144 Years of Research

Across the Isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Simon E. Coppard and Juan José Alvarado
5 Echinoderms of Colombia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

Milena Benavides-Serrato, Giomar Helena Borrero-Pérez,
Jaime R. Cantera K, Mishal Cohen-Rengifo and Raúl Neira
6 Echinoderms of Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

Jorge Sonnenholzner, Margarita Brandt, Vanessa Francisco,
Alex Hearn, María Luzuriaga, Paulina Guarderas
and Juan Carlos Navarro
xv
xvi Contents
7 Echinoderms from Venezuela: Scientific Recount,

Diversity and Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
César Lodeiros, Alberto Martín, Vanessa Francisco, Nicida Noriega,
Yusbelly Díaz, Jeny Reyes, Orangel Aguilera and José Alió
8 Echinoderms of Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277

Yuri Hooker, Elba Prieto-Rios and Francisco A. Solís-Marín
9 Echinoderm from Brazil: Historical Research and the Current

State of Biodiversity Knowledge . . . . . . . . . . . . . . . . . . . . . . . . . 301
Carlos R. R. Ventura, Michela Borges, Lúcia S. Campos,
Leticia V. Costa-Lotufo, Carolina A. Freire, Valéria F. Hadel,
Cynthia. L. C. Manso, José R. M. C. Silva, Yara Tavares
and Cláudio G. Tiago
10 Echinoderm Research in Uruguay . . . . . . . . . . . . . . . . . . . . . . . 345

Sergio Martínez
11 Echinoderms from Argentina: Biodiversity, Distribution

and Current State of Knowledge . . . . . . . . . . . . . . . . . . . . . . . . 359
Martín I. Brogger, Damián G. Gil, Tamara Rubilar,
Mariano I. Martinez, María Enriqueta Díaz de Vivar,
Mariana Escolar, Lucía Epherra, Analía F. Pérez
and Alejandro Tablado
12 Echinoderms of the Cuban Archipelago. . . . . . . . . . . . . . . . . . . 403

Rosa del Valle-García, Mercedes Abreu-Pérez, Francisco
Alonso Solís-Marín and Alfredo Laguarda-Figueras
13 Recent Echinoderms from Hispaniola . . . . . . . . . . . . . . . . . . . . 425

Alejandro Herrera-Moreno and Liliana Betancourt
14 Review of Echinoderm Research in Puerto Rico,

with the Focus on Biological and Ecological Aspects . . . . . . . . . 437
Stacey M. Williams, Milena Benavides-Serrato, Jose E. García-Arrarás,
Edwin A. Hernández-Delgado and Ruber Rodriguez Barreras
15 Echinoderms of the Canary Islands, Spain. . . . . . . . . . . . . . . . . 471

José Carlos Hernández, Sabrina Clemente, Fernando Tuya,
Angel Pérez-Ruzafa, Carlos Sangil, Leopoldo Moro-Abad
and Juan José Bacallado-Aránega
Contents xvii
16 Latin America Echinoderm Biodiversity and Biogeography:

Patterns and Affinities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511
A. Pérez-Ruzafa, J. J. Alvarado, F. A. Solís-Marín, J. C. Hernández,
A. Morata, C. Marcos, M. Abreu-Pérez, O. Aguilera, J. Alió,
J. J. Bacallado-Aránega, E. Barraza, M. Benavides-Serrato,
F. Benítez-Villalobos, L. Betancourt-Fernández, M. Borges,
M. Brandt, M. I. Brogger, G. H. Borrero-Pérez,
B. E. Buitrón-Sánchez, L. S. Campos, J. R. Cantera, S. Clemente,
M. Cohen-Renfijo, S. E. Coppard, L. V. Costa-Lotufo,
R. del Valle-García, M. E. Díaz de Vivar,
J. P. Díaz-Martínez, Y. Díaz, A. Durán-González, L. Epherra,
M. Escolar, V. Francisco, C. A. Freire, J. E. García-Arrarás,
D. G. Gil, P. Guarderas, V. F. Hadel, A. Hearn,
E. A. Hernández-Delgado, A. Herrera-Moreno, M. D Herrero-Pérezrul,
Y. Hooker, M. B. I. Honey-Escandón, C. Lodeiros, M. Luzuriaga,
C. L. C. Manso, A. Martín, M. I. Martínez, S. Martínez,
L. Moro-Abad, E. Mutschke, J. C. Navarro, R. Neira, N. Noriega,
J. S. Palleiro-Nayar, A. F. Pérez, E. Prieto-Ríos, J. Reyes,
R. Rodríguez-Barreras, T. Rubilar, T. I. Sancho-Mejia, C. Sangil,
J. R. M. C. Silva, J. I. Sonnenholzner, C. R. R. Ventura, A. Tablado,
Y. Tavares, C. G. Tiago, F. Tuya and S. M. Williams
Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 543
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 655
Chapter 1
Echinoderm Research and Diversity
in Latin America
Juan José Alvarado and Francisco Alonso Solís-Marín
1.1 Introduction
The phylum Echinodermata consists of approximately 7,000 living species and

13,000 extinct species (Pawson 2007). Studies on echinoderms have focused on
experimental ecology due to their important role in marine ecosystems, with
several species identified as ‘‘keystone species’’ (Uthicke et al. 2009). These
species can have effects on community structure and function through several
trophic levels and on a wide geographic scale (Birkeland 1989). However, growing
global pressures resulting from fishing echinoderms have put them under threat
(Micael et al. 2009).
Latin America (LA) comprehends all the countries in America that speak
Spanish or Portuguese; including the French speaking country of Haiti, that is
located in Hispaniola Island, shared with Dominican Republic. LA includes
Mexico, Central America, and the islands of the Caribbean and South America.
However, the term has been used with different meanings that have created some
confusion (Colburn 2002). In total it includes 22 countries with an area of
J. J. Alvarado (&)
Centro de Investigación en Ciencias del Mar y Limnología (CIMAR),
Universidad de Costa Rica, San Pedro, 11501-2060, San José, Costa Rica
e-mail: juanalva76@yahoo.comjuan.alvarado@ucr.ac.cr
J. J. Alvarado
Posgrado en Ciencias Marinas y Costeras, Universidad Autónoma
de Baja California Sur, La Paz, México
F. A. Solís-Marín
Colección Nacional de Equinodermos ‘‘Ma. E. Caso Muñoz’’,
Laboratorio de Sistemática y Ecología de Equinodermos, Instituto de Ciencias
del Mar y Limnología (ICML), Universidad Nacional Autónoma de México (UNAM),
Apdo. Post 70-305, 04510, México, DF, Mexico
e-mail: fasolis@cmarl.unam.mx
J. J. Alvarado and F. A. Solís-Marín (eds.), Echinoderm Research and Diversity 1

in Latin America, DOI: 10.1007/978-3-642-20051-9_1,
2 J. J. Alvarado and F. A. Solís-Marín
21,069,501 km2, and according to the Sea around Us project. It possesses

17,010,693 km2 of exclusive economic zones and 2,706,433 km2 of shelf area
(www.seaaroundus.org). LA is a megadiverse region, including six of the 17
megadiverse countries around the world: Brazil, Colombia, Ecuador, Mexico, Peru
and Venezuela (Michán and Llorente-Bousquets 2010). However, scientific pro-
duction in terms of science and engineering is considerably lower (3.2 %) in 2000
than in the United States and Canada (36.7 %) (Hermes-Lima et al. 2007).
This book presents a change in the research paradigm of echinoderms in this
region. It gives an exhaustive review of all studies that has been done. All chapters
have been written by local scientists. It compiles reports, thesis, and papers in
regional and international journals. John Pearse (personal communication) noted
that the common opinion that the echinoderms knowledge in LA was ‘‘scarce with
information often fragmented, but quite the contrary, showing that there is an
impressive amount of work’’. Until today the greatest regional reviews about
echinoderms were the compilation by Linda Yvonne Maluf (1988) on composition
and distribution along the Eastern Pacific and the masterpiece presented by Clark
and Downey (1992) about taxonomy and distribution of the starfishes of the
Atlantic. The contribution by Hendler et al. (1995) about the echinoderms in the
Caribbean Sea not only compiled taxonomic and distribution information, but also
provided ecological data and high quality pictures of the species and their iden-
tifiable features.
This book is divided in 16 chapters (including this one) and two taxonomic lists
(Appendix) (one from the Pacific Ocean and one from the Caribbean Sea and
Atlantic Ocean including the Canary Islands) with information about presence/
absence, bathymetric range and the type of habitat or substrate where the species
have been found. These taxonomic lists were built following the criteria of Clark
(1931, 1947) for Crinoidea; Clark (1989, 1993, 1996), Clark and Downey (1992),
Clark and Mah (1999) for Asteroidea; Matsumoto (1915), Stöhr and O’Hara
(2007) for Ophiuroidea; Smith (1984) for Echinoidea and Panning (1929, 1934a,b,
1935), Pawson and Fell (1965) for Holothuroidea. The World Register of Marine
Species (Appeltans et al. 2011) was also consulted for all species. From those
taxonomic lists we present only the species that are fully identified, avoiding the
presence of ‘‘sp.’’ reports. However, in each chapter the authors included ‘‘sp.’’
reports in their species numbers. Fourteen chapters deal with studies done in
countries, regions or islands (Fig. 1.1): Mexico, Central America (Belize, Guate-
mala, Honduras, El Salvador, Nicaragua, Costa Rica) and Panama, Colombia,
Ecuador, Venezuela, Peru, Brazil, Uruguay, Argentina, Cuba, Hispaniola Island
(Haiti and Dominican Republic), Puerto Rico and the Canary Islands. The final
chapter has a biogeographical and diversity analysis of the two taxonomic lists.
Unfortunately a chapter about Chilean Echinoderms could not be included in this
book, but the Pacific taxonomic list includes all the species present in that country
plus Easter Island.
Each country chapter contains six sections. 1. An introduction with a brief
description of: the marine environments, length of the coast, continental shelf area
and oceanographic conditions. 2. A historic account of the investigations that have
1 Echinoderm Research and Diversity in Latin America 3
Fig. 1.1 The American continent with the countries participating on the book indicated in colors
been done on echinoderms with information on the first expeditions and collections,
as well as ecology, reproduction, paleontology, bioerosion, competition, among
others studies. 3. Distributional patterns and diversity. 4. If specific fishery or
aquaculture activity exists, this is explained in a section, giving the name of the
species extracted, numbers, and seasons of extraction, regulations, aquaculture
activities, marketing and the benefits to local communities from these activities.
5. A section on threats to echinoderms, explaining the main environmental and
anthropogenic threats that are affecting the echinoderm populations and how. It also
examines whether some echinoderms are affecting the environment (overpopula-
tions, extreme grazing, bioerosion). 6. The final section contains general conclu-
sions from the review and recommendations for future research to meet the needs of
the country.
This book covers a large part of America, excluding only Canada, the United
States, the Lesser Antilles, Guyana, Surinam and French Guiana. For the latter two
countries and the French overseas territory, information on echinoderms is
available from the publications by Cherbonier (1959a, b, c, d, e), Walenkamp
(1976, 1979), Jangoux (1978) and Le Loeuff and Von Cosel (2000).
1.2 Echinoderm Research in LA: An Overview
Latin American echinoderm studies started in the 13th century. Bernardino de

Sahagún (1499–1590) a Spanish Franciscan friar who wrote about the Aztec
culture in Mexico (The General History of the Things of New Spain) before the
Spanish conquerors arrived in Central America (Florentine Codex). He describes
and explains ancient native religion, beliefs, practices, deities of the Aztecs. His
magnificent document was prepared in 12 books. Book number 11 describes the
‘‘Earthly Things’’ (properties of animals, birds, fish, trees, herbs, flowers, metals,
stones, and colors). In this volume he draws, describes and explains some inver-
tebrate forms used by the Aztecs during their daily life, including some sea
urchins, starfish and probably sea cucumbers. This is probably the earliest record
of echinoderms in the Americas.
Louis Éconches Feuillée (1660–1732), a French member of the Order of the
Minims, was an explorer, geographer, and botanist who travelled to Chile,
Argentina and Peru on different expeditions from 1709 to 1711. He gave the first
echinoderm references for these Latin American countries. The first formal
taxonomic description of a Latin American echinoderm was made by the famous
naturalist Carl Nilsson Linnæus (1707–1778) in his 1758 publication of Systema
Naturae. In this classification of all living organisms, he formally described several
echinoderm species from Latin American coasts (i.e. Oreaster reticulatus,
Echinometra lucunter and Clypeaster rosaceus). Jean-Baptiste Pierre Antoine de
Monet de Lamarck (1734–1829), a French naturalist and an early proponent of the
idea that evolution occurred and proceeded in accordance with natural laws,
published in 1816 one of his masterpieces, ‘‘Histoire Naturelle des Animaux sans
Vertébres’’. He reported new echinoderm species collected in LA. A contemporary
of Lamarck was the Chilean Juan Ignacio Molina (1740–1829) better known as
‘‘Abate Molina’’. In 1782 he published his work ‘‘Saggio sulla Storia Naturale del
Chile’’ where he described two new species of sea urchins from Chilean waters,
Echinus albus (=Loxechinus albus), and Tetrapygus niger.
Jean-Louis-Rodolphe Agassiz (1807–1873), a Swiss paleontologist, geologist
and a prominent zoologist, was a professor at Harvard University in the United
States. In several papers he described echinoid fauna from the LA. Rudolf
Amandus Philippi Krumwiede (1808–1904) was a German-Chilean paleontologist
and zoologist who described and published new and already known echinoderm
species from Chile. His most ‘‘famous’’ species is probably the sea urchin
Diadema antillarum, from the Caribbean, which he described in 1845. Theodore
Lyman (1833–1897), an U.S. zoologist who studied under the supervision of

Professor Louis Agassiz. He worked many years at the Museum of Comparative
Zoology of Harvard University and described many species of Latin American
echinoderms, mainly ophiurans. Alexander Emanuel Rodolphe Agassiz
(1835–1910), a Swiss-born U.S. marine zoologist and oceanographer, was the son
of Louis Agassiz. He emigrated in 1849 to the United States where he conducted
significant systematic zoological work on Latin American echinoderms (e.g.
starfish and echinoids).
Rafael Arango y Molina (1837–1893), a Cuban zoologist who wrote the first
echinoderm records of the archipelago, worked together with L. F. Pourtalés
(1824–1880), a Swiss who also worked on LA echinoderms and visited Cuba
around 1867.
The transition from 19th to 20th centuries was characterized in general by large
expeditions that reported new discoveries on the echinoderm fauna of LA. These
included studies by several of the most prominent echinoderm researchers such as
Addison Emery Verrill (1839–1926), Theodoro Mortensen (1868–1952), Hubert
Lyman Clark (1870–1947), Austin Hobart Clark (1880–1954), Elizabeth
Deichmann (1896–1975), María Codocedo Rojas (1909–1989), Irene Bernasconi
(1896–1989), Gustave Cherbonier (1909–1995), María Elena Caso Muñoz
(1915–1991), Maureen E. Downey (1920–2000), Ailsa McGown Clark (1926),
Ignácio M. Brito (1927–2001), Patricio Sánchez Reyes (1928–1999), Cinthya
Anne Gust Ahearn (1952–2008) and Isabel Pérez Farfante de Canet (1916–2009).
In this century, many active echinoderm specialists still work on the Latin
American fauna: Luiz Roberto Tommasi (Universidade de São Paulo, Brasil),
David L. Pawson (Smithsonian Institution, Washington, D.C, U.S.A.), Claude
Massin (Institut Royal des Sciences naturelles de Belgique, Brussels, Belgium),
John M. Lawrence (University of South Florida, U.S.A.), Alfredo Laguarda
Figueras (Instituto de Ciencias del Mar y Limnología, Universidad Nacional
Autónoma de México), Gordon Hendler (Los Angeles County Museum,
Los Angeles California, U.S.A), Rich Mooi (California Academy of Sciencies,
California, U.S.A.), Alberto Larraín Prat (Universidad de Concepción, Chile),
Harilaos A. Lessios (Smithsonian Tropical Research Institute, Panama) and many
others who are still actively increasing our knowledge of this fascinating inver-
tebrate group.
1.3 General Results
Most Latin American countries have a common beginning in echinoderm research,

with the presence of European and United States expeditions, followed by a period
of research by regional scientists in the middle of the twentieth century.
Researchers like María Elena Caso, Irene Bernasconi and Luiz Roberto Tomassi
are prominent. Most of the specimens collected by these expeditions are in foreign
museums in the United States (Los Angeles County Museum of Natural History,
Museum of Comparative Zoology at Harvard University, California Academy of

Sciences Natural History Museum, Smithsonian Institution National Museum of
Natural History, The Scripps Institution of Oceanography Benthic Invertebrate
Collection), France (Museum National d’Histoire Naturelle), Germany (Museum
für Naturkunde) and Denmark (Zoologisk Museum-Staten Naturistorike Museum).
However the Coleccción Nacional de Equinodermos Ma. Elena Caso Muñoz at the
Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autonoma of
Mexico, holds one of the largest collection of Latin American specimens with
more than 81,000.
During the last three decades each country has set its own path, being spe-
cialized in one or two areas. The common line of research is basic ecology
(distribution, population sizes, and diversity) and taxonomy. However some
countries stand out in particular lines of research: physiology (Puerto Rico and
Argentina), larval development (Venezuela and Mexico), aquaculture (Venezuela,
Mexico, Brazil), fisheries (Mexico), evolution (Panama), biogeography (Panama,
Mexico), paleontology (Mexico, Uruguay), reproduction (Panama, Mexico,
Brazil) and secondary metabolites (Argentina). Most of the research has focused
on the Echinoidea and Asteroidea, with very few works on Crinoidea. This last
class needs an increased research effort. Regular echinoids have taken much of the
attention. Among them is the genus Diadema, due to the massive die-offs of
D. antillarum in the Caribbean, the barrens grounds produce by over populations
of this species on the Canary Islands, and the predominance along the coral reefs
of the Eastern Tropical Pacific of D. mexicanum with differential bioerosives
impacts depending on its abundance. There is a need to increase the research on
echinoderms in Uruguay, Central America, Haiti, Dominican Republic, Cuba, and
Peru. It is also necessary to develop taxonomical, ecological (i.e. behavior,
predation), physiological and development studies. It is essential to improve
research in the Ophiuroidea and Holothuroidea.
On the other hand, most of the threats to echinoderms and their needs are
shared. The most common threats are bad coastal planning, sedimentation, coastal
pollution, illegal extraction, inefficient fisheries management, overfishing of
predators, lack of protection, lack of baselines studies before the establishment of a
sea cucumber fishery, lack of resources and lack of taxonomists. It is extremely
urgent to support fishery practices with basic ecological studies, to understand the
biology and ecology of each species before the establishment of an extraction
activity or quota.
A great part of the research that has been done in LA can be found in BSc.,
‘‘Licenciatura’’, MSc. and PhD thesis. Some are regional and national reports or
abstracts of projects that have not had international distribution. The majority of
those publications are in Spanish, and published in regional journals like Acta
Científica Venezolana, Boletín de Investigaciones Marinas y Costeras, Ciencias
Marinas, Gayana Zoológica, Hidrobiológica, Océanides, Revista de Biología
Marina y Oceanografía, Revista de Biología Tropical and Revista Ciencia y Mar.
Many publications are in journals of museums or research centers (e.g. Revista del
Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia, Anales del
Table 1.1 Number of species in each Echinodermata class from the taxonomic list (TL) in the
Appendix
Total Pacific Caribbean and Atlantic Common species between
TL TL TLs
Crinoidea 82 21 63 2
Asteroidea 392 193 228 29
Ophiuroidea 521 211 340 30
Echinoidea 242 118 137 13
Holothuroidea 302 174 158 30
Total 1539 717 926 104
spp. with – 620 836 –
bathymetric (86 %) (92 %)
information
spp. with habitat – 473 630 –
information (66 %) (70 %)
Insituto de Ciencias del Mar y Limnología de la Universidad Nacional Autónoma

de México). These have a good level of national and local distribution, but not
regionally. There is a need to promote the publication of thesis in local and
regional journals, as well to encourage publication in English to have a broader
audience and distribution.
A total of 1,539 species (Table 1.1) (21.8 % of the worldwide species
accordingly to Pawson 2007) can be found in the taxonomic lists (see Appendix),
with 717 species in the Pacific and 926 in the Caribbean and Atlantic, and 104
species shared in both ocean basins. The richest class is Ophiuroidea with 521
species, followed by the class Asteroidea with 392 species. All classes have the
richest number of species in the Caribbean and Atlantic in contrast with the
Pacific, except for the class Holothuroidea, which is richest in the latter.
From the Pacific list, 86 % of the species give bathymetric information and
66 % give information on the habitat. From the Caribbean and Atlantic list, 92 %
and 70 % of the species give bathymetric and habitat information, respectively.
This indicates there is a greater understanding of the environment conditions were
this organisms inhabits in the Caribbean and Atlantic than in the Pacific. This is
probably due to a greater and continuous research effort in this region (i.e. Pawson
et al. 2009).
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Chapter 2
The Echinoderms of Mexico: Biodiversity,
Distribution and Current State of
Knowledge
Francisco A. Solís-Marín, Magali B. I. Honey-Escandón,

M. D. Herrero-Perezrul, Francisco Benitez-Villalobos,
Julia P. Díaz-Martínez, Blanca E. Buitrón-Sánchez,
Julio S. Palleiro-Nayar and Alicia Durán-González
F. A. Solís-Marín (&) M. B. I. Honey-Escandón A. Durán-González

Laboratorio de Sistemática y Ecología de Equinodermos, Instituto de Ciencias del Mar y
Limnología (ICML), Colección Nacional de Equinodermos ‘‘Ma. E. Caso Muñoz’’,
Universidad Nacional Autónoma de México (UNAM), Apdo. Post. 70-305, 04510, México,
D.F., México
A. Durán-González
e-mail: aliciad@cmarl.unam.mx
M. B. I. Honey-Escandón
Posgrado en Ciencias del Mar y Limnología, Instituto de Ciencias
del Mar y Limnología (ICML), UNAM, Apdo. Post. 70-305, 04510, México, D.F., México
e-mail: mbihoney@hotmail.com
M. D. Herrero-Perezrul
Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico Nacional,
Ave. IPN S/n Col., Playa Palo de Santa Rita, CP. 23096, La Paz, BCS, México
e-mail: dherrero@ipn.mx
F. Benitez-Villalobos J. P. Díaz-Martínez
Instituto de Recursos, Universidad del Mar, Carretera a Zipolite km 1.5,
Puerto Ángel, Oaxaca, México
e-mail: fvb@angel.umar.mx
J. P. Díaz-Martínez
e-mail: lobi2@hotmail.com
B. E. Buitrón-Sánchez
Departamento de Paleontología, Instituto de Geología UNAM,
04510, Mexico, D.F., México
e-mail: blancab@servidor.unam.mx
J. S. Palleiro-Nayar
Centro Regional de Investigación Pesquera, Instituto Nacional de Pesca,
Apartado postal 1305, Ensenada, BC, México
e-mail: juliopalleiro@yahoo.com.mx

12 F. A. Solís-Marín et al.
Fig. 2.1 Mexico with its principal islands, archipelago, marine lagoons and coasts. Fishing area of
sea urchins: Strongylocentrotus franciscanus (grey areas), fishing areas of Isostichopus fuscus in the
Gulf of California and Bahia Magdalena (grey circles), and the geographic location where ecological
studies of echinoderms have been carried out. (1) Salcedo-Martínez et al. (1988); (2) Caso et al.
(1994); (3) Reyes-Bonilla and Calderon-Aguilera (1999); (4) Benítez-Villalobos (2001); (5) Reyes-
Bonilla et al. (2005); (6) Cintra-Buenrostro et al. (2005); (7) Zamorano and Leyte-Morales (2005a);
(8) Zamorano and Leyte-Morales (2005b); (9) Holguín-Quiñones et al. (2008); (10) Rios-Jara et al.
(2008); (11) López-Uriarte et al. (2009); (12) Zamorano and Leyte-Morales (2009); (13)
Herrero-Perezrul et al. (2010); (14) Caso et al. (1996); (15) Celaya-Hernández et al. (2008); (16)
Vazquez-Bader et al. (2005); (17) Espino-Barr et al. (1996); (18) Herrera-Escalante et al. (2005); (19)
Benitez-Villalobos et al. (2008); (20) Benitez-Villalobos et al. (2009); (21) Tapia-Vasquez et al.
(1996); (22) Fajardo-Leon et al. (2008); (23) Herrero-Perezrul et al. (1999); (24) Reyes-Bonilla et al.
(2008); (25) Herrero-Perezrul and Reyes-Bonilla (2008); (26) Skarbnik-López et al. (2010)
2.1 Oceanography and Marine Environments in Mexico
Mexico is the twelfth largest country in the world in terms of extent of coastline and
marine surface area. It has an exclusive economic zone of 314,992,000 ha, a con-
tinental platform of 39,460,300 ha and a coastline of 11,500 km. 1,600,000 ha are
estuarine areas and about 1,250,000 ha are coastal lagoons. Approximately
6,000 km2 of islands are found in the Pacific Ocean (including the remote Guadalupe
Island and the Revillagigedo Islands), Gulf of Mexico, Caribbean, and Gulf of
California (Fig. 2.1), which explains the country’s extraordinary biodiversity in
terms of coastal and marine resources and ecosystems (wetlands, mangrove forests,
barrier islands, dunes, coral reefs, sea grass meadows and nearshore islands) that are
distributed along these four seas (INEGI 2007).
2 The Echinoderms of Mexico 13
The Pacific coast of Mexico extends from 14° to 33°N, and from 92° to 117°W.
This includes the Baja California Peninsula, the second largest in the world (about
1,600 km in length), numerous islands and the eastern coast of the Gulf of Cali-
fornia; the Gulf of Tehuantepec, making a total coast line well above 8,000 km in
length (Fig. 2.1).
The Gulf of California (GC) has several oceanographic features that make it
unique among semi enclosed seas of similar latitude and dimensions, the most
important being strong tidal mixing, some of it close to the deep stratified oxygen
concentration zone. The GC is 1,400 km long and its width in the inner region is
150–200 km, with an area of 177,000 km2, a mean depth of 818 m, and a volume
of 145,000 km3. The GC is an evaporative basin, but in general it gains heat
through the surface. Lacking a sill at the point of connection with the Pacific
Ocean, the GC is constantly affected by a wide spectrum of signals coming from
the Pacific Ocean, including tides and El Niño, which is the most important
interannual anomaly (Lavin and Marinone 2003) (Fig. 2.1).
The Revillagigedo Archipelago or Revillagigedo Islands are a group of four
volcanic islands off the Mexican Pacific coast, known for their unique ecosystem.
They have been part of the Mexican State of Colima since 1861, but are under
Mexican federal jurisdiction. They lie 386 km southwest of Cabo San Lucas, the
southern tip of Baja California and between 720 and 970 km west of Manzanillo.
They are located around 18°490 N and 112°460 W. The Revillagigedo Islands are
home to many endemic plant and animal species and often are called Mexico’s
‘‘little Galapagos’’. They are recognized as a distinct terrestrial ecoregion, part of
the Neotropic ecozone. Socorro Island is the largest and has the most diverse flora,
fauna, and topography (Fig. 2.1).
The Gulf of Mexico (GM) consists of several ecological and geological prov-
inces, such as the continental shelf, the continental slope, and the abyssal plain.
The coastal zone consists of tidal marshes, sandy beaches, mangrove-covered
areas, and many bays, estuaries, and lagoons. The continental shelf forms an
almost continuous terrace around the margin of the GM. Its width varies from a
maximum of more than 320 km to a minimum of about 40 km. Off the coast of
Campeche and the Yucatan Peninsula, the continental shelf consists of a broad
area composed primarily of carbonate material. Off the states of Campeche and
Yucatan, the bank of Campeche extends from 198 and 238 N, and 898 and 938W in
the southern GM (Gore 1992). It is delimited in the north by the Campeche slope;
in the south by the shoreline of the states of Campeche, Yucatan and Quintana
Roo; in the west by the Campeche canyon; and in the east by the Yucatan strait.
This area is an almost flat carbonate plateau that has an average depth of about
51 m (Fig. 2.1).
The remainder of the GM shelf consists of sand, silt, and clay sediments. On the
shelf and the slope that dips downward to the abyssal plain, buried salt domes
occur at various depths. The abyssal plain, which forms the floor of the gulf,
consists of a large triangular area near the centre, bounded by abrupt fault scarps
toward Florida and the Yucatan Peninsula and by more gentle slopes to the north
and west. The basin is very flat, having a gradient of only about 0.3 m per
2,440 m. The deepest point in the Mexico Basin (Sigsbee Deep), is 5,203 m.
The Terminos Lagoon, the largest lagoon-estuarine ecosystem in Mexico, is
located in the southern GM. Terminos Lagoon is a large (*2,500 km2), shallow
(mean depth 3.5 m) coastal lagoon bordering the southern Gulf of Mexico in
Campeche, with 200,108 ha of open water including associated lagoons and
channels, surrounded by about 259,000 ha of mangroves and cattail marshes. The
lagoon borders two geological provinces: to the east the Yucatan Peninsula (low
rainfall, calcareous soils, and no significant surface drainage); to the west and
south, the lowlands of Tabasco and the highlands of Chiapas and Guatemala.
Offshore is the Bay of Campeche, a region supporting one of the largest marine
fisheries in Mexico. The Usumacinta-Grijalva river systems discharge into the
Gulf by three main rivers: the Candelaria, the Chumpan, and the Palizada. The
lagoon is surrounded by shores and is fully incorporated into a National Flora and
Fauna Reserve that comprises 705,016 ha of open water and associated wetlands
and upland (Yañez-Arancibia et al. 1983) (Fig. 2.1).
The Mexican Caribbean and the Yucatan Peninsula is a northeastern projection
of Central America, lying between the GM to the west and north and the Caribbean
Sea to the east, encompassing some 197,600 km2. It includes the Mexican states of
Campeche, Quintana Roo, and Yucatan and, to the south, large parts of Belize and
Guatemala. The peninsula has a mean width of about 320 km and a coastline of
about 1,100 km (De la Lanza-Espino 1991). The Yucatan Peninsula subsurface
Caribbean water (16–20 °C, 36.1–36.5 %) upwells along the eastern slope of the
Yucatan shelf. This water, which originates at depths of about 220–250 m in the
Yucatan Channel, rises at about 10 cm s-1 into the euphotic zone and only
occasionally breaks the ocean surface. This upwelling follows a seasonal cycle.
During spring and summer the upwelled water intrudes over the Yucatan shelf to
create a two-layered water column. Yucatan is one of the most important
upwelling regions on a western oceanic margin (Merino 1997) (Fig. 2.1).
2.2 Echinoderm Fauna of Mexico: An Inventory and Analysis
2.2.1 Historical Review
Studies on Mexican echinoderms started in the nineteenth century, when some

specimens were collected and analyzed by Louis Agassiz in 1841. He reported the
presence of the sand dollar Mellita quinquiesperforata (as Mellita hexapora) in the
Gulf of Mexico (Veracruz). Later, Verrill (1870, 1871) published on some spec-
imens collected by Yale University in Mexican waters. The Blake expeditions to
the Gulf of Mexico and Caribbean (1878 and 1884) produced several papers, such
as Perrier (1881) on Asteroidea, Lyman (1883) on Ophiuroidea, Théel (1886) on
Holothuroidea and Agassiz (1878–1879, 1888) on Echinodea. The Challenger
expeditions in the Mexican Pacific produced papers by Lyman (1879, 1882), Théel
(1879), Agassiz (1881a, b), Carpenter (1884) and Sladen (1889). The Albatross
expedition in 1884 to the Mexican Caribbean resulted in the reports of Rathbun
(1885) for some sea urchins. The Mexican Pacific expedition in 1891 resulted in
reports by Ludwig (1893) and Hartlaub (1895).
Few expeditions have been made during the twentieth century. Several non-
Mexican authors wrote different reports that included echinoderms from Mexican
waters. The reports of Ludwig (1905) and Clark (1917, 1920a, b) were based on
material collected by the last Albatross expedition from 1899 to 1905. The
expeditions to the East Pacific made by the Velero III under the direction of
Captain Allan Hancock during 1931–1941 resulted in the description of Mexican
echinoderms by Clark (1948), Ziesenhenne (1940, 1942), Deichmann (1941, 1958)
and Domantay (1953, 1961). The Templeton Crocker Expedition on board of the
Zaca took place in the Gulf of California in 1936, Deichmann (1937) and
Ziesenhenne (1937) described the holothuroids and echinoids of that area. In 1937
and 1938, again on board of the Zaca, the Eastern Pacific Expeditions of the New
York Zoological Society resulted in publications by Deichmann (1938) and Clark
(1940). In 1940, Steinbeck and Ricketts made an expedition to the Gulf of
California on board the Western Flyer and cataloged more than 550 species of
animals, including echinoderms (Steinbeck and Ricketts 1941). Fisher (1906),
Clark (1913, 1923a, b, 1933, 1916, 1918, 1954), Boone (1926) and Deichmann
(1930) published on Mexican echinoderms from different collections.
During the second half and the end of the twentieth century, several authors
continued to describe new species or published checklists of the echinoderms of
Mexico, such as Clark (1940), Deichmann (1941, 1954, 1963), Fell (1962),
Thomas (1962), Parker (1963), Schroeder (1964), Phelan (1970), Downey (1972,
1973), Brusca (1973, 1980), Litvinova (1975), Lessios et al. (1984a, b), Hendler
and Turner (1987), Hendler and Peck (1988), Maluf (1988), Kerstitch (1989),
Solís-Marín et al. 1993), Hendler et al. (1995), Cutress (1996), Hendler (1996),
Roux and Pawson (1999), Sagarin et al. (2008) and Pawson et al. (2009).
Massin and Hendrickx (2010) described a new species of the genus Synallactes
(S. virgulasolida) collected during a deep-water benthic fauna survey off the
Pacific coast of Mexico in the East Pacific, on board the R/V El Puma. This was
the first record of a Synallactes in the Gulf of California.
Massin and Hendrickx (2011) studied the deep-water holothurians
(377–2,200 m) collected during the research cruises aboard the R/V ‘‘El Puma’’
from the Mexican Pacific and Gulf of California, recording 13 species, two of which
were new to science (Ypsilocucumis californiae and Mitsukuriella unusordo).
The small number of specimens collected during this survey compared to other
areas of the world could be linked to the limiting effect of the Pacific Mexico
Oxygen Minimum Zone.
Honey-Escandón et al. (2011) described a new of species of holothuroid
(Holothuria carere) from the shallow waters of the Mexican Pacific.
Mexican expeditions did not take place until 1939 and added more species to
the list of echinoderms. The Universidad Nacional Autónoma de México (UNAM)
Fig. 2.2 Maria Elena Caso

Muñoz (December 1915–
October 1991)
started the first attempt to organize and classify mainly the Mexican echinoderm
fauna into a comprehensive scientific collection. Dr. Maria Elena Caso began a
series of taxonomic studies that lasted for more than 50 years and that resulted in
more than 50 papers.
Maria Elena Caso Muñoz (Fig. 2.2) was born in Mexico City (December 18,
1915). She was the daughter of Antonio Caso (a noted Mexican philosopher
and rector of the former Universidad Nacional de Mexico, now the National
Autonomous University of Mexico) and Josefina Muñoz. Maria Elena childhood
was in a warm and loving family environment. The end of the Spanish Civil War,
with Franco’s victory in 1939, sparked the start of one of the most dramatic
exoduses in contemporary history. Exiled scientists carried out tremendously
important work in the countries offering them asylum, helping to promote scien-
tific and technological progress. Enrique Rioja Lo-Bianco arrived to Mexico in
1939 to work at the Instituto de Biologia, Universidad Nacional de Mexico
(UNAM). There, Ma. Elena Caso fell under the tutelage of Enrique Rioja and
immediately started the systematic study of the Mexican echinoderms by col-
lecting specimens from Mexican waters, initiating the Mexican Echinoderm
National Collection (MENC).
She studied in the Faculty of Sciences at the UNAM (Mexico City) (1937–1940)
where she obtained her Masters degree in 1943. In 1961 she received the degree of
Doctor of Philosophy at the same University, presenting a thesis on the present
knowledge of the Mexican echinoderm fauna. In the course of her career she
published 56 papers on echinoderms and one book honoring Rioja’s scientific
endeavor. She described eleven new taxa (five species, one subfamily, one subgenus
and two varieties). It was Ma. Elena Caso’s intention to provide for
future researches reference works that would eliminate the need to study the older
literature. Consequently, her work (all exclusively written in Spanish) contains a
staggering amount of details pictured in highly illustrated volumes. After her death
(October 23, 1991) some of her papers were still in press. They were published
postmortem (Caso 1992, 1994a, b, 1996a, b, c, d). Her ideas on the taxonomy of
echinoderms were often controversial but were always respected. Maria Elena Caso
will always be regarded as the pivotal figure in the systematic study of echinoderms
in Mexico.
Currently, the Mexican Echinoderm National Collection (MENC) that Dr. Caso
started holds more than 74,000 specimens belonging to *620 species distributed
along Mexican coasts. The MENC also holds specimens from different parts of the
world. Recently, the collection has entered a rapid growth phase both in terms of
numbers and of geographic and taxonomic coverage. Since 1984, yearly expedi-
tions of Mexican research vessels ‘‘Puma’’ in the Pacific Ocean and ‘‘Justo Sierra’’
in the Atlantic, and since 1992, twice-yearly expeditions to both shallow and deep
sea areas in Mexican waters, have added numerous taxa previously unrepresented
in the collection. Important collections previously minor at MENC, such as frozen
tissues, have grown quickly, establishing MENC as an important national repos-
itory. The MENC also possesses an excellent echinoderm library, with more than
5,100 reprints, books, proceedings, dissertations, etc., covering a great variety of
topics from Aristotle to the present. Such records have been much more complete
with generous requests from people around the world since 1939.
Knowledge of Mexican echinoderms continued to increase in the MENC at the end
of the twentieth century and the beginning of the twenty-first century, with new curators
and several students. The papers published include description of new species,
checklists, taxonomic and biogeographic studies as well as morphological and
molecular phylogenies. Some examples are Buitron and Solis-Marin (1993),
Reyes-Bonilla (1995), Solís-Marín et al. (1997a, b, 2003, 2005, 2007a, b, 2009), Cintra-
Buenrostro et al. (1998), Bravo-Tzompantzi et al. (1999), Godinez-Dominguez and
Gonzalez-Sanson (1999), Solis-Marin and Laguarda-Figueras (1999, 2008, 2010a, b),
Barbosa-Ledesma et al. (2000), Laguarda-Figueras et al. (2001, 2002, 2004, 2005a, b,
2009), Duran-Gonzalez et al. (2005), Fuentes-Farias et al. (2005), Luna and Gonzalez-
Vallejo (2006), Hernandez-Herrejon et al. (2008, 2010), Honey-Escandón et al. (2008),
Solis-Marin (2008), Torres-Martinez et al. (2008), Laguarda-Figueras and Solis-Marin
(2009) and Martinez-Melo and Solis-Marin (2010).
The number of thesis or dissertations on the taxonomy of echinoderms
increased during this period, with the highest number between 2001–2010:
Orbe-Mendoza (1971), Gamboa-Contreras (1978), Worbis-Torres (1986),
Olivares-Gonzalez (1986), San Juan-Ruiz (1988), Herrero-Perezrul (1990),
(Sánchez-Domínguez 1993), Anzo-Martinez (1994), Bravo-Tzompantzi (1996),
Castañeda-Sarabia (1996), Hernandez-Pliego (1998), Solis-Marin (1998),
Cortes-Fernandez (1999), Benitez-Villalobos (2000), Gomez-Carriedo (2001),
Frontana-Uribe (2002, 2005), Celaya-Hernandez (2006), Arriaga-Ochoa (2007,
2010), Dominguez-Castanedo (2007), Hernandez-Herrejon (2007, 2010),
Fig. 2.3 Echinoderm

composition (number of
species per class) in Mexican
waters
Gonzalez-Azcarraga (2009), Bribiesca-Contreras (2011), Estrada-Rodriguez

(2011), Hernandez-Diaz (2011), Pineda-Enriquez (2011) and Zarate-Montes
(2011).
2.2.2 Biodiversity and Distribution of the Mexican Echinoderms
Mexico’s extensive littoral zone and great diversity of habitats supports 643
species of echinoderms (see Appendix), approximately 10 % of the world total.
Class Crinoidea is less well represented with 29 species (4 % of the world total),
while the Class Ophiuroidea is the richest of all with 197 species (31 %); Class
Asteroidea is second with 185 species (29 %); Class Echinoidea is the third most
diverse group with 119 species (19 %); Class Holothuroidea is represented by 113
species (17 %) (Fig. 2.3). Diversity and distribution of echinoderms in Mexico is
described for the four coastal regions of this country: Gulf of California, Pacific
Ocean, Gulf of Mexico and Caribbean Sea (Fig. 2.4).
2.2.3 Gulf of California
Although the Gulf of California is a diverse zone for some classes of echinoderms,
there are no records of crinoids in this area. The asteroids in Mexico are well
represented in all the coasts. In the Gulf of California there is a vast variety and
abundance of specific generic types of asteroids higher than the Caribbean Sea.
There are 58 species. Twenty nine occur in deep waters ([200 m depth). The most
conspicuous genera are: Astropecten, Luidia, Nidorellia, Oreaster, Pharia,
Phataria and Heliaster (Solis-Marin et al. 1993) (Fig. 2.5).
Seventy one ophiuroid species occur in the Gulf of California in two orders,
14 families and 34 genera. Ophiactis savignyi has the widest distribution in shallow
Fig. 2.4 Number of

echinoderm species per class
in each region of Mexico.
C Caribbean, GM Gulf of
Mexico, P Pacific Ocean and
GC Gulf of California
waters. Ophiothrix spiculata is the most abundant and frequent species in the deep
waters (Fig. 2.6).
The Gulf of California is the second area in terms of diversity of echinoids, with
a total of 39 species (Solis-Marin et al. 2005), 12 of them in deep water. The most
characteristic are: Hesperocidaris asteriscus, H. perplexa, Arbacia incisa, Astro-
pyga pulvinata, Centrostephanus coronatus, Echinometra vanbrunti, Lovenia
cordiformis, Encope wetmorei, E. micropora, Clypeaster europacificus, Agassizia
scrobiculata and Mellita grantii (Figs. 2.6 and 2.7).
This region holds the greatest diversity of the sea cucumbers in Mexico, with 64
species. Most (48 species) have wider distributions; they occur from the Gulf of
California to Central America or Galapagos Islands. Only two have a northern
distribution, from Alaska to the Gulf of California. The majority of the species
have shallow bathymetric distributions. Only 20 of the 58 species were collected
from depths greater than 200 m. The most characteristic species of sea cucumbers
are Holothuria (Selenkothuria) lubrica, H. (Thymiosycia) arenicola,
H. (T.) impatiens, H. (Halodeima) inornata, H. (Stauropora) fuscocinerea,
H. (Platyperona) difficilis, Isostichopus fuscus, Neothyone gibbosa and N. gibber
(Solis-Marin et al. 2005, 2009) (Fig. 2.8).
2.2.4 Mexican Pacific Ocean
On the Pacific coast, the crinoids are represented by four species (13.7 % of
the total for Mexico) off the western coast of the Baja California peninsula, and in
the polymetallic nodule fields near the Revillagigedo archipelago and the Clarion
Fig. 2.5 a Davidaster sp., Mexican Caribbean. b Nidorellia armata (Gray, 1840), Gulf of
California. c Heliaster kubiniji Xantus, 1860, Gulf of California. d Amphiaster insignis Verrill,
1868, Gulf of California. e Linckia columbiae Gray, 1840, Gulf of California. f Astropecten
armatus Gray, 1840, Gulf of California. Images a by Valeria Mas, b and e by Carlos Sanchez-
Ortiz, c, d and f by Israel Sanchez
fracture zone. These species, one stalked crinoid and three comatulids, are
well adapted to thrive in extreme depths. The stalked crinoid is known from 3,000
to 4,500 m, while the comatulids occur from 12 to 3,234 m. The hyocrinid species
Hyocrinus foelli Roux and Pawson, 1999, has been reported only near the
Revillagigedo archipelago.
Fig. 2.6 a Ophiothrix galapagensis Lütken and Mortensen, 1899, central Gulf of California.
b Astrodictyum panamense (Verrill, 1867), central Gulf of California. c Ophiothela mirabilis
Verrill, 1867, Guerrero, Mexican Pacific. d Ophiothrix rudis Lyman, 1874, Guerrero, Mexican
Pacific. e Echinometra viridis A. Agassiz, 1863, Veracruz, Gulf of Mexico. f Echinometra
vanbrunti A. Agassiz, 1863, Jalisco, Mexican Pacific. Images a and b by Carlos Sanchez-Ortiz,
c and d by Valeria Mas, e by Fred Boulag and f by Magali Honey-Escandón
The Class Asteroidea off the Mexican Pacific coast is of great interest, not only
because of the high number of genera characteristic of this area, but also for the
close relationships that the endemic species have with species in the Indo-Pacific,
the Mediterranean and the Caribbean (Laguarda-Figueras et al. 2004). There are 59
species in total; 68 % (41 species) are shared with the Gulf of Baja California, 47
Fig. 2.7 a Tripneustes depressus A. Agassiz, 1863, Jalisco, Mexican Pacific. b Toxopneustes roseus
(A. Agassiz, 1863), Jalisco, Mexican Pacific. c Astropyga pulvinata (Lamarck, 1816), Guerrero,
Mexican Pacific. d Holothuria pluricuriosa Jaeger, 1833, Gulf of California. e Isostichopus fuscus
Ludwig, 1874, central Gulf of California. f Cucumaria flamma Solis-Marin and Laguarda-Figueras,
1999, Jalisco, Mexican Pacific. Images a by Magali Honey-Escandón, b by Francisco Solis, c by
Valeria Mas, d and e by Carlos Sanchez-Ortiz, f by Mauricio Valdes
are distributed in shallow waters and 12 occur only in the West coast of Baja
California Peninsula (Fig. 2.4). Only 10 species are found also in the Revilla-
gigedo Islands (Honey-Honey-Escandón et al. 2008). Conspicuous genera of
asteroids include: Astropecten, Luidia, Nidorellia, Pharia, Phataria, Heliaster, and
Henricia (Solis-Marin et al. 1993) (Fig. 2.5).
Fig. 2.8 a Euapta godeffroyi (Semper, 1868), central Gulf of California. b Holothuria arenicola
Semper, 1868, Jalisco, Mexican Pacific. c Labidodemas cf. macullochi (Deichmann, 1958),
Jalisco, Mexican Pacific. d Holothuria kefersteini Selenka, 1867, south of Gulf of California.
e Pentamera chierchia (Ludwig, 1887), Guerrero, Mexican Pacific. f Holothuria cf. inhabilis
Selenka, 1867, central Gulf of California. Images a by Offer Keter, b and c by Francisco Solis,
d by Luis Carballo, e by Valeria Mas and f by Carlos Sanchez-Ortiz
The Pacific Ocean has the lowest diversity of ophiuroids of all Mexican waters,
with only 63 species distributed in two orders, 12 families and 28 genera. Only 23
species are restricted to the Mexican Pacific Coast and the rest (40 species) are
common to the Pacific Ocean and the Gulf of California. The order Ophiurida is
the richest from the class Ophiuroidea, with nine families. The most speciose
genus is Amphiodia, with seven species. Ophiactis savignyi is the ophiuroid

species with the widest distribution in the shallow waters of the Pacific Ocean; it is
distributed along the Gulf of California, the Gulf of Mexico, and also in the
Mexican Caribbean. It has been recorded for eight States, the Revillagigedo
Islands, and for the Maria Islands. Also, Ophiocoma aethiops is widely distributed
in the shallow waters of the Pacific Ocean and it has been recorded for seven
States, as well as for the Revillagigedo Islands.
Thirty seven species of echinoids are found in the Mexican Pacific Ocean,
including the west coast of Baja California. These are classified in 25 genera, 14
families and seven orders. Twenty seven species are shared with the Gulf of
California. The most common species are: E. thouarsii, D. mexicanum, C. coronatus,
A. incisa, Echinometra vanbrunti, Clypeaster europacificus, Encope micropora,
E. perspectiva and E. wetmorei (Honey-Escandón et al. 2008) (Figs. 2.7 and 2.8).
The Pacific coast of Mexico is the second in biodiversity for holothuroids, with
53 species (Fig. 2.4). Ten occur only on the west coast of Baja California. Seven
occur also in the Pacific Coast. Of these 53 species, 46 also occur in the Gulf of
California and the range of distribution can extend south to the Galapagos Islands.
The characteristic species are almost the same for the Gulf of California:
Afrocucumis ovulum, Holothuria (Selenkothuria) lubrica, H. (Semperothuria) imi-
tans, H. (Thymiosycia) arenicola, H. (T.) impatiens, H. (Halodeima) inornata, H.
(Mertensiothuria) hilla, H. (Stauropora) fuscocinerea, Isostichopus fuscus, Neo-
thyone gibbosa and N. gibber (Honey-Escandón et al. 2008) (Figs. 2.7, 2.8).
2.2.5 Gulf of Mexico
The Gulf of Mexico is the most diverse area for the Classes Crinoidea, Asteroidea,
Ophiuroidea and Echinoidea. Twenty five species (86 % of the total for Mexico)
of crinoids live in the Gulf of Mexico and most of them occur in Campeche Bank.
This Bank holds 22 of the 25 species present in the entire Gulf; about 75.8 % of
the total species of crinoids in Mexican waters. Most of those crinoids can be
found in both shallow (\200 m), and deep ([200 m) water environments; 17
species (68 % of the species in the Gulf of Mexico) are found in waters of less than
200 m depth, while 23 species inhabit the deepest part of the gulf. The unstalked
or comatulid crinoids represent 76 % (19 species) of the total number of species
present in the Gulf of Mexico, the stalked crinoids comprise 21 % (six species)
and there is one species (4 %) of cyrtocrinid crinoid, Holopus rangii, present on
Campeche Bank (Fig. 2.4).
There are 96 species of sea stars in the Gulf of Mexico. Half (48 species) are
distributed in deep waters ([200 m depth), and 29 species have a wide bathy-
metric range. The most common genera are Luidia, Astropecten, Cheiraster,
Linckia, Pteraster and Echinaster (Duran-Gonzalez et al. 2005).
Seventy nine species of ophiuroids occur in the Gulf of Mexico. Yucatan has
the greatest number of species (46). Ophiactis savignyi is the species most fre-
quently found in the Gulf of Mexico, especially in the State of Veracruz.
The number of species of echinoids in the Gulf of Mexico is 59. Of these, 30
(52 %) are present in deep waters ([200 m depth). The most common shallow
water species with wide bathymetric (0–80 m) distribution are Eucidaris tribu-
loides, Astropyga magnifica, Arbacia punctulata, Lytechinus variegatus carolinus
and Encope michelini. On the outer shelf Clypeaster ravenelii and Brissopsis are
present in 80–190 m. Finally, Plesiodiadema antillarum and Brissopsis atlantica
occur on the slope to 200 m. Geographically, Encope aberrans, Clypeaster
subdepressus and Echinolampas depressa are restricted to Campeche Bank,
Yucatan and Cape Catoche, Quintana Roo (Laguarda-Figueras et al. 2005a).
The Gulf of Mexico has the lowest diversity of sea cucumbers with only 33
species. Two species also occur off the Pacific coast and 20 are distributed also in
the Mexican Caribbean in shallow waters. The rest (15 species) are present in deep
waters of the Gulf of Mexico (Fig. 2.4), more than 200 m deep. The most common
species are: Holothuria (Halodeima) grisea, H. (H.) floridana, H. (Selenkothuria)
glaberrima, H. (Semperothuria) surinamensis and Isostichopus badionotus
(Duran-Gonzalez et al. 2005).
Mexican Caribbean
In the Mexican Caribbean Sea, the crinoid fauna is not as diverse as it is in the
Gulf of Mexico; nonetheless, the diversity of ecosystems (coral reefs, sandy bot-
tom plains, deep slopes) provide habitats for 13 species, that represent 45 % of the
sea lilies and feather stars reported in Mexican waters. In the Caribbean,
the stalked crinoids represent 23 % with three species reported, inhabiting the
Arrowsmith Bank and the Yucatan Channel area; while the comatulids comprise
the major percentage (76 %) with 10 species in the same area. In this area, the
isocrinid stalked crinoid Endoxocrinus parrae is the most frequently found
species, along with the unstalked Davidaster rubiginosus (Fig. 2.5), Comactinia
meridionalis and Crinometra brevipinna. A total of 13 species (84 %) are shared
with the Gulf of Mexico (Fig. 2.4). Stalked crinoids are present in the Bank of
Campeche and the Caribbean at depths that range from 86 to 143 m in the shallow
areas; and from 747 to 1,245 m as the deepest limit. In the same region, the
comatulid crinoids can be found from 2 to 200 m in the shallow range; and from
200 to more than 3,500 m in the deepest areas.
For the Asteroidea, the Mexican Caribbean is least diverse, with only 51
species. Forty two (79 %) of these are shared with the Gulf of Mexico. Of these 51
species, 14 occur in deep waters ([200 m depth) and 29 have a batimetric range at
depths from 0 to more than 200 m. The most common genera are almost the same
as in the Gulf of Mexico: Luidia, Astropecten, Cheiraster and Echinaster
(Laguarda-Figueras et al. 2005b) (Fig. 2.5).
The Mexican Caribbean is the second most species-rich area of ophiuroids with
78 species. The distribution patterns of the species are diverse. The ones with the
widest distribution in shallow waters of the Gulf of Mexico and Mexican Carib-
bean Sea are Ophiolepis elegans, O. impressa, Ophiocoma echinata, Ophioderma
cinereum, Ophiactis savignyi, and Ophiothrix angulata. The deep-sea species with
the widest distribution for both areas are Ophiolepis elegans, O. impressa,
Ophioderma cinereum, and Ophiothrix angulata. Forty nine species are shared
with the Gulf of Mexico.
Echinoids of the Mexican Caribbean coast are less studied than the other areas.
Nevertheless, it is the second area in diversity with 50 species; 39 of them shared
with the Gulf of Mexico. The most characteristic species occur in shallow waters.
These are Encope aberrans, E. michelini, Clypeaster subdepressus, Cassidulus
caribaearum, Diadema antillarum, Echinometra lucunter, E. viridis, Eucidaris
tribuloides and Tripneustes ventricosus (Duran-Gonzalez et al. 2005).
The Mexican Caribbean has 33 species of holothuroids. The great majority of
them (27 species) also occur throughout the Caribbean Sea and the Florida Keys. The
most characteristic species are Holothuria (Halodeima) floridana, H. (H.) grisea,
H. (H.) mexicana, H. (Semperothuria) surinamensis, H. (Thymiosycia) arenicola,
H. (T.) impatiens, H. (T.) thomasi, Isostichopus badionotus, I. macroparentheses and
Euapta lappa (Laguarda-Figueras et al. 2005b).
Endemism
The delimitation of endemism in marine species is difficult due to dispersion of

larvae through oceanic currents. Nevertheless, Mexico possesses a few species that
are endemic to a biogeographic province.
The Gulf of California, due to its unique oceanographic and geologic charac-
teristics, has the majority of endemismic species (5). The echinoid Encope grandis
is reported only for the Gulf of California and is present both along the continental
coast and the islands within the Gulf. Three species of ophiuroids are endemic.
These are Ophiacantha hirta, Amphiophiura oligopora and Amphiura seminuda.
One species of shallow-water holothuroid Athyone glasselli is endemic.
For the North Pacific Ocean of Mexico, two species of crinoids, Fariometra
parvula and Florometra taneri appear as endemic to the Californian province. In the
southern part of Mexico, the sea star Luidia latiradiata (formely Platasterias
latiradiata) seems to be endemic to the region from Chiapas, Mexico to Colombia.
This species possesses archaic characters that can describe it as a ‘‘living fossil’’
(Caso 1945, 1970, 1972a; Fell 1962; Solis-Marin et al. 1993).
The only reported endemic species of ophiuroid in the Gulf of Mexico is
Amphiodia guillermosoberoni from Terminos Lagoon, in Campeche State. This
species was discovered and described by Caso (1979b). The species is more
abundant inhabiting sites were the salinity ranges from 16 to 21 ppm. Amphiodia
guillermosoberoni is the only Mexican ophiuroid species known to inhabit low

salinity sites.
Only one species of sea star is endemic to the Mexican Caribbean, Copidaster
cavernicola. This species, described by Solis-Marin and Laguarda-Figueras
(2010a) is the only known asteroid that inhabits an anchialine-water cave
(‘‘Aerolito’’) in Cozumel Island (Solis-Marin et al. 2007a, Solis-Marin and
Laguarda-Figueras 2008).
2.2.6 Mexican Fossil Echinoderms
The records of fossil Mexican echinoderms have been the subject of numerous
publications since Nyst and Galeotti (1839). Buitron (1968) produced the first
catalog on fossil echinoids of Mexico. Buitron (1990a) published an illustrated
catalog of the echinoids from the Upper Jurassic and Lower Cretaceous of Mexico.
Nieto-Lopez and Garcia-Barrera (2006) reviewed the Cretaceous echinoids of
Mexico. Research on this matter has been carried out in extended areas such as the
northern region (Boese 1910; Boese and Cavins 1927; Cooke 1955; Buitron 1970a,
1974a), the eastern region (Dickerson and Kew 1917; Lambert 1928; Maldonado-
Koerdell 1953) and the northeastern region (Caso 1951b). Studies on the Baja
California fossil echinoderms are mainly by Jordan and Hertlein (1926), Jackson
(1937), Durham (1950, 1961), Wilson and Rocha (1955), and Chase (1956).
The fossil crinoids of Sonora State have been reported by Vachard et al. (2004)
and Buitron et al. (2007a, 2007b, 2008). Strimple (1971) mentioned the crinoids
from Coahuila State. The fossil echinoderms from Tamaulipas State have been
reported by Dumble (1918), Israelsky (1924), Muir (1936) and Caso (1956). The
fossil echinoderms of Veracruz State were studied by Boese (1906), Dickerson and
Kew (1917), Caso (1956), Buitron (1978a) and Buitron and Silva (1979). Buitron
et al. (1987) studied the fossil crinoids of Hidalgo State. Buitron (Buitrón 1973a,
b) and Garcia-Barrera and Pantoja (1991) described the fossil echinoid fauna from
Michoacan, Guerrero and Colima States. Fossil echinoderms from Puebla State
were reported by Alencaster and Buitron (1965), Buitron (1970b), Esquivel-Ma-
cias et al. (2004), Martin-Medrano and Garcia-Barrera (2006), Applegate et al.
(2009), and Martin-Medrano et al. (2009). Buitron (1976) and Garcia-Barrera and
Pantoja (1991) studied the fossil echinoids from the State of Guerrero, and Buitron
(1990b) from Oaxaca State. Lambert (1936), Jackson (1937), Muellerried (1951)
and Buitrón (1974b, 1974c, 1978b) have recorded the fossil echinoids from
Chiapas State.
The fossil echinoderm fauna in Mexico is represented by approximately 248
species. The Eocrinoidea (an extinct class of echinoderms that lived between the
Early Cambrian and Late Silurian periods) is represented by one species (Gogia
granulosa Robison 1965) (Nardin et al. 2009). Seventeen species of fossil
Crinoidea have been recorded for the Mexican Paleozoic and Mesozoic; Class
Asteroidea with only one species reported for the Cretaceous (Pentasteria sp.).
Six species of Ophiuroidea have been recorded from the Mexican Cretaceous
and Paleogene-Neogene (Pliocene) from different localities in the mainland
territory (Ophiactis applegatei, Ophiura sp., aff. Amphiura, aff. Ophiomusium; aff.
Stegophiura and aff. Ophiura). The Echinoidea is the most diverse and widely
distributed echinoderm fossil group in Mexico, with 221 species. One hundred of
these are from the Mesozoic and 121 species are from the Cenozoic. Both groups
are represented in 17 States of Mexico (Baja California, Sonora, Chihuahua,
Coahuila, Nuevo Leon, Tamaulipas, San Luis Potosi, Hidalgo, Veracruz, Jalisco,
Michoacan, Colima, Puebla, Guerrero, Oaxaca, Chiapas and Yucatan States). The
Holothuroidea is represented by two Cretaceous species (Applegate et al. 2009).
The distribution of Mexican echinoderms in the Mesozoic and Cenozoic indicates
that there was a large marine faunal province in eastern North America, which
differs at the species level from the marine faunal province on the west coast of
North America. This implies the existence of some kind of barrier during the Tertiary
that prevented the faunal interchange between these two provinces. Echinoderms
and other invertebrate groups in the region of the Atlantic and Pacific show the
mark of a moderate exchange, superposed on the old pattern of Tertiary provincial
aforementioned.
2.3 Research
2.3.1 Ecology and Reproduction Studies on Mexican Echinoderms
Ecological studies of the Echinodermata in Mexico have contributed to the

knowledge of species richness, diversity and communities of echinoderms in
several zones, mainly in the Gulf of California, southern Pacific and the Gulf of
Mexico (Fig. 2.1).
Gulf of California
Salcedo-Martinez et al. (1988) generated a benthic inventory of echinoderms in

Zihuatanejo, Guerrero. Data for each species were provided on bathymetric dis-
tribution, substrate, and some characteristics of the habitat.
Morgan and Cowles (1996) studied the effects of temperature on the behaviour
and physiology of Phataria unifascialis and its implications for species distribution
in the Gulf of California. They determined whether cool temperature was an
important factor limiting this species distribution in the northern regions of the Gulf.
Pyloric caecae and gonads showed that food limitation or lack of reproductive
potential was a principal limiting factor. They also suggested that low temperatures
of the northern gulf may well be an important factor limiting the northern distribution
of this species in the Gulf.
Caso et al. (1996) performed qualitative and quantitative analyses of the

echinoderm fauna from the littoral of Mazatlan Bay, Sinaloa and identified 25
species. The results showed that P. unifascialis, Ophiocoma alexandri and
Echinometra vanbrunti were conspicuous because of their density and abundance.
Cluster analysis showed the existence of three groups: the first included species
with a limited distribution and low abundance, the second and the third had species
widely distributed on sandy substrates in the bay. The highest diversity and
specific richness index values corresponded to the species located in protected
areas of the Bay, whereas the lowest diversity and specific richness index values
corresponded to the species found in the exposed areas.
Reyes-Bonilla and Calderon-Aguilera (1999) analyzed the abundance, spatial
distribution and consumption rate of Eucidaris thouarsii and Acanthaster planci at
Cabo Pulmo, Gulf of California. The results showed that the action of corallivore
organisms in terms of their population density and consumption rates was not a
key factor determining scleractinian abundance in the reef of Cabo Pulmo.
Holguin-Quiñones et al. (2000) analyzed the spatial and temporal variation of
Asteroidea, Echinoidea and Holothuroidea from shallow water at Loreto Bay, Gulf
of California. The greatest mean abundance by transect and mean densities in order
of importance for Echinoidea were E. vanbrunti, Centrostephanus coronatus,
Tripneustes depressus, E. thouarsii and Diadema mexicanum; and for Asteroidea,
P. unifascialis, Mithrodia bradleyi and A. planci. Asteroidea was the dominant
class with 12 species, followed by Echinoidea.
Reyes-Bonilla et al. (2005) examined and compared the community structure of
asteroids in four regions of the Gulf of California, characterized by the presence
of rocky reefs. They found Loreto Bay had the highest richness and abundance of
asteroids. However, there were no statistically significant interregional differences
among ecological indices and no groups of locations singled out in terms of
species composition. Phataria unifascialis was the dominant species followed by
Pharia pyramidatus. There were statistically significant positive associations
between three pairs of species, indicating competition between these asteroids in
the Gulf of California is not particularly relevant.
Cintra-Buenrostro et al. (2005) determined the correlation between species
richness of shallow water (depth \ 200 m) asteroids and key oceanographic
factors in nine sections of one degree latitude (from 23 to 31°N) of the Gulf of
California. They reported continental shelf area (at 100 and 200 m depth), mean
temperature and range at three depth levels (0, 60 and 120 m), thermocline depth,
surface nutrient concentrations (nitrates, phosphates and silicates), surface
photosynthetic pigment concentration, and productivity. Sea star species were
assigned to different feeding guilds: predators of small mobile invertebrates,
predators of colonial organisms, generalist carnivores, detritivores, and plankti-
vores. The results showed that total species richness and guild species richness had
strong latitudinal attenuation patterns and were higher in the southernmost Gulf.
Nutrients and surface pigments always presented negative relationships with
species richness, indicating that productive environments set limits to the diversity
of sea stars in the study area.
The number of echinoderm species varies along the Gulf of California,

increasing towards tropical latitudes (Cintra-Buenrostro 2001). However it seems
the community structure of echinoderms in the Gulf is relatively homogeneous and
dominated by asteroids (Reyes-Bonilla et al. 2005; Herrero-Perezrul et al. 2008).
However, echinoderm distribution is probably related to habitat type. For instance,
shallow bays favor coral communities and species associated with them.
Holguin-Quiñones et al. (2008) analyzed the spatial and temporal variation of
Echinoidea and Holothuroidea in shallow water at Isla San Jose, Gulf of California.
The most abundant species were T. depresus, E. vanbrunti, P. unifascialis,
C. coronatus, D. mexicanum and E. thouarsii. There were no statistically significant
spatial or temporal differences in diversity, evenness and species richness.
The analysis identified two groups from locations of the eastern and western coasts.
Herrero-Perezrul et al. (2010) analyzed the community structure of conspicuous
species of echinoderms of three islands in the southern Gulf of California:
San Jose, Espiritu Santo and Cerralvo. They identified 23 species. The ecological
indices showed similar values between the islands. However, the species
composition was different. The community structure was stable in the study area.
It seems that habitat type has a strong influence on the distribution and abundance
of echinoderms.
Luna-Salguero and Reyes-Bonilla (2010) compared the community and trophic
structure of rocky bottom asteroids in two regions of the Gulf of California. The
starfish assemblages in both zones were dominated by a single species, Phataria
unifascialis, and the Loreto area had higher values of richness, abundance and
diversity of asteroids than the Ligüi area. In both areas, herbivores predominated
above detritivores and carnivores. Abundance at each trophic level was statisti-
cally significantly higher in Loreto with more trophic groups per transect. These
higher values of richness, abundance and diversity in Loreto are probably a con-
sequence of a higher number of habitats and food resources.
South Pacific
Benitez-Villalobos (2001) characterized the community of echinoderms at two

coral reef communities (La Entrega and Casa Mixteca) in Bahias de Huatulco
region, Oaxaca State. The La Entrega community showed the lowest values of
diversity and evenness and the highest values of dominance, associated with a
more intense human activity. At the same locality Zamorano and Leyte-Morales
(2005a) repeated the analysis of Benitez-Villalobos (2001) and compared the
results obtained by both studies. Species richness and abundances they recorded
were higher than those obtained in 2001. However there were no statistically
significant differences in the diversity calculated by both studies. Ophiuroidea was
the dominant class of echinoderms in 2001, whereas Holothuroidea was dominant
in 2005. The authors attributed this change in dominance to the intense dredging
that had occurred in Santa Cruz Bay. This probably produced an increase in
suspended matter in the water column, supporting the holothuroids, whereas

ophiuroids had been affected by the loss of natural refuges.
Zamorano and Leyte-Morales (2005b) worked in La Entrega and assessed
population densities at two depth ranges (0–6 m and 6–12 m) of four echinoid
species: E. thouarsii, E. vanbrunti, T. roseus and D. mexicanum. Density of the
four populations varied considerably with the highest densities in the deep zone.
Total abundance of sea urchins, including shallow and deep samples, did not vary
significantly over time. However, the abundance per depth range was different
over time for all species except for E. thouarsii. The authors established that the
reef at La Entrega was undergoing an advanced state of erosion that could favor an
increase in sea urchin abundance and therefore an increase in bioerosion activity.
Rios-Jara et al. (2008) described the specific composition of echinoderms from
Isla Isabel National Park, in the central Mexican Pacific, on rocky and sandy
substrates in subtidal areas to 19 m depth. The most important species according to
the biological value index were D. mexicanum, T. roseus, P. unifascialis, Pharia
pyramidatus, Ophicoma alexandri, Holothuria lubrica, Isostichopus fuscus and
E. thouarsii. They established these species showed more affinity to the Gulf of
California than to the central-south Pacific region.
Lopez-Uriarte et al. (2009) described the specific composition and community
structure of the macrobenthic invertebrates from Bahia de Chamela, Jalisco, using
quadrants along transects of 50 m length. Echinoderms showed the highest species
richness, and the abundance and number of species were related to the heterogeneity
of the substrate, which was characterized by stable mixed substrates including
rocks, medium sized sand and scattered coral colonies. These kinds of substrates
provide shelter in their cavities and crevices, and surfaces for the attachment of
sessile species.
Zamorano and Leyte-Morales (2009) described and characterized the com-
munity of echinoderms associated with coral reefs in Zihuatanejo and Acapulco,
Guerrero during the rainy and dry seasons. The authors proposed the mean values
obtained for diversity in the 13 study sites are low compared to the maximum
possible diversity, due to the dominance of the sea urchin D. mexicanum and the
sea star P. unifascialis. These species, T. roseus and Hesperocidaris asteriscus
were the most frequent and predominant species. Compared to other sites of the
Eastern Tropical Pacific, densities were considered low. However, they did not
find statistically significant differences in the densities of D. mexicanum between
localities, depths, and seasons, although in some localities, the sea urchin densities
increased considerably during the rainy season. The results also showed that, in
localities close to Acapulco, Guerrero, a heavily visited touristic place, the
diversity of echinoderms was lower and the abundance of D. mexicanum was
higher.
Gulf of Mexico
In terms of ecology of Echinodermata, the Gulf of Mexico is one of the less

studied zones. Only four studies have been published presenting ecological anal-
yses of the communities of echinoderms in the region (Fig. 2.1).
Caso et al. (1994) carried out qualitative and quantitative studies on the echino-
derm community from the Laguna de Terminos, Campeche. The relative abundance,
geographic and local distribution, type of nutrition and habitat of 12 species of
Echinodermata were analyzed. Lytechinus variegatus, Luidia clathrata, Echinaster
serpentarius, Ophiophragmus würdermanii and Amphiodia guillermosoberoni stood
out because of their relative importance and abundance. The higher diversity and
specific richness index values corresponded to the species located in the areas with
major oceanic influence, and the low diversity and specific richness index values
corresponded to species found in the lagoons.
Celaya-Hernandez et al. (2008) analyzed the diversity, abundance, distribution
and substrate preferences of the regular sea urchins found at the southern region of
the Isla Verde lagoon reef, Veracruz. The substrate types considered in the analysis
were: coral-rocks, rocks, rocks-sand, and sand and Thalassia testudinum beds.
Diadema antillarum, Echinometra lucunter lucunter and E. viridis were mainly
associated with coral-rock, rock and sand substrates, whereas Lytechinus variegatus
and Tripneustes ventricosus were mainly associated with sea grass. The authors
established that the distribution of the sea urchins is dependent upon substrate type
and feeding habits which are intimately linked. The seven species reached their
largest abundances during the rainy season. Jorgensen et al. (2008) recorded the
high population density survival of D. antillarum after a category five hurricane in
the southern Mexican Caribbean.
Vazquez-Bader et al. (2008) described the distribution and relative abundance
of epifaunal echinoderms in the southwestern region of the Gulf of Mexico, based
on 181 collections by the use of an Otter Trawl during different years. This study
recorded 25 species of Asteroidea, 15 species of Ophiuroidea, 14 species of
Echinoidea, four species of Crinoidea, and one species of Holothuroidea. The most
frequently found species was L. clathrata, reported at 68 % of all stations. The
highest seasonal density of echinoderms was recorded in autumn on carbonate
substrate, whereas the highest species richness was found in summer.
Gonzalez-Azcarraga (2009) compared the community structure and morpho-
logical diversity of regular sea urchins in the Gulf of Mexico ‘‘Veracruz Reef
System’’ and the Cozumel reefs in the Mexican Caribbean. They found statistically
significative differences for all indexes among localities and at all depths. Veracruz
Reef System showed the highest values due to their greater substrate heterogeneity
and a high organic material deposition.
2.3.2 Population Ecology Studies
Few population ecology studies focused on particular species of echinoderms have

been carried out in Mexico. This is regrettable, especially in the case of echinoids
and holothurians, considering their importance as key organisms in the structure
and dynamics of benthic communities, and their potential for commercial
exploitation. Important species include the sea urchin D. mexicanum and the sea
cucumbers Isostichopus fuscus and Parastichopus parvimensis.
Echinoids
Espino-Barr et al. (1996) analyzed the density and population structure of

D. mexicanum on the rocky shore of the state of Colima in the Mexican Pacific as
part of a technical assessment for requests of permission for sea urchin fishing
along Colima and Jalisco shores. The results indicated an average density of
17,000 ind ha-1. There was an increase in gonad weight with respect to body
weight from May to August, followed by a decrease, indicating spawning during
those months. They propose the commercial exploitation of the species between
May and September when the gonads reach the greatest weight. Nevertheless they
proposed it be restricted to protect the species.
Herrera-Escalante et al. (2005) evaluated the importance of D. mexicanum as a
bioerosive agent of coral carbonate at Bahias de Huatulco and the relative magnitude
of coral accretion and bioerosion. The results showed that, in general, D. mexicanum
did not exert a significant role on coral reef community structure at the study area,
probably because of their relatively small size and low numbers. Regarding the
balance between bioerosion and carbonate production, they established that coral
accretion exceeded sea urchin erosion at all sites examined, indicating that at Bahias
de Huatulco coral reef communities were actively growing.
Benitez-Villalobos et al. (2008) evaluated the temporal variation of population
densities and bioerosion by D. mexicanum at Isla Montosa, La Entrega, Isla
Cacaluta and San Agustin, Bahias de Huatulco. Mean density of the sea urchins
was relatively constant during the study at all localities and there were significant
differences between La Entrega and the other localities. The calculated amount of
CaCO3 removed by D. mexicanum showed variations between localities. The
lowest rate of bioerosion occurred at San Agustin and the highest at La Entrega.
They compared their results with those obtained by Herrera-Escalante et al. (2005)
and established that the recent urchin population decrease means a reduction in
their removal of coral CaCO3, and this reduction could be beneficial for coral reef
growing and recovery from stress. Nevertheless it could also signify an increase in
algal biomass and competition with corals for light and space.
Benitez-Villalobos et al. (2009) reported that until May 2009, the density of
D. mexicanum at the different reef communities of Bahias de Huatulco remained
relatively constant over time, with the highest density at La Entrega, where up to
aggregations of up to 100 ind m-2 were observed. However, in May 2009, not a
single individual was observed in the entire reef area (4–12 m) between 2.30 and
3.20 m and surrounding rocky bottom. Instead there were a high number of spines
and hundreds of bare tests scattered over the reef, indicating a mass mortality event
in the sea urchin population. Observations made by the authors during further
monitoring suggested disease as the cause (Benitez-Villalobos et al. 2009).
The observed signs were similar to those described during the mass mortality event
of D. antillarum that occurred in the 1980s throughout the geographical range of
this species in the Western Atlantic.
Fodrie et al. (2007) studied the mechanisms that regulate the feedings position and
feeding mode selection of the sand dollar Dendraster excentricus in some Mexican
localities. Zamorano and Leyte-Morales (2009) worked on the association of the
echinoderms with reef formations on the central Mexican Pacific (Guerrero). Torres-
Martinez and Solis-Marin (2010) analyzed sediment utilization and feeding niche
breadth of the echinoid Meoma grandis in the central Mexican Pacific (Guerrero).
Ebert (2010) studied the demographic patterns of Strongylocentrotus purpuratus
along a latitudinal gradient and included one site on the Mexican Pacific territory.
Francisco and Herzka (2010) measured the role of physical and biological
factors regulating the mode of feeding of the sand dollar D. excentricus in a
shallow estuary in Baja California. The percentage of inclined sand dollars was
strongly and positively correlated with tidal level but no relationship was found
with current velocity, density and organic matter content of the water. The prone
position, indicative of deposit feeding, was largely limited to low tidal levels. Sand
dollars were only oriented parallel to the prevailing currents during the strongest
currents of spring tides ([20 cm s-1). The regulation of the feeding mode of sand
dollars in shadow and hydrodynamically complex estuarine systems differs from
the feeding mode found in exposed coastal environments.
Sonnenholzner et al. (2010) evaluated the effect of three tagging methods on the
growth and survival of Strongylocentrotus purpuratus. The tags tetracycline and
two internal tags, Decimal Coded Wire Tags (CWT) and Passive Integrated
Transponder tags (PIT) were used alone or in combination to evaluate growth
rates, survival, tag retention and post-tagging stress. The results suggested that the
tags used did not affect growth or survival of the urchins. Post-tagging stress was
significantly affected by urchin size but not by tagging methods.
Palleiro-Nayar et al. (2011) analyzed spatial variation in Strongylocentrotus
franciscanus recruitment and assessed the impact of substrate availability and
adult sea urchin density on recruitment along the west coast of Baja California.
There were significant differences in recruit density among sites. Substrate
structure was the main factor that explained these differences. Adult densities did
not explain recruitment differences among sites. Temporal analysis showed that
both substrate structure and adult densities were important in explaining recruit
densities at both sites. The absence of a clear spatial pattern suggests other factors
may explain the differences observed in density and recruitment among sites.
Holothuroids
Reproduction is maybe the most studied trait of sea cucumbers in Mexico, espe-
cially for commercial species (Isostichopus fuscus and Parastichopus parvimensis)
(Herrero-Perezrul 1994, 2004; Fajardo-León et al. 1995; Tapia-Vasquez et al.
1996; Herrero-Perezrul et al. 1999). However, some other sea cucumbers (Hol-
othuria pluricuriosa and H. lubrica) have been studied (Herrero-Perezrul 1994;
Skarbnik-Lopez 2006; Skarbnik-Lopez et al. 2010).
Tapia-Vasquez et al. (1996) analyzed the gonadal cycle of P. parvimensis at the
west coast of Baja California. They showed that the gonad develops during spring
and spawning occurs during summer (July and August). They estimated first
maturity at lengths between 25 and 29 cm. The reproduction in this species was
similar at Isla Natividad and Bahia Tortugas, Baja California Sur (Fajardo-Leon
et al. 2008). One interesting trait they described is evisceration prior to gonad
development from August to October. Therefore they proposed a ‘‘reproductive
ban’’ from February to May and another from August to October because the
evisceration process.
Herrero-Perezrul et al. (1999) analyzed the reproduction and growth of I. fuscus
at Isla Espiritu Santo, Baja California Sur. They described an annual reproductive
pattern, with a single spawning period during summer, which was also influenced
by sea surface temperature. The reproductive cycle was monitored histologically.
Five gonadal stages were described according to cell types present in the gonad.
Reyes-Bonilla et al. (2008) evaluated the abundance of I. fuscus in the National
Park Bahia de Loreto, Gulf of California, during the fishing season 2005–2006 at
29 sites or ‘‘banks’’. The average abundance of I. fuscus in the park was 1.41 ±
0.02 individuals per transect, with an average density of 0.0280 ± 0.0004 ind m-2.
The density was slightly higher in the southern banks. The lowest density was at
Isla Montserrat, which is close to the coastline of the Baja California Peninsula
where most of the fishery occurs. They concluded that the low levels of density do
not represent a critical situation for the populations.
Herrero-Perezrul and Reyes-Bonilla (2008) estimated the relative condition of
an exploited population of I. fuscus at Isla Espiritu Santo, Baja California Sur.
Length and weight were monitored each month and the weight-length relationship
and the index of relative condition were calculated. The weight-length relationship
showed that I. fuscus grew allometrically at this site. The index of relative con-
dition exhibited a parabolic relationship with total length, and peaked at 21 cm
length, which is the size of first maturity. They concluded this pattern may indicate
the condition of individuals improves slowly with age up to a certain point and
then decreases gradually. This finding is important because provides possible
evidence of aging. It is the first indication of senescence in holothurians.
Skarbnik-Lopez et al. (2010) described the reproductive biology and growth of
Holothuria (Selenkothuria) lubrica at Bahia de la Paz, Baja California Sur.
According to data of the length-weight relationship they determined this species
grows allometrically at the site. Five gonadal stages were described according to
cell types present in the gonad. The results showed this holothurian spawns
annually during summer. Variations of the gonad index were directly related to
gonad development and sea-surface temperature, with highest values in summer
corresponding with ripe gonads and the highest sea-surface temperature.
Some other studies have been directed to chemical analysis. Bakus (1974)
demonstrated the toxicity to fishes of some holothurian species (Isostichopus
fuscus, Holothuria impatiens, H. imitans, Neothyne gibbosa and Parastichopus
parvimensis) occurring in the Mexican Pacific (Gulf of California). The evidence
supports the hypothesis that toxicity in this species has evolved as a chemical
defense mechanism against predation by fishes.
The small number of publications about ecology of echinoderms in Mexico
provides evidence of unequal and limited development of ecological studies
related to this group (Fig. 2.4). There are few papers analyzing echinoderm
communities in Mexico, and only four species have been studied in terms of
population ecology. Furthermore, basically all the studies have been focused
mainly on the Gulf of California and the Mexican Pacific, leaving the rest of the
shoreline practically unstudied ecologically, except for two projects performed in
the Gulf of Mexico (Fig. 2.1). This is an indicator of the geographical location and
field of influence of the institutions that hold specialists who are dedicated to the
study of ecology of echinoderms; therefore it is important and essential to become
involved with other institutions in order to develop collaborations that make
possible an increase of knowledge of the current status and the dynamics of
populations of echinoderms, in terms of their relationships with their biotic and
abiotic environments, in order to obtain a more comprehensive knowledge of
echinoderm communities in the entire country.
2.3.3 Chemical Studies of Echinoderms in Mexico
There are few studies about the chemistry on Mexican echinoderms. The first was
by Lara-Guadarrame (1985), who determinated the basic biochemical compounds
in the gonads of the sea urchin Strongylocentrotus franciscanus. Encarnacion et al.
(1989) isolated the first marine natural product from a Mexican echinoderm,
‘‘Neothyside A’’ from the sea cucumber Neothyone gibbosa. This tryterpenoid
tetraglycoside was isolated with cromatographic techniques. Another natural
product isolated from N. gibbosa is Neothyoside B, a triterpenoid diglycoside
(Encarnacion et al. 1996).
Encarnacion et al. (1996) wrote a book about the traditional southern California
medicine. They mentioned that Indians used some marine organism as traditional
drugs, including the sea urchin Echinometra vanbrunti and the sea star Astropecten
armatus. Both are still used as traditional medicines as anti-abortion agents, as
agents to remove scars and as a preventive for vaginal bleeding.
Table 2.1 Mexican echinoderms used in the assays of biological activity by Bryan et al. (1996,
1997)
Antibacterial Antifouling Feeding deterrence
B. B. L. C. L.
amphirite neritina rhomboidies variegatus emarginata
Class Asteroidea
Anthenoides piercei H H
Echinaster sp. H nt nt nt
Astropecten H H H H
articulatus
Goniaster tesselatus H H H H H
Linckia nodosa H H
Luidia clathrata H H
Narcissia trigonaria H H H
Oreaster reticulatus H H H
Tamaria halperni H H H
Tethyaster grandis H
Tosia parva H H
(=Pawsonaster
parvus)
Class Holothuroidea
Holothuria H H
lentiginosa
Isostichopus H H
badionotus
Class Ophiuroidea
Astrocyclis caecilia H H
Asteroporpa annulata H H H H
Astrophyton H H H
muricatum
Class Crinoidea
Comactinia H H H
merdionalis
H Activity significative, nt No tested
Bryan et al. (1996) studied the antibacterial and antifouling activity of the
ethanolic extracts of 20 species of echinoderms (11 asteroids, three holothuroids,
three ophiuroids and one crinoid) collected in the Gulf of Mexico. They used
cultures of Deleya marina and Alteromonas luteo-violacea for antibacterial assays,
and larvae of the barnacle Balanus amphirite and the bryozoan Bugula neritina in
the antifouling assays. They found that only three species had significant anti-
bacterial activity and 11 species antifouling activity (Table 2.1). Bryan et al.
(1997) determinated the chemical defense of 19 species of echinoderms
(10 asteroids, three holothuroids, three ophiuroids and crinoid). For this assay they
made pellets with a tissue of every species and used them in feeding assays with
two marine fish Lagodon rhombodies and Cyprinodon variegatus and a crustacean
Libinia emarginata. They found that 15 species deterred feeding by L. rhombodies
and 17 species deterred feeding by C. variegatus. Only two species deterred

feeding L. emarginata (Table 2.1). Juárez-Espinoza (2010) screened antibacterial,
antioxidant and hemolitic activity from ethanolic extracts of D. antillarlum.
2.3.4 Molecular Systematics Studies on Mexican Echinoderms
McCartney et al. (2000) sequenced 630 bp of the cytochrome oxidase I (COI)

mitochondrial gene of seven species of the genus Echinometra (including E.
vanbrunti from the Mexican Pacific) to provide comparable information on the
eastern Pacific and Atlantic species, to estimate dates for cladogenic events. They
found that an earlier split, assumed to have been coincident with the completion of
the Isthmus of Panama, separated the eastern Pacific E. vanbrunti from the Atlantic
common ancestor and concluded gene flow within species is generally high.
Lessios et al. (2001) reconstructed the phylogeny of the sea urching genus
Diadema using mitochondrial DNA sequences from individuals around the world
to determine the efficacy of barriers to gene flow and to ascertain the history of
dispersal and vicariance events that led to speciation. They used D. mexicanum
from the Gulf of California in the analysis. The D. mexicanum specimens grouped
with species from the West Pacific and western and central Atlantic (D. antillarum,
D. savignyi and D. paucispinun). They found that Indo-Pacific populations
apparently maintained genetic contact with Atlantic populations around the
southern tip of Africa for some time after the Isthmus of Panama was completed.
2.3.5 Biogeography Studies
Studies on echinoderm distribution patterns in Mexico are relatively recent. In

addition, they are poorly documented. The first approaches have been made to
understand the affinities between faunal provinces or regions in the Mexican ter-
ritory. One of the earliest works in Mexico was made by Parker (1963). His study
analyzed the zoogeography and ecology of benthic invertebrates in the Gulf of
California, recognizing 11 faunal assemblages based on the characterization of its
environments. He found that, compared with benthic communities elsewhere in the
world, the diversity of shallow-water species in the Gulf of California is striking.
No single species is dominant. Comparisons between the macro-invertebrate
assemblages of the Gulf of California and those of the Gulf of Mexico and other
parts of the world demonstrate that great similarities, generally at the subgeneric
level, exist in similar environments throughout the subtropical and tropical regions
of the world.
Laguarda-Figueras et al. (2002) analyzed the geographical and bathymetric
distribution of Mexican Caribbean asteroids, evaluating the distributional patters
and their affinities. They found that the ‘‘extended-Western Atlantic’’ and ‘‘warm
Fig. 2.9 a Echinometra vanbrunti A. Agassiz, 1863. Test, primary spine and teeth from the
offerings at the great temple of the Aztecs; b Clypeaster speciosus Verrill, 1870. Test from the
offerings at the great temple of the Aztecs
water Eastern Atlantic’’ were the patterns best represented, suggesting that the
asteroid fauna in the Mexican Caribbean includes an important group of tropical
species that have extended towards the north in zones of colder waters like the
northern Gulf of Mexico and east coast of USA. Work by Caballero-Ochoa and
Laguarda-Figueras (2010) used Parsimony Analysis of Endemism (PAE) to ana-
lyze the holothurian faunal affinities between the provinces of the Mexican tropical
Pacific Ocean.
They found that the Cortes Province was the most diverse, including 62
holothurian species. The Panamic Province was not closely related with the
Mexican, Cortes, Californian, Clipperton, Revillagigedo and Islas Marianas
Provinces. Another important work is that of Hernandez-Diaz (2011) who also
used PAE method to solve the relationship between echinoderms of the reefs of
Yucatan. She found a nested pattern of distribution, which suggests they all belong
to the same zoogeographic region with faunal elements from the Gulf of Mexico
and Caribbean Sea.
2.3.6 Ethnobiology of Echinoderms in Mexico
Animal remains have been part of the archaeological discoveries around the world
and are present in different cultures. Ethnobiology is defined as the study of the
past and present interrelationships between human cultures and the animals in their
environment. It includes classification and naming of zoological forms, cultural
knowledge and use of wild and domestic animals (Seixas and Begossi 2001).
In 1978, the Aztec Great Temple ruins were discovered beneath the central
plaza of Mexico City. Over the next few decades, this sacred structure was
excavated and studied, revealing a wealth of information about Aztec religious
life. Five-hundred years ago, this multi-tiered pyramid marked the literal center of
the Aztec universe, where elaborated ceremonies were performed to maintain
cosmic order and sustain the gods. Among the abundant offerings entombed in the
foundations, the seventh constructive stage of the Great Temple held the remains
of complete tests and fragments of three species of echinoids: Echinometra
vanbrunti (Fig. 2.9a), Clypeaster speciosus (Fig. 2.9b), Mellita sp., Meoma
grandis. Echinometra vanbrunti represents the most abundant echinoderm remains
buried beneath the Great Temple (Solis-Marin et al. 2010). Only one arm tip
fragment of the starfish Astropecten sp. has been found, showing that starfish were
also used during the burial ceremonies. The remains are dated between the years
1502–1521 and were found at the deepest excavation level.
Because the Aztecs received tribute of other towns that they had dominated, we
believe these echinoderms came from the tribute payment of some group dominated
by the Mexicas. For example, C. speciosus presently occurs along the northwestern
coast of Mexico. We believe it was transported from that zone. The Mexica Empire at
that time (1502–1521) had conquered part of the national territory (Guerrero,
Oaxaca, Chiapas States) and other coastal areas of the Mexican Pacific. This record is
very important because it indicates they received materials far more to the north than
normally considered (limits of the Michoacan and Guerrero States). Some specimens
found in the offerings probably have they origin from the west coast of North Mexico,
perhaps Mazatlan, Sinaloa. Although the Aztecs did not conquer the groups of these
areas, we believe that the settler groups of ‘‘Zihuatlan’’ province gave tributes to
the Aztecs. Although they never knew the exact limits of their empire, they knew
that it was in the North of Mexico. The Mexicas never defeated the Tarascos, but
they dominated the province of Zihuatlan. There is reason to believe they received
tribute of some group or groups that were more to the north of the country.
Animal offerings in the burials were made by the highest priests. Their rituals
were closed events. In the massive events they were not celebrated with this type
of offerings. The offerings were made to inaugurate and to consecrate new building
structures and possibly to commemorate festivities of the ritual calendar. The
marine animals, including the sea urchins and starfish (and very probably other
classes of echinoderms) were brought alive to the site. Echinmetra vanbrunti found
in the offerings have remains of pedicellariae on the top of the test and some
fragments of the Aristotle’s lantern. The transport was made using emissaries who
ran very long distances and for a time. In average they ran 30 km daily.
2.4 Aquaculture and Fisheries
Echinoderm fisheries in Mexico are based mainly on the sea urchins Strongylo-
centrotus franciscanus and S. purpuratus, the sea cucumbers I. fuscus and
Parastichopus parvimensis, on the Pacific coast of Mexico and in the Gulf of
California. Except for P. parvimensis, the other species were heavily exploited
during the 1990s. It is important to mention that all echinoderm catch is exported
to other countries, mostly Eastern countries. In the same way, other echinoderms
are also captured for aquarium trade (Micael et al. 2009). Mexico is considered
one of the main exporters together with Indonesia, Singapore, Fiji, Sri Lanka,
Philippines and Vanuatu. For instance, the populations of the Atlantic sea star
Oreaster reticulatus, used as souvenir, have declined in the Mexican Caribbean
(Lunn et al. 2008). Also in the Gulf of California there are other sea stars
(P. unifascialis, P. pyramidatus and Pentaceraster cumingi) that are currently
extracted for ornamentation purposes. However, artisanal fishing has no regula-
tions and the limited information available comes from the results of a survey of
the Mexican echinoderm trade. Currently, an estimated 62 stakeholders in Mexico
collect an average of 12,000 sea stars annually for the souvenir industry.
Some studies have focused on the effect of chemicals related to metamorphosis of
commercially important sea urchins. Carpizo-Ituarte et al. (2002) tested the response
of three species of sea urchin (L. pictus, S. purpuratus, and S. fransicanus) to excess
K+ in sea water as artificial inducers of metamorphosis. The results showed that K+
was an effective metamorphic inducer in all three species. The response depended on
concentration and length of exposure. Castellanos-Kotkoff et al. (2004) showed the
effect of thyroxine on metamorphosis and the presence of the nuclear receptor
COUP-TF in the larvae of three species of sea urchins (L. pictus, S. purpuratus,
and S. fransicanus) from the Mexican Pacific.
Salas-Garza et al. (2005) summarize some of the basic studies and main
achievements in the larval development of the red sea urchin S. franciscanus in
Baja California. Spawning of red sea urchins was routinely induced with KCl; eggs
were fertilized using a 100,000 sperm ml-1 solution. KCl proved the most con-
sistent metamorphic inducer, regularly yielding metamorphosis percentages higher
than 90 %. Metamorphosis was considered complete when the functional jaw that
juveniles use for first benthic feeding appeared (as soon as 20 days after induction).
With this method several thousands of red sea urchin juveniles were produced.
Amador-Cano et al. (2006) used artificial inducers to study signal-transduction
pathways involved in metamorphosis of S. purpuratus postlarvae. Participation of
protein kinase C (PKC), G-protein-coupled receptors (GPCRs), and calcium were
investigated during its metamorphosis, showing that the GPCRs may be shared
between the artificial (KCl) and natural (biofilm) inducers.
2.4.1 The Fishery of Sea Urchins in the Baja California

Peninsula, Mexico
The Baja California fishery is based on two species, the red urchin S. franciscanus
and the purple urchin S. purpuratus. Both species are found in the Pacific coast of
North America, including the Baja California peninsula (Ebert and Southon 2003).
The former is one of the largest species in the world, growing to about 200 mm test
diameter, weighing more than 1 kg and can live for more than 100 years. The
purple sea urchin is small, growing to about 70 mm test diameter.
Fig. 2.10 Red and purple sea urchin catch in Baja California, Mexico
The fishery is limited from Coronado Islands to Isla Natividad in Baja

California Sur (Fig. 2.10). The fishery for S. franciscanus began in 1972 to supply
the Japanese market. The fishery grew quickly and is characterized by three peaks
in catch, in 1979 (5,707 tons), 1986 (8,493 tons) and 1989 (5,536 tons). On the
other hand, S. purpuratus has been harvested only since 1993. Although catches
have remained relatively small (Fig. 2.10). Densities of this echinoid have risen in
the last ten years, but inconsistent roe quality has retarded development of the
fishery (Palleiro-Nayar et al. 2008).
Sea urchins are collected by ‘‘hooka’’ or air compressor to depths up to 30 m.
The divers use short-handled rakes to dislodge the sea urchin from the reef.
Because other fishermen are paid on the basis of roe recovery rather than the whole
animal, divers test roe quality as they work. There is no recreational fishery and
illegal harvest has not been estimated. However, is believed to be relatively small
(Palleiro-Nayar 2009). To date there are some management measures, including
reduction in fishing effort (300 divers in 1987, 229 divers in 2010). Catch rates for
S. franciscanus have declined significantly from 309 kg day-1 in 1988 to 87 kg
day-1 in 2002 and to 176 kg day-1 in 2009. The sea urchin roe produced over 1.3
million US dollars in 2009. However, the maximum profits were obtained in 1994
with 10.4 million US dollars (Fig. 2.10).
2.4.2 The Brown Sea Cucumber Fishery (Isostichopus fuscus)

in the Gulf of California, Mexico
The brown sea cucumber I. fuscus is a common inhabitant of coral and rocky
bottoms from the Gulf of California, Mexico to Ecuador (Maluf 1988; Hearn et al.
2005; Solis-Marin et al. 2009). The fishery of this sea cucumber is without a doubt
a most peculiar story. From the beginning, there were no regulations for fishing
activities, and the only information that existed was related to taxonomy (Caso
1961b, 1967a). The fishery started in the late 1980s, mostly in the Gulf of
California, although some trade was done in Jalisco and Oaxaca (Sierra-Rodriguez
1994; Nuño-Hermosillo 2003) (Fig. 2.1). The maximum recorded catch occurred
in 1991, with 1,800 tons but mean annual catch was around 300 tons (Herrero-
Perezrul and Chavez-Ortiz 2005). In 1994, due to overfishing, authorities closed
the fishery and included I. fuscus in the list of protected species (Anonymous
1994). The fishery was closed until 2000. With biological data that had been
generated by then, some regulations were established and the fishery reopened in
Baja California Sur. Stakeholders were required to record biological data (size,
fishing logs, reproduction, etc.) in exchange for the license. Currently the fishery in
Baja California Sur represents a sustainable activity which is been going now for
10 years with no evidence of overfishing (Herrero-Perezrul 2010). The fishery
management plan in Baja California Sur includes a minimum size of 20 cm and
400 g, total allowable quotas, estimations of biomass and population density, and a
reproductive ban from June 1st to October 31.
2.4.3 The warty Sea Cucumber Fishery (Parastichopus

parvimensis) in the Baja California Peninsula, Mexico
The warty sea cucumber P. parvimensis can grow to a length of 60 cm and a width
of 5 cm. Mobility is limited, though individuals can move up to 4 m day-1 while
feeding (Morris et al. 1980). It is commonly found in rocky bottoms from 0 to
60 m (Woodby et al. 2000). However, exploitation takes place mostly at depths no
deeper than 30 m. The distribution along the west coast of Baja California Pen-
insula is from Coronado Islands to Punta Abreojos (Fajardo-Leon and Turrubiates-
Morales 2009).The fishery started in 1989. In general, fishing takes place when the
sea urchin season closes, from March to June. The stock is exploited by the same
fishermen exploiting sea urchins. Until 2004, the fishery was only in Baja Cali-
fornia, but expanded to Baja California Sur in 2004 (Fajardo-Leon and Turrubi-
ates-Morales 2009). The maximum catch of warty sea cucumber in Baja California
was 1992 with 723 tons. However, landings have decreased to around 230 tons in
recent years with around 100 divers (Salgado-Rogel et al. 2009). In Baja California
Sur the maximum catch was in 2008 with 381 tons. The sea cucumber is boiled,
dried and salted before export, while lesser quantities are frozen. In Asia, sea
cucumber are claimed to have a variety of beneficial health enhancing properties
like lowering high blood pressure, aiding proper digestive function, and others
(Rogers-Bennett and Ono 2006).
2.5 Echinoderm Threats in Mexico
The marine biodiversity of Mexico is threatened by various human activities.

The main factors are pollution and fishing activities.
2.5.1 Over Exploitation
Of the five classes of echinoderms, sea urchins and sea cucumbers are most
heavily exploited. The sea cucumbers (bêche-de-mer or trèpang) that are fished
commercially include 42 species from around the world. They are consumed for
their high-protein content of the body wall, which is boiled and dried (Purcell et al.
2010). During the 1990s, the number of producing countries increased worldwide
with established fisheries in Canada, USA, Mexico and Ecuador. Despite efforts to
regulate the sea cucumber fisheries in Mexico, illegal fishing continues. One of the
greatest dangers to unregulated fishing of this resource is potential population
collapse and therefore the risk of species extinction (Pech 2010). This situation is
not new. In the late 80’s began in the Gulf of California: I. fuscus catches exceeded
one thousand tons in 1991 (Reyes-Bonilla et al. 2008). However, only 5 years later
the fishery authorities declared it endangered and prohibited its capture. This
species is currently on the list of protected species (NOM-059-ecol-2001).
So far, there have been four species of sea cucumbers exploited in the Gulf of
Mexico and the Mexican Caribbean: Holothuria mexicana, H. floridana, Astichopus
multifidus, Isostichopus badionotus (Zetina-Moguel et al. 2003) and two species in
the Mexican Pacific, I. fuscus and P. parvimensis.
Fisheries are also a threat to sea urchins in Mexico. Two species of sea urchins
from the Mexican Pacific have been exploited: S. purpuratus and S. franciscanus.
One of these fisheries occurs in Baja California Mexico, with the exploitation of
S. purpuratus, the purple sea urchin. Gonads are exported to the Northeast Pacific
(Micael et al. 2009). The organisms are collected by hand or with dredges. Since
the quantity and quality of roe cannot be assessed externally, the individuals are
cracked open, causing a high level of mortality and discard of animals.
2.5.2 Trawling and Accidental Catch
The catch depends on the gear used in particular fishery and the amount of effort
applied. One of the major problems in trawl fisheries is the large numbers of other
species that are caught and discarded. Northwestern Mexico, and in particular the
Gulf of California, is the most important fishing area in the country (Robadue
2002). Seventy-seven percent of the volume of fish production in Mexico is
obtained from the Pacific Ocean, and discharged in Baja California, Baja
California Sur, Sonora, Sinaloa and Nayarit States. Most of the catch (80 %) is
from the Gulf of California, where coastal fishing generates 114,000 tons y-1:
31 % in Baja California Sur, 28 % in Sonora, 22 % in Sinaloa and 19 % in Baja
California. In general, 10 % of coastal catches are sharks and dogfish, 17 % bony
fish, 15 % mollusks, 6 % crustaceans and 52 % unidentified (Rodriguez-Valencia
and Cisneros-Mata 2006).
Echinoderms are affected directly by incidental catch when fishing with trawls
because of the significant bycatch effect. The number of species collected this way
is difficult to estimate because they are not recorded in catches. Discards from
trawling are dumped at sea and sink to the bottom where their decomposition
reduces the levels of oxygen and make the habitat less suitable for benthic
organisms. Invertebrates account for up to 90 of the numbers of animals discarded
and up to 73% of these are echinoderms (Pranovi et al. 2001).
2.5.3 Ornamental Species
Many species of echinoderms are popular in aquarium trade. Mexico is included in

the main exporters together with Indonesia, Singapore, Fiji, Sri Lanka, Philippines
and Vanuatu. One Caribbean sea star captured for souvenir industry is Oreaster
reticulatus. Other countries in the Caribbean, including Mexico, allow the
extraction of this species without apparent restrictions.
Little is known about the use of echinoderms as souvenirs or for aquariums. The
limited information available comes from the results of a survey of the Mexican
echinoderm trade. Currently an estimated 62 fisheries in Mexico collect an average
of 12,000 sea stars annually for the souvenir industry (Lunn et al. 2008).
2.5.4 Management and Conservation
Among the possible actions for conservation and management of marine resources,
one of the most important is the recognition of priority sites. Protected areas have
been advocated as one of the most important and effective tools for safeguarding
the world’s biodiversity. A major reason for this is that they protect species from
their greatest threat: habitat loss. In March 2007, Mexico had 61 Marine Protected
Areas with valid establishment decrees, occupying 13,336,390 ha or 58.5 % of its
total federal protected area (*42 % of its total marine exclusive economic zone)
(Ortiz-Lozano et al. 2009). It is noteworthy that most of the known species of
echinoderms are distributed in shallow water areas, and many of them are located
within protected sites.
Until now, no comprehensive work on the population dynamics of the different
exploited species have been made. There is no current information on the number
of possible species that might suggest its economic importance and its distribution
along the Mexican littoral (for example the genus sea star Pisaster). Likewise, we
need to control and regulate the sea cucumbers fisheries in Mexico. For the sea
cucumber I. fuscus, the permit holders have to fish in specific areas called man-
agement unit for wild life or UMA (Unidad de Manejo Ambiental). The exploi-
tation permits are given to organized fishermen who must submit reports on this
activity. UMA controls a harvest quota for each area. It has restricted collection to
sea cucumbers with a minimum 370 g of total weight and has established a closing
season from June to September. On the other hand, there is no official management
plan for P. parvimensis, although the permit holders must collect the sea cucumber
in specific areas and observe a minimum size limit of 200 g body wall (without
gut) (Perez-Plascencia 1995). Assessments of the sea cucumber fishery are based
on annual analyses of fishery-derived information such as catch rate and mean size
of individuals in the landed catch as well as fishery-independent surveys. These are
used to determinate biomass and set a quota of no more 10 % of the total biomass
estimation.
The main life history traits of holothurians suggest that they constitute fragile
stocks. For instance, they are big, slow growing organisms which hardly move,
and therefore easily detected by divers. Population density is less than 1 ind m-2,
similar to values for many of the most important species around the world (Her-
rero-Perezrul et al. 1999; Uthicke et al. 2009). Consequently overfishing is likely
to occur due to their high vulnerability. Conservation efforts have been made,
especially for the fished populations. The permits are based on management
regulations. They limit the number of boats and divers in order to control fishing
effort, and assess the population before and after the fishing season. A private
company has been successful in culturing larvae to produce juvenile I. fuscus
(F.A. Solis-Marin pers. obs.).
The red sea urchin fishery was unregulated until 1987 when a suite of man-
agement reforms transformed the fishery from an open-access competitive model
to one in which individual permit holders (with one or more divers) had exclusive
access to one area. Prior to 1987, divers were able to fish everywhere, with the
result that many areas became severely overfished. In 1994, the Federal Govern-
ment strengthened the holders permit rights by giving them long term concessions
(20 years) for exclusive areas access. The permits are tradable. These changes
were designed to provide investment security and promote greater commitment to
long-term sustainable use. Other management measures introduced in 1987
included the introduction of a minimum legal size of 80 mm test diameter, a
closed season between April and June (later extended to March–June), and a catch
and effort reporting scheme (Palleiro-Nayar 2004).
Assessments of the red sea urchin fishery are based on annual analyses of
fishery-derived information such as catch rate and mean size of individuals in the
catch as well as fishery-independent surveys. Video surveys of reefs also indicate
declines in densities of red sea urchins and increased in density of purple sea
urchins that have replaced red urchins in deeper water (Salgado-Rogel and
Palleiro-Nayar 2008).
In addition, FAO recognizes that some countries have established measures to

reduce bycatch in certain fisheries, through which the ships find it much easier to
avoid capture of unwanted species (Sommer 2005). The pervasive trend of
overfishing, and increasing examples of local extinctions requires immediate
action for conserving stocks, biodiversity, ecosystem functioning and resilience
from other stressors than overfishing (e.g. global warming and ocean acidification)
to ensure the ecological, social and economic benefits of these natural resources
(Toral-Granda et al. 2008).
2.6 Concluding Remarks and Recommendations
The geographic position of Mexico surrounded by two oceans, two gulfs and its
tropical location, explains its extraordinary biodiversity in terms of coastal and
marine resources and ecosystems, making this country one of the most biodiverse
on the planet.
The earliest published article that included work on echinoderms from Mexico
dates back to Louis Agassiz in 1841, who reported the existence of the sand dollar
Mellita hexapora (as M. quinquiesperforata) for the Gulf of Mexico (Veracruz).
Thus began a period of exploration in which most of the species from Mexican
waters were described (Verrill 1870, 1871; Perrier 1881; Lyman 1883; Théel 1886;
Agassiz 1878–1879, 1888). During the twentieth century, few expeditions took
place but several non-Mexican authors wrote reports that included echinoderms
from Mexican marine waters (Ludwig 1905; Clark 1913, 1917, 1920a, b, 1923a, b,
1933, 1948, 1916, 1918, 1954; Boone 1926; Ziesenhenne 1937, 1940, 1942;
Deichmann 1930, 1937, 1951, 1958; Steinbeck and Ricketts 1941; Domantay
1953, 1961). It was not until 1939 when Mexican expeditions began and more
species were added to the developing list of species (Caso 1941, 1943, 1944, 1945,
1946, 1947, 1948a, b, 1949, 1951a, 1953, 1954, 1955, 1958, 1961a, b, 1962a, b,
1963, 1964, 1965, 1967a, b, 1968a, b, c, 1970, 1971a, b, c, 1972a, b, c, 1974a, b, c,
1975, 1976, 1977, 1978a, b, 1979a, b, c). The UNAM began the first attempt to
organize and classify Mexican echinoderm fauna into a comprehensive scientific
collection. The knowledge of the Mexican echinoderms continued to increase in
the Mexican Echinoderm National Collection at the end of the twentieth century
and the beginning of the twenty-first century, with new curators and the incor-
poration of several students. The papers published include description of new
species, checklists, taxonomic and biogeographic studies as well as morphological
and molecular phylogenies (Caso 1979a, b, c, 1980a, b, 1983a, b, 1984, 1986a, b,
1990, 1992; Buitron and Solis-Marin 1993; Caso et al. 1994a, b, 1996; Caso
1996a, b, c, d; Solis-Marin et al. 1997a, b, 2003, 2005, 2007a, 2007, 2009; Cintra-
Buenrostro et al. 1998; Bravo-Tzompantzi et al. 1999, Solis-Marin and Laguarda-
Figueras 1999, 2010a, b; Godinez-Dominguez and Gonzalez-Sanson 1999;
Barbosa-Ledesma et al. 2000; Laguarda-Figueras et al. 2001, 2002, 2004, 2005a,
b, 2009; Duran-Gonzalez et al. 2005; Trujillo-Luna and Gonzalez-Vallejo 2006;
Solis-Marin 2008; Honey-Escandón et al. 2008; Torres-Martinez et al. 2008;

Hernandez-Herrejon et al. 2008, 2010; Laguarda-Figueras and Solis-Marin 2009;
Martinez-Melo and Solis-Marin 2010; Honey-Escandón et al. 2011).
Mexico has 643 species of echinoderms in its territorial seas. The Class Cri-
noidea is the least represented with 29 species (4 %) while the Class Ophiuroidea
is the richest class with 197 species (31 %). The Class Asteroidea is the second
richest class with 185 species (29 %) and the Class Echinoidea is the third most
diverse group with 119 species (19 %) reported in both Pacific and Atlantic
oceans. The Class Holothuroidea has 113 species (17 %).
It is important to make some recommendations: It is necessary to carry out
more ecological studies. The low number and limited scope of publications about
the ecology of echinoderms in Mexico provides evidence of an unequal and
limited development of ecological studies of echinoderms. Few papers exist
concerning analyses of echinoderm communities in Mexico. It is important to
generate more information about their density and other population parameters,
including growth, reproduction, feeding and their relation with the biotic and
abiotic environment. Evaluation must be made of environmental factors that could
influence their presence in a certain location, including studies about their
vulnerability to climatic change. This information will be of great importance, not
only for the scientific knowledge but for verification of ecological theories and a
better description of the autoecologic (individual) and synecologic (community)
relations. It could also establish strategies leading towards the sustainable pro-
duction and administration of economically important echinoderms as a resource
and search for alternatives to increase their production through aquaculture.
The study of sea cucumber aquaculture in Mexico has not begun despite their
high market value and their detritivorous feeding which makes them attractive for
cultivation. Since the body wall is the commercial product from sea cucumbers, it
is necessary to perform studies on the optimization of its production. Considering
that the techniques for holothurian aquaculture are being optimized, particularly in
Asiatic regions, we should take advantage of this technological expansion to set up
culture programs for our own species (i. e. I. fuscus and I. badionotus). This could
also bring environmental profit, since it would diminish the illegal fishery that is
currently observed extensively in Mexican waters.
It is very possible that the list of echinoderms present in Mexican waters is
larger than the one presented here. In particular, the deep sea of the country has
been little studied. Most expeditions covering this area have been made by
institutions of other countries where a great amount of information that is difficult
to access is deposited. Because of this, it is necessary to have available catalogues
of echinoderms with information about their biology, ecology, distribution and
fisheries, in order to improve the access to information and with it the promotion of
studies on echinoderms of Mexico.
As we prepare and train more specialists in echinoderm studies in our country,
it will be possible to amply advance the range of studies that are now required.
Acknowledgments To Dr. Alfredo Laguarda-Figueras for his critical review during the earliest
drafts of this chapter. Students of the Laboratory of Systematics and Ecology of Echinoderms at
UNAM made possible the creation of this chapter, together with the realization of the date extensive
review, figures and plates design, and data base reports: J.I. Alvarez-Rojas, J.A. Arriaga-Ochoa,
G. Bribiesca-Contreras, A.A. Caballero-Ochoa, N. Escandón-Flores, K. Evangelista-García,
P. Garcés-Solchaga, L.A. Hernández-Herrejón, Y.Q. Hernánez-Díaz, P.A. Juárez Espinoza,
C. Martín Cao-Romero, A. Martínez-Melo, T. Pineda-Enríquez, V. Tapia-Ramírez and M. Valdés-
DeAnda. BEBS thanks DGAPA-PAPIIT (UNAM, Mexico) project IN105012, and CONACyT
165826 for funding field trips. FASM thanks DGAPA-PAPIIT IN207011 for funding field trips.
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Chapter 3
Central America Echinoderms: Diversity,
Ecology and Future Perspectives
Juan José Alvarado, Enrique Barraza and Tito Isaac Sancho-Mejías
3.1 Introduction
Central America (CA) is a narrow strip of land that divides the two largest oceans
of the world, which are the Atlantic and the Pacific (Cortés 2007). It include seven
countries: Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica and
Panama. According to the ‘‘Sea Around US Project’’ (www.seaaroundus.org) CA
posseses an Exclusive Economic Zone of 1,518,705 km2 (1,005,922 km2 on the
Pacific and 512,783 km2 on the Caribbean), which is three time larger than its
terrestrial area (521,876 km2). CA also posseses 156,251 km2 of continental shelf
(56,217 km2 on the Pacific and 100,034 km2 on the Caribbean), with 3.7 % of the
coral reefs area of the world (0.4 % on the Pacific and 3.3 % on the Caribbean)
(www.seaaroundus.org), and 6,603 km of coastline (2,856 km on the Caribbean
Sea and 3,747 km on the Pacific Ocean) (www.cia.gov/library/publications/the-
world-factbook).
J. J. Alvarado (&)
Universidad de Costa Rica San Pedro, San José, 11501-2060, Costa Rica
J. J. Alvarado
Posgrado en Ciencias Marinas y Costeras, Universidad Autónoma de Baja California Sur,
La Paz, Mexico
E. Barraza
Gerencia de Vida Silvestre, Ministerio de Medio Ambiente y Recursos Naturales,
San Salvador, El Salvador
e-mail: ebarraza@marn.gob.sveulaliabarr@yahoo.com
T. I. Sancho-Mejías
Museo de Zoología, Escuela de Biología, Universidad de Costa Rica,
San José, Costa Rica
e-mail: sancho.ekin02@gmail.com

68 J. J. Alvarado et al.
The formation of the Isthmus of Panama united South and North America, and
gave rise to the great biological interchange of terrestrial faunae and floras, but
cutting the seaway between the Atlantic and the Pacific (O’Dea et al. 2007). This
process was the result of two geologic processes: plate tectonics and volcanisms
(Coates and Obando 1996). Twenty million years ago (m.y.a.; Miocene), Central
and South America were separated by a wide and deep seaway. This seaway
became narrower due to the movement of the South American plate to the
Northwest until the arc of Panama collided with South America 10 m.y.a. Sub-
sequently, it began to rise and produced the emergence of land around 5 m.y.a.,
causing a final closure of the isthmus between 3.5 and 2.5 m.y.a. (Coates and
Obando 1996; O’Dea et al. 2007).
The Pacific coast is characterized by extensive mangrove systems; sandy bea-
ches, rocky points, cobbles beaches, deltas, and isolated coral reefs of limited
development (Cortés 2007). This coast experiences seasonal upwelling of cold,
nutrient-rich waters from December to April, and produces an increase of the
plankton productivity (D’Croz and Robertson 1997; Fiedler 2002; Amador et al.
2006). This elevated productivity caused by upwelling adds to the high production
from river run-off caused by high precipitation, and increases the nutrients con-
tribution of terrestrial origin (O’Dea et al. 2007). This coast is under the influence
of the Intertropical Convergence Zone (ITCZ), characterized by high irradiance,
variable winds and high precipitation (Amador et al. 2006). The depth of the
mixing layer is shallow (30–40 m) with a permanent thermocline (Longhurst 1998),
and localized near the surface (25 m depth) (Brenes et al. 1990). Productivity is
considered as moderate to high, due to the presence of upwelling areas such as the
Costa Rica Thermal Dome (Fiedler 2002) and seasonal upwelling in the Gulfs of
Tehuantepec, Papagayo and Panama (Bakun et al. 1999). The Pacific coast has six
mayor gulfs: (1) The Gulf of Fonseca (shared by El Salvador, Honduras and
Nicaragua), (2) the Gulf of Papagayo, (3) the Gulf of Nicoya and (4) the Golfo
Dulce (2, 3 and 4 in Costa Rica), and the Gulfs of (5) Chiriquí and (6) Panama
(in Panama) (Cortés 2007). Golfo Dulce is one of four fjord-like gulfs in the tropics
with anoxic conditions at the bottom (Quesada-Alpízar and Cortés 2006).
The Pacific side of CA also has an oceanic island, Cocos Island (Isla del Coco), of
volcanic origin, and located approximately 500 km Southwest of Costa Rica and
630 km Northeast of the Galápagos Islands (Fig. 3.1). Cocos Island is the only
subaereally exposed portion of the aseismic Cocos Ridge. Its volcanic origin is
associated with the Galápagos hot spot (Castillo et al. 1988). This oceanic ridge is
more than 1,000 km long and approximately 200 km wide with depths of 2,000 m
(Harpp et al. 2005; Marcaillou et al. 2006). Moreover, the middle America Trench,
in front of the Pacific coast of CA extends from Mexico to Costa Rica with a maxi-
mum depth of 6,669 km (Astiz et al. 1987). Hydrothermal and cold seeps and oxygen
minimum zones have been reported in the continental shelf (Bohrmann et al. 2002;
Helly and Levin 2004, respectively).
The currents on the Pacific side of CA (Fig. 3.1) are under the influence of the
North Equatorial Countercurrent (NECC), and are responsible for the heat trans-
port from the Tropical Pacific to CA. A portion of the water is first carried out to
3 Central America Echinoderms: Diversity, Ecology and Future Perspectives 69
Fig. 3.1 Central America main currents and distribution of reefs (R), mangroves (M) and
seagrass (S) ecosystems (modified from Lessios et al. 1984; Glynn et al. 1996; Cortés 2007)
the Gulf of Panama and then moved northward by the Costa Rica Coastal Current
along the coast of CA, and finally joins the North Equatorial Current (Wyrtki
1967; Brenes and Gutiérrez 1998; Kessler 2006).
The Caribbean coast of CA is characterized by relatively stable water tem-
peratures, extensive coral reefs and seagrass beds, and large areas of carbonate
platforms (Cortés 2007). This side of CA is considered a passive margin with few
elevated outcrops (Coates 1997). This coast is characterized by lower levels of
nutrients than the Pacific and is less productive, although there are some embay-
ments that have nutrient enrichment due to river run-off (D’Croz and Robertson
1997; O’Dea et al. 2007). The principal currents of the Caribbean Sea go from east
to west. The isthmus deviates some water to the south along Nicaragua, Costa Rica
and Panama (Fig. 3.1). The rest of the water continues to the west towards
Honduras and then north into Belize (Jackson and D’Croz 1997). This coast has two
main Gulfs: (1) the Gulf of Honduras shared by Belize, Guatemala and Honduras
(Harborne et al. 2001; Heyman and Kjerfve 2001) and (2) the Gulf de los Mosquitos
in Panama which includes the semi-enclosed bay of Bocas del Toro (Cortés 2007).
The Caribbean of CA has four carbonate platforms in Belize, Honduras and
Nicaragua (Miskitos), and Panama (Bocas del Toro and San Blas) (Cortés 2007).
Belize has the largest atolls of the Caribbean and one of the largest barrier reefs of
the world (Gibson and Carter 2003). Long stretches of this coast are low-lying and
may be associated with coastal lagoons or canals and swamps (Cortés 2007).
The aim of this chapter is to review and detail echinoderm research in CA, from
Belize to Costa Rica on the Caribbean side, and from Guatemala to Costa Rica on
the Pacific side. The echinoderm research from Panama is reviewed by Coppard
and Alvarado (Chap. 4) due to the greater amount of information from that
country. In order to highlight future needs of new research lines, we present a
review of the main expeditions in the region, and describe ecological and other
studies from both coasts as well as the other research efforts carried out.
3.2 Echinoderm Research
The knowledge of the CA echinoderm fauna is disperse and scarce for some of its
groups. Alvarado and Cortés (2004, 2009) divided the research of echinoderms in
the region into two periods. The first period, from mid-nineteenth century to the
mid-twentieth century, was characterized by expeditions from North America and
Europe, in a period when most of the local species were described. The second
period, from mid-twentieth century to the present, has been characterized by
widespread interest in ecology done by local scientists at research stations in CA.
Echinoderm research in CA started on the Caribbean side in 1884 (although the
first report was from Panama by Verrill (1867) when the US Fish Commission
Steamer Albatross expedition collected the asteroid Sclerasterias tanneri from
Honduran waters. Between 1888 and 1891, the Albatross visited the Pacific side of
CA. Those expeditions, led by Alexander Agassiz, collected mostly from Cocos
Island and Cocos Ridge (Ludwig 1894, 1905; Lütken and Mortensen 1899;
Alvarado 2010). Ludwig (1894, 1905), Agassiz (1898) and Lütken and Mortensen
(1899) described many new species of asteroids, ophiuroids, echinoids and holo-
thurians from the eastern Pacific. The asteroid Pauliella aenigma, the echinoids
Hesperocidaris panamensis and Centrocidaris doederleini, and the holothurian
Psolus diomedeae were species described from specimens collected at Cocos
Island, Costa Rica (Alvarado 2010).
Between 1930s and 1940s, as a result of the expeditions to the eastern Pacific by
the Allan Hancock Foundation and the New York Zoological Society, there was a
significant increase of research on echinoderms (Ziesenhenne 1937, 1940, 1942,
1955; Clark 1939, 1940, 1948; Deichmann 1938, 1941, 1958). The Velero III from
the Allan Hancock Foundation collected 37 species of echinoderms from the
Pacific of CA (35 species from Costa Rica, one species from Nicaragua and one
species from Guatemala), from eight type of species localities (seven from Costa
Rica and one from Nicaragua). The species described from Costa Rica were the
ophuiroid Ophionereis dictyota, the asteroid Tamaria obstipa, the echinioids
Encope cocosi (endemic to Cocos Island) and the holothurians Labidodemas
americanum, Euthynodium veleronis, Psolus diomedae and Pentamera beebei, and
the irregular echinoid Encope laevis from Corinto Port, Nicaragua.
The Zoological Society of New York expeditions aboard the SY Arcturus and
SY Zaca were led by William Beebe. They reported the presence 58 species of
echinoderms (Fisher 1928; Clark 1940). However, Zaca collected a total of 822
specimens: 73 asteroids, 645 ophiuroids and 104 echinoids. This included 57
species: 13 species of asteroids, 29 species of ophiuroids and 16 species of
echinoids. From that material Clark (1940) described two new species of ophiu-
roids: Amphiodia vicina from the Gulf of Nicoya, Costa Rica, and Ophiolepis
grisea from the Gulf of Fonseca (Monypenny Point) and the presence of Echin-
aster (Othilia) aculeata, a species that was thought to be extinct.
Between 1979 and 1981, as part of a study of the megabenthic invertebrates
communities of Gulf of Nicoya on board the R/V Skimmer, nine species of echi-
noderms were found (two species of asteroids: Luidia tessellata and Astropecten
armatus; five species of ophiuroids: Ophiocoma sp., Ophiothrix sp., Ophiopallus
sp., one species each of Ophiocanthidae, Ophiodermatidae and Ophiuridae; and
two species of echinoids: Astropyga pulvinata and Arbacia incisa). All the echi-
noderms together accounted for only 0.5 % of the total number of individuals and
only 1.6 % of the total biomass. They were more abundant in the inner parts of the
gulf (Maurer et al. 1984). Between 1980 and 1981, Maurer and Vargas (1983)
continued with the study of the benthic communities, but reported only four species
of ophiuroids (Amphipholis sp., Amphipholis squamata, Ophiactis savignyi and
Ophiothrix spiculata), which accounted for 0.5 % of the biomass.
Later expeditions at the end of the twentieth century and beginning of the twenty
first centuries were carried out on board the RV Urracá from the Smithsonian
Tropical Research Institution in collaboration with authorities in Costa Rica
(University of Costa Rica) (Cortés 2009) and El Salvador (Ministerio de Medio
Ambiente y Recursos Naturales) (Barraza and Hasbún 2005). The El Salvadorian
vessels, the RV Fengur, collected many new records of echinoderms (Barraza
1995), and the RV Miguel Olivier collected new deep-sea asteroids and ophiuroids
at 900 m in December 2009. The Costa Rican expeditions from the Centro de
Investigaciones en Ciencias del Mar y Limnología collected specimens at Cocos
Island at depths between the years 2005 and 2009. Other Pacific expeditions
that visited and collect echinoderms were the RV Boric (1927), RV Stranger
(1939, 1969), and the RV Te Vega (1968), last one from Stanford University
(Cortés 2009). Most of these expeditions did not publish accounts of their expe-
ditions, but the species collected are recorded in the data base of the National
Museum of Natural History, Smithsonian Institution, Washington, DC. The species
are recently reported by Alvarado et al. (2010).
On the Caribbean side, Devaney (1974) reviewed the first expeditions that
visited Belize. He indicated that the first research ship was the Pawnee I of the
Harry Payne Bingham Institute in April 1925, from which Boone (1928) described
two species of echinoids, three species of asteroids and eight species of crinoids
from Glover’s Reef. From the 1937–1938 Rosaura expedition, John and Clark
(1954) reported four species of asteroids, five species of ophiuroids, two species of
echinoids, and one species of holothuroids. Some of them from depths of more
than 914.4 m (500 fathoms). During the 1959–1960 Cambridge Expedition,
Stoddard (1962) made brief remarks on echinoderms in his comparisons of
ecological zonation (Devaney 1974). These expeditions were complemented with
the extensive taxonomic work carried out at Carrie Bow Cay (Kier 1975; Pawson
1976; Hotchkiss 1982; Macurda 1982).
According to Miller (1984), the asteroid family Ophidiasteridae is represented
by 13 species in the tropical western Atlantic, Caribbean and Gulf of Mexico. For
four species, no new material has been reported from the western Atlantic since the
description of the holotypes. Miller (1984) collected specimens of Copidaster
lymani at Carrie Bow Cay, Belize. Thus, these are the first specimens reported for
western Atlantic since the holotype had been described from Florida. Miller (1984)
also included a key to the western Atlantic species of Ophidiasteridae.
During the 1980s, Gordon Hendler and a number of colleagues made several
studies on Belizean echinoderms diversity and distribution, and included the
descriptions, of at least, five new species of ophiuroids species. Hendler and Miller
(1984) described two new ophiuroid species from the western Atlantic: Ophioderma
devaneyi collected from the outer continental shelf and upper slope off Florida,
and Ophioderma ensiferum collected the reef slope of the Belize Barrier Reef, off
Carrie Bow Cay. Hendler and Littman (1986) examined the size distribution of 33
ophiuroid species from distinct substrata and reef zones located in four zones on the
Belize Barrier Reef. They related size-distribution and population density data to the
mode of reproduction and the biological interactions between the ophiuroids, and
their habitat in order to understand the biology of juvenile individuals and other reef
cryptofauna.
Hendler and Turner (1987) described two new Ophiolepis species from the
Caribbean Sea and the Gulf of Mexico, O. gemma from deep-reef habitats and
O. ailsae from bathyal depths. Ophiolepis gemma was collected at Carrie Bow Cay
located on the seaward side of the Belize Barrier Reef, Barbados, and the Gulf of
Mexico. They found two brooding specimens from Belize that held embryos in
their bursae. They concluded that the occurrence of embryos at the same stage of
development in one specimen suggests synchronous brooding in the individual.
Ophiolepis ailsae was collected at several locations through the Caribbean Sea.
Bursae examination did not reveal indication of brooding. Thus the reproductive
mode of this species is not known. They described specialized hooked plates on
the distal arms in both new species.Moreover, they suggested the hooks are used in
feeding.
Hendler (1988a) described Ophiolepis pawsoni, bringing to seven the number
of Ophiolepis species reported from the western Atlantic. The specimens were
collected on the seaward slope of the Belize Barrier Reef where it occurs simp-
atrically with O. gemma. Hendler (1988a) also included a key to the species of the
genus Ophiolepis that comprises three groups based on different morphology of
the disk, arms, and central rosette.
Hendler and Pawson (2000) made an exhaustive description of the echinoderms
present at several localities in Belize and reported the presence of more than 90
species, mostly from barrier reefs localities (including Carrie Bow, Rhomboidal
and Pelican Cays). They described the environments and gave some information
about the natural history of the species. Most of this information is also present in
the compilation of echinoderms of the Caribbean by Hendler et al. (1995).
Between 1935 and 1978, research vessels visited the Caribbean coast: RV
Leonidas (1935), RV Oregon (1962, 1967), RV Oregon II (1970), RV Alaminos
(1970), RV Pillsbury (1971), RV Pescanica 25 (1974) and RV Alpa Helix (1977).
These expeditions had several collection stations in Honduras, Nicaragua, Costa
Rica and Panama. No scientific accounts of the echinoderms collected were
published. Most of the specimens are deposited at the National Museum of Natural
History, Smithsonian Institution, at Washington, DC. More recently those species
are reported by Alvarado (2011).
From 1867 to 2011, 230 papers related to echinoderms in CA (including
Panama) have been published (Fig. 3.2). Between 1860 and 1970, there were less
than ten publications per decade (Fig. 3.2a). Most of those publications are related
to expedition’s reports and monographs of specific groups. Their contribution was
enormous, setting the foundations for future research, describing the majority of
species that nowadays we know for CA. The last four decades (1970–2011) have
been the most productive in terms of numbers of publications. This period was not
only dedicated to describing new species, but also their relationship with their
habitat, biological processes, reproduction, evolution, and their relationship with
disturbances like the El Niño-Southern Oscillation. In these four decades echi-
noderm research in the region was greatly advanced by (1) the Smithsonian
Tropical Research Institute (STRI) in Panama with their laboratories since 1960s
in Naos, Punta Galeta and Bocas del Toro (Robertson et al. 2009), (2) the Carrie
Bow Cay Marine Field Station that was established in 1972 by the Smithsonian
National Museum of Natural History (Rützler 2009), and (3) the Marine Science
and Limnology Research Center (Centro de Investigaciones en Ciencias del Mar y
Limnología-CIMAR) at the University of Costa Rica that was established in 1979
(Cortés 2009). The research was carried out mostly by Gordon Hendler, Peter W.
Glynn and Harilaos Lessios, and they without any doubt were the pioneers in the
study of echinoderm ecology and evolution in CA and established the base line for
future research. The effect of the presence of these institutions is evident in the
number of publications per country in CA. Panama (92), Belize (49) and Costa
Rica (39) published most of these, with only 13 publications for the remaining
Fig. 3.2 a Number of echinoderm publications per decade since 1860 until 2011 in Central
America, b per country and by c research area
countries (Fig. 3.2b). Forty-four publications are common to all CA countries

resulted from those major early expeditions to the region or reviews. The main
focuses of research have been ecology (94 publications), taxonomy (57 publica-
tions), evolutionary processes (37 publications), diversity (20 publications) and
reproduction (17 publications). Very little research has been done on physiology,
anatomy or fisheries, and none in either conservation issues or aquaculture
(Fig. 3.2c). These indicate future areas of research.
3.2.1 Ecology
Crinoid Ecology
Crinoid research has being carried out only in Belize. There are no reports on the
presence of crinoids on the Pacific side, with the exception of Thalassometra
agassizii and Fariometra parvula found in deeper waters at Cocos Ridge (Alvarado
and Cortés 2009; Alvarado 2010).
Macurda (1982) described the presence of four shallow water crinoids at Carrie
Bow Cay, Belize: Davidaster discoidea, Davidaster rubiginosa, Ctenantedon
kinziei and Analcidometra armata. Both Davidaster species were the most abun-
dant crinoids on the forereef slope, and located on the top of the outer ridge, and in
the spur and groove zone of the inner fore reef. Macurda (1982) argued that this
distribution reflects their need of moderate water motion without direct wave surge.
He described three color varieties for D. discoidea and seven for D. rubiginosa.
The three forms of D. discoidea have black-tipped pinnules, but differ in the color
of the arms (green, yellow and dark green). For D. rubiginosa, he describes only
three forms that are all orange, but with different pinnules colors (black, yellow and
white). The black form is the most common. For these six color forms, he provided
the depth range, and the average arm numbers with their length. Ctenantedon
kinziei was a rare species, occurring only on the top of the outer ridge and down the
fore-reef slope, from depths of 13–25 m, and completely hidden within coral
colonies (mostly Montastraea). Analcidometra armata was associated with the
gorgonians (Pseudopterogorgia spp., Muriceopsis sp. and Gorgonia mariae)
(Macurda 1982).
Humes (2000) described a new parasitic species of siphonostomatoid copepod
(Asterocheres crinoidicola) on Nemaster grandis and D. rubiginosa. This was the
first report of a copepod parasitic on a crinoid in the Caribbean.
Asteroid Ecology
Caribbean
The asteroid Oreaster reticulatus (Fig. 3.3) has been recorded as common on
shallow waters throughout the Caribbean Sea. Increased collection for ornamental
shell and aquarium trade has resulted in a striking decline in Caribbean popula-
tions (See Scheibling and Metaxas 2010 for references). In Belize, Scheibling and
Metaxas (2010) examined the potential role of mangroves and fringing patch reefs
Fig. 3.3 Common Central American Caribbean echinoderms. a Oreaster reticulatus; b Lytechi-
nus variegatus; c Diadema antillarum; d Tripnesustes ventricosus; e Echinometra lucunter;
f Echinometra viridis; g Actinopyga agassizi; h Holothuria mexicana. (Photographs by I.S.
Wehrtmann, C. Fernández and J.J. Alvarado)
as recruitment sites for O. reticulatus. They sampled populations at sites within

these habitat types. They included a population in a seagrass bed and another on an
open sand flat for comparison. The radius of each individual was measured. They
also recorded the coloration pattern to distinguish juveniles from adults. Their
results showed that O. reticulatus density varied among sites. It was lowest at the
seagrass bed and highest at sand bottom, while intermediate densities were
recorded at mangrove and fringing reef sites. They noted that asteroid populations
in mangrove and coral habitats were mainly composed of juveniles, but lowest in
the seagrass beds and intermediate on the sand flat. Based on the abundance of
juveniles on mangrove banks and in fringing reefs, they concluded that these are
important habitats for recruitment and serve as nurseries for O. reticulatus.
Furthermore, they proposed that species such as O. reticulatus may serve as
sentinels of ecological change, and they may have broad-ranging effects in the
health of coastal ecosystems.
Pacific
Guzman and Cortés (1989), as part of a study of the coral reefs at Caño Island,
indicated the presence of the sea star Acanthaster planci. This species was found in
densities between 0.4 and 1.8 ind ha-1, and feeding mainly on the corals Pavona,
Psammocora and Pocillopora. Acanthaster planci was also observed at Cocos
Island, where it had densities of 10.4 ± 3.1 ind ha-1in 1987. It fed on Porites
lobata, a coral that was not part of its diet (Guzman and Cortés 1992). Guzman and
Cortés (2007) reported densities of 7.4 ± 2.3 ind ha-1 in 2002. Alvarado and
Chiriboga (2008) reported a density of 0.02 ind m-2 on only two of 19 study sites
around the island. In other studies the seastar was present in four of five reefs
studied (Guzman and Cortés 1992, 2007).
Ophiuroid Ecology
Caribbean
During the 1990s, Richard B. Aronson evaluated the impact of hurricanes and
storms on predator–prey dynamics in Caribbean coral reefs. Aronson (1991)
assessed predation, physical disturbance and sublethal arm damage in ophiuroids
(Ophioderma spp.) in Belize and Jamaica in order to compare to Jurassic-Present
trends. This study supported the hypothesis that predation on ophiuroids increased
when durophagous teleostean fishes and decapods crustaceans diversified in the
Late Mesozoic.
Aronson (1992) investigated predation intensity experienced by Ophiothrix
oerstedi within and among back reef sites in Jamaica, Barbados and Belize, and
provided assessment of the impact of severe storms on predator–prey interactions
in back reef habitats. Aronson (1998) evaluated the ecology of predation on
ophiuroids in the Caribbean on two temporal scales: seasonally over a 3 years
period and annually over a decade. His research suggests that some ecological
relationships are stable at multiple temporal scales. Aronson’s investigations

concluded that strong hurricanes did not cause detectable damage to the ophiuroids
subpopulations and did not affect predator–prey interactions (Aronson 1991, 1992,
1998).
Hendler (1982) studied the feeding behavior of the basketstar Astrophyton
muricatum at Carrie Bow Cay, Belize. He described the sequence of events in
nocturnal behavior from 18:16 to 05:40 h, by filming an indivdual and measuring
the direction and intensity of currents. Astrophyton muricatum fed during the
periods of moderate water speed associated with ebb and flood currents. It changed
feeding position in response to tidal flow by directing its arms into the current.
Hendler (1984a) examined the mutualistic association between the sponge
Callyspongia vaginalis and its obligate commensal Ophiothrix lineata on Carrie
Bow Cay. The diet of O.lineata consisted of detrital particles adhering to the
sponge which were too large for it to use as food. Thus, the brittlestar cleans
the inhalant surface of the sponge as it feeds, increasing filtration capability of
the sponge. This association allows O. lineata to hide on the sponge from pred-
atory fishes. Hendler (1984a) reports the presence of six species of ophiuroids
(Ophiothrix lineata, O. suensoni, O. angulata, O. oerstedi, Ophiactis savignyi and
O. quinqueradia) associated with the sponges C. vaginalis and Neofibularia
nolitangere, and provides their size frequency (disk diameter) and their location
inside or outside of the sponges. Ophiothrix lineata and O. oerstedi were restricted
to C. vaginalis, while only O. quinqueradia was restricted to N. nolitangere.
Ophiothrix lineata activity outside the sponges begins at dusk and diminishes in
the morning (Hendler 1984a).
Hendler and Peck (1988) compared Belize Barrier Reef ophiuroids collected at
a depth of about 24 m on the fore-reef slope with those collected in shallow waters
(0–15 m depth) by Hendler and Littman (1986). They found 44 species, eight
species were restricted to the shallow reef, and nine species were found only on the
fore-reef. They measured the disk diameter of each individual and grouped the
species according to four modes of reproduction: (1) planktotrophic (with pluteus
larvae), (2) lecitotrophic (with vitellaria larvae), (3) brooders, and (4) fissiparous.
They noted that species richness of brooding species increased with depth, but the
number of fissiparous, planktotrophic and lecitotrophic species did not vary with
depth. Their results showed that species richness increases with depth. They found
differences in the species richness composition, relative species abundance, and
reproductive modes between the ophiuroids occurring on the shallow-reef and the
fore-reef slope.
Hendler et al. (1999) described a previously unreported phenomenon of
planktonic dispersal of juvenile ophiuroids. This observation suggests that juve-
niles reenter the plankton and drift, or perhaps raft on algal fragments, after having
been recruited to the benthos. The water-borne juvenile ophiuroids are compared
to plantigrade and planktonic postlarva stages of other marine invertebrates.
Parasitic crustaceans in ophiruorids from Belize were studied by Maddocks
(1987), Humes and Hendler (1999) and Hendler and Kim (2010). Maddocks
(1987) described the ostracod Pontocypria hendleri as a commensal of the
ophiuroid Ophioblenna antillensis from Carrie Bow and Wee Wee Cays. Humes
and Hendler (1999) described the presence of three copepods (Ophiosyllus latus,
O. reductus and Pseudanthessius deficiens) which are parasitic on four species of
Belizian ophiuroids (Ophioderma cinereum, O. brevispinum, Ophiocoma echinata
and Ophiocomella ophiactoides). Pseudanthessius deficiens live in O. cinereum
and O. brevispinum. The other two species of copepods are host-specific: O. latus–
Ophiocomella ophiactoides and O. reductus–O. echinata. One to four O. latus
occurred on 17–57 % of O. ophiactoides examined. These copepods generally
clasp the first or second arm joint near the edge of the disk. Hendler and Kim
(2010) described the presence of a parasitic metanauplius larva of thaumatipsyllid
copepods (Caribeopsyllus chawayi and Caribeopsyllus sp.) in Ophiothrix angulata
and Ophiactis savignyii in Carrie Bow. This parasitic larva used mandibular chelae
to grasp the host’s stomach wall. The larvae of C. chawayi were found only in
O. savignyi, and the larvae of Caribeopsyllus sp. was found only in O. angulata
living on Halimeda spp. Between 18–27 % of O. angulata were parasitized. Most
of the parasites were small compared to their hosts. Infected O. angulata tended to
be small and were both sexually mature and immature (Hendler and Kim 2010).
Hendler (2005) described two new species of ophiuroids from the Southern
Caribbean of Costa Rica and Panama, Ophiothrix cimar and O. stri were named in
honor of the Centro de Investigaciones en Ciencias del Mar y Limnología
(CIMAR) and the Smithsonian Tropical Research Institute (STRI) respectively.
Individuals or both species are cryptic and were found beneath coral rubble in
shallow water fringing coral reefs. Ophiothrtix stri was found also in the bore holes
of the sea urchin Echinometra lucunter, but the association was not obligatory.
Ophiothrix cimar was relatively more common than O. stri in back reef micro-
habitats, but neither species occurred in nearby lagoonal and mangrove habitats.
Both species are gonochoric and do not brood. Their eggs are of a size indicative
of abbreviated larval development. The gametes of O. stri developed into demersal
embryos that developed within a large, adhesive fertilization envelope. Hendler
(2005) also discussed their systematic affinities, and indicated that O. stri and
O. synoecina could be sister species with a greater systematic affinity to O. lineata
than to O. cimar. He also states that the reefs where the species were previously
observed have deteriorated because of environmental degradation and produced a
reduction of the populations.
Most recent research on Belizean ophiuroids involves genetic connectivity and
phylum diversity on a regional scale of the Caribbean. Richards et al. (2007)
assessed gene flow between Florida and Belize of the broadcast-spawing brittle
star Ophiothrix lineata. They concluded this species showed significant genetic
structure between Florida and Belize.
Pacific
Between 1987 and 1996, a series of studies of different benthic communities were
conducted on the mud flats at Punta Morales, Gulf of Nicoya, Costa Rica (Vargas
1987, 1989). In those studies the sand dollar Mellitella stokesii (as Encope stok-
essi) and the ophiuroid Microphiopholis geminate (as Amphipholis germinata)
appeared as regular members of those communities. This research continued until
1996, including 76 collecting dates from February 1984 until September 1996
(Vargas and Solano 2011). Using sediment cores (265.5 cm3; 14 cores/date),
Vargas and Solano (2011) studied the presence of both species in an area of
400 m2. They found a total of 149 individuals of M. geminata and 63 of M.
stokesii. Both species had an irregular temporal pattern of abundance, with slight
increases at the end of the rainy season or during the dry seasons.
Echinoid Ecology
Caribbean
Strathmann (1981) studied the role of spines in preventing structural damage to

echinoids tests at Carrie Bow Cay and Twin Cay, Belize. He analyzed echinoid
tests by breaking them by way of dropping lead fishing sinkers on living sea
urchins submerged in sea water. He observed specimens from five species rep-
resenting four echinoid orders: Eucidaris tribuloides, Diadema antillarum,
Tripneustes, Lytechinus variegatus and Echinometra viridis (Fig. 3.3). The results
suggest that spines and tissue apparently protect the test apparently by both
spreading the load and absorbing energy. He also noted breaks on podial pore
tracks are common in all the species. The breaks at sutures were common in living
specimens of E. tribuloides. In contrast, the breaks on sutures in D. antillarum,
L. variegatus, T. ventricosus, and E. viridis were often absent and comprised a
small portion when present. Thus, he suggests that tracts of podial pores are sites
of relative weakness. In addition, he evaluated the strength of the test of the
different species under impact. He found that, within species, strength appears to
increase with size. Strathmann’s (1981) observations on test strength and thickness
led him to set up a ranking of strength under impact. Eucidaris tribuloides was the
strongest, followed by E. viridis and D. antillarum, then T. ventricosus. Lytechinus
variegatus was the weakest species. Thus, it appears that, among the species
examined, the most primitive design is the strongest. He extrapolated the results
from modern echinoids to extinct classes and he argued that is speculative, because
it is difficult to determine the causes of failure of skeletons in most fossil forms.
Information about the decline and recuperation of populations of D. antillarum
in Central America is scarce because there are few records prior to the mass
mortality and only for several of localities in Belize (Hay 1984; Lewis and
Wainwright 1985); Honduras (Hay 1984), and Costa Rica (Valdez and Villalobos
1978; Cortés 1981). Post mortality information comes from Costa Rica (Cortés
1994; Fonseca 2003; Alvarado et al. 2004; Bolaños et al. 2005; Fonseca et al.
2006; Myhre and Acevedo-Gutiérrez 2007; Cortés et al. 2010), Nicaragua (Ryan
et al. 1998), Honduras (Lessios 1998), Guatemala (Fonseca and Arrivillaga 2003)
and Belize (Aronson and Precht 1997; Carpenter and Edmunds 2006).
Valdez and Villalobos (1978) studied the spatial distribution, and the correla-
tion with the substrate and the degree of aggregation of D. antillarum on the coral
reef at Cahuita National Park, Costa Rica in 1977. They reported densities between
3.83 and 8.8 ind m-2 and a higher degree of aggregation during morning hours that
could be a protection mechanism. In the afternoon, D. antillarum started to dis-
perse to the reef substrate to feed. Cortés (1981) reported D. antillarum densities in
this same location in 1980 between 4 and 33 ind m-2. In June 1983, a high
mortality of this sea urchin was observed in Cahuita decreasing the densities to
0.25–2 ind m-2 (Murillo and Cortés 1984). By 1992, the densities were low (0.01
ind m-2) and D. antillarum had almost disappeared from this reef (Cortés 1994).
These low densities remained from 1999 until 2003 (Fonseca 2003; Alvarado et al.
2004; Bolaños et al. 2005). Bolaños et al. (2005) indicated a low density of
D. antillarum (0.03 ind m-2) on the reef lagoon of Cahuita National Park,
in October 2003. In this same area, they reported the presence of 15 echinoderms
species (one asteroid, seven ophiuroids, five echinoids and two holothurians) that
were more diverse at sites with coral substrate and algae than seagrass or sand
sites.
Between January and November, 1984, Morales and Murillo (1996) performed
the first study on the distribution, abundance and composition of the zooplankton
present at the coral reef of Cahuita National Park. As part of the merozooplankton,
they described the echinopluteus larvae of the sea urchins D. antillarum and
L. variegatus. In general, the larvae were scarce, with a single peak in abundance in
November (36 ind m-2). This peak probably corresponded to spawning during the
new moon as observed in Panama (Lessios 1981). The low numbers of D. antillarum
larvae are presumed to be related to the high adult mortality in 1983 (Morales and
Murillo 1996).
At the CARICOMP monitoring site in Cahuita (Meager Shoal), between 1999 and
2008, the density of D. antillarum oscillated between 2 and 7 ind m-2, while den-
sities of E. viridis decreased significantly from 20 to 0.6 ind m-2 from 2000 to 2008
(Fonseca et al. 2006; Cortés et al. 2010). At this same site, densities of E. tribuloides
were always low (\1 ind m-2; Cortés et al. 2010). Thus there has been a recovery of
urchin grazers at Cahuita, and probably with a replacement of the main grazers.
For the Gandoca–Manzanillo Wildlife Refuge, Costa Rica, to the south of
Cahuita National Park, Myhre and Acevedo-Gutiérrez (2007) reported a slight
increase on D. antillarum from 2000 to 2004. In 2000, Fonseca (2003) reported a
mean density of 0.10 ± 0.22 ind m-2, that increased to 0.2 ± 0.2 in adult urchins
(40–70 mm test diameter) m-2 in 2004 (Myhre and Acevedo-Gutiérrez 2007). An
increased abundance of live coral within the refuge seems to be associated with
this small recovery in density of urchins, because in zones with high densities of
D. antillarum, the cover of non-calcareous macroalgae was low, presumably as a
result of herbivory (Myhre and Acevedo-Gutiérrez 2007).
At Corn Island, Nicaragua, Ryan et al. (1998), reported D. antillarun densities
being below 4 ind m-2 in some shallow (\ 5 m) nearshore reefs but not at the
CARICOMP sites. On those sites the mean density of E. viridis was 29.2 ± 25.2
ind m-2.
Carpenter and Edmunds (2006) reported intermediate densities (0.5–4 ind m-2)
of D. antillarum between 2003 and 2004 at Turneffe Atoll, Belize compared to
other Caribbean localities. The size frequency of the urchins ranged between \ 20
and 100–110 mm, with a modal size of 40–49 mm, a normal distribution, and with
high recruitment (ind \ 30 mm). According to their data, D. antillarum in Belize,
as well as other five sampling sites along the Caribbean, is recovering and helping
with the recovery of corals reefs by facilitating coral recruitment.
Lessios et al. (2001), using mtDNA from Diadema populations from the
Caribbean (D. antillarum), eastern Pacific (D. mexicanum) and eastern Atlantic
(D. aff. antillarum), and including specimens from Belize, Honduras, Costa Rica
and Panama, showed that D. antillarum was abundant in the Caribbean long before
(100,000 years before present) humans could have affected ecological processes.
This conclusion agreed with Jackson’s (1997) hypothesis in that D. antillarum was
abundant before humans began to remove fish from the Caribbean and con-
tradicting the idea of Hay (1984) that past high abundances, before the 1983 mass
mortality were a recent phenomenon due to the removal of its fish predators and
competitors by humans. However, it is possible that their populations were not
affected by human pressure, but their grazing impact is, reason why they became
known as one of the most important carbonate removers form the Caribbean reefs
(Ogden 1977; Scoffin et al. 1980; Bak 1994).
Vadas et al. (1982) studied the presence of Lytechinus variagetus in seagrass
beds at the Miskitos Cays, Nicaragua, reporting densities between 0.8 and 10.8 ind
m-2, test diameters and weights between 43.2 and 51.6 mm and 49.4–63.4 g
respectively, and a gonad volume between 0.7 and 2.9 cm3. In 2001, Fonseca
(2008) visited this area to describe the coral reef localities using the AGGRA
protocol. She reported the presence of nine species of echinoderms at the sampling
sites (O. reticulatus, Linckia guildingii, Asterina folium, E. viridis, E. lucunter,
D. antillarum, Meoma ventricosa, Isosticopus badionotus and Holothuria mexi-
cana; Fig. 3.3). No density or distribution was reported.
In Cayos Cochinos Biological Reserve, Honduras, Lessios (1998) reported the
presence of 13 echinoids species (E. tribuloides, D. antillarum, Arbacia punctu-
lata, L. variegatus, Tripneustes ventricosus, E. lucunter, E. viridis, Clypeaster
rosaceus, Paraster doederleini, P. jloridiensis, Brissopsis elongata, Brissus uni-
color and Meoma ventricosa). He provided information about field identification
characteristics, habitat and density. Lessios (1998) indicated that the shallow water
echinoid fauna of this Reserve appears to be typical of the Caribbean. However,
the presence of A. punctulata (0.010 ind m-2) is noteworthy, because this species
is often absent or very rare elsewhere in the tropical West Atlantic. Moreover,
L. variegatus, that is normally abundant in Thalassia tetudinum beds everywhere
in the Caribbean, was rare (0.005 ind m-2) at Cayos Cochinos in 1995, as was
D. antillarum (0.190 ind m-2). Echinometra lucunter had the highest density
(3.805 ind m-2), followed by E. tribuloides (0.467 ind m-2).
Fig. 3.4 Astropyga pulvinata aggregation in Punta Ulloa, Cocos Island, August 2007 (Photo-
graph by J. Garay)
Pacific
On the Pacific side, Alvarado (2008) studied the seasonal occurrence of Astropyga
pulvinata in Bahía Culebra, located on the north Pacific side of Costa Rica. During
upwelling months (December to April), this sea urchin forms aggregations as a
protection mechanism from predators while feeding. During the rainy season when
non-upwelling occurs, the populations of A. pulvinata were reduced in the bay, and
probably migrating into deeper waters. Alvarado (2008) calculated an average of
679 ind month-1 for the upwelling season and 159 ind month-1 for the non-
upwelling season. The average number of aggregations per month was 21.6, with
an average of 27 ind per aggregation. Most individuals have a test diameter
between 70 and 120 mm. These aggregations act as refuges for other animals like
fish juveniles and crustaceans, thus having an important conservation value. This
aggregation behavior was also observed in Cocos Island in August 2007 (J. Garay
pers. comm.; Fig. 3.4).
Alvarado and Fernández (2005) reported the presence of 25 species of echi-
noderms (four species of asteroids, six species of ophiuroids, five species of
echinoids and ten species of holothuroids) from Marino Ballena National Park, on
the south Pacific of Costa Rica. Echinoderms densities were low (0.01–1.92 ind
m-2). Diadema mexicanum (Fig. 3.5) had densities between 0.05 and 1.03 ind
m-2, being highest on the farthest site from the coast. They argued that the lower
densities were probably the result of the high sedimentation, producing a negative
effect on the diversity of echinoderms and on the development of coral reefs
(Alvarado and Fernández 2005; Alvarado et al. 2009).
In Cocos Island, Costa Rica, Alvarado and Chiriboga (2008) reported the pres-
ence of 28 species of echinoderms (seven species of asteroids, six species of
ophiuroids, nine species of echinoids and six species of holothuroids) located
between 3 and 18 m deep at 18 sites around the Island. The sea urchin D. mexicanum
was the only echinoderm present at all sites and depths studied, and provided a
density between 0.05 and 6.53 ind m-2.
For the Pacific side of Nicaragua, on the Rivas Department, Alvarado et al.
(2011) reported the presence of 11 species of echinoderms (one species of asteroid,
six species of echinoids, and four species of holothuroids). The sea urchin
D. mexicanum was present in nine of the ten study sites, with a density between
0.04 and 1.10 ind m-2, while Echinometra vanbrunti had the highest density (4.09
ind m-2) of all echinoderms studied.
Carballo-Bolaños and Pocasangre-Orellana (2007) describe the echinoderm
intertidal fauna to a depth of 2 m in the Los Cobanos rocky reefs system, a natural
protected area in El Salvador. They reported the presence of 13 echinoderms
species (five species of ophiuroids, two species of echinoids and six species of
holothuroids). The ophiuroids were the most abundant group with 1,771 individ-
uals at the study sites, and followed by the sea cucumbers with 877 individuals,
while there were only 22 individuals of Echinometra vanbrunti and Astropyga
pulvinata. The ophiuroids Ophiocoma aethiops and O. alexandri (Fig. 3.5),
and the sea cucumber Holothuria kefersteini had the highest densities (0.15, 0.14
and 0.13 ind m-2, respectively). These species were more common in the upper
intertidal zone than in the lower intertidal zone. They also reported, for the
first time in El Salvador, the presence of the ophiuroid Ophiothrix rudis.
Moreover, underwater surveys in the same protected area found subtidal echinoids:
A. pulvinata, Toxopneustes roseus, D. mexicanum and Eucidaris thouarsii.
Astropyga pulvinata was absent on subtidal boulders at Pirigallo and Menaguera
Islands, Gulf of Fonseca (2001–2008) (E. Baraza pers. observ). In November 2009
some scattered individuals were found and in 2011 A. pulvinata were widespread
at both places (E. Baraza pers. observ).
Holothuroid Ecology
Caribbean
Hasbún and Lawrence (2002) describe the presence of five holothurians

(Isostichopus badionotus, Holothuria mexicana, H. thomasi, H. arenicola and
Actinopyga agassizi) in Cayos Cochinos Biological Reserve, Honduras Caribbean.
They provide detailed information about the spicule width, length and height from
Fig. 3.5 Common Central America Pacific echinoderm species. a Phataria unifacialis;
b Nidorellia armata; c Pharia pyramidata; d Pentaceraster cumingi; e Echinothrix calamaris;
f Diadema mexicanum; g Ophiocoma aethiops; h Ophiocoma alexandri. (Photographs by
G. Edgar, S. Sunagawa, B. Bezy and J.J. Alvarado)
the tegument and tentacles. They also reported the presence of the commensal
pearlfish, Carapus bermudensis, living in H. mexicana and A. agassizi (Fig. 3.3).
Pawson and Miller (1988) describe, with specimens of Thyone tanyspeira

(Phyllophoridae), collected in Honduras, Colombia and Trinidad. The specimens
were collected during the expedition of the RV Pillsbury in 1969 and 1971 into the
Caribbean.
Pacific
Bakus (1974) compared the toxicity of holothurianss in order to test the hypothesis
that toxicity in holothurians is inversely related with latitude and that this phe-
nomenon is in part a reflection of fish predation. He conducted the study at Cocos
Island (Costa Rica), Bahía de Bacochibampo (México), Santa Catalina Island
(California), and Friday Harbor (Washington). Six of the seven species of holo-
thurians at Cocos Island were toxic to fishes, three of six holothurians were toxic to
fish in Mexico, while one of two were toxic to fish in California and three of
twelve species in Washington were mildly toxic to fish. At Cocos Island the toxic
species were Holothuria theeli, H. atra, H. impatiens, H. sp. 1, H. sp. 2 and
Bohadschia sp. The body wall, viscera, and Cuverian tubules (in H. theeli and
H. sp.2) were the most toxic to fishes.
3.2.2 Grazing and Bioerosion
Caribbean
In the Miskitos Cays, Nicaragua, Vadas et al. (1982) studied the abundances,
feeding and reproductive activities of the sea urchin Lytechinus variegatus on a
seagrass bed (Thalassia testudinum). Intermediate (2–4 m) and deeper (4–8 m)
depths contain fewer urchins and have higher plant diversities. Although highest
plant diversities were correlated with intermediate grazer densities (*5 ind m-2).
The diet of these urchins appears to be governed primarily by availability and the
palatability of food items. Detrital T. testudinum and sediment were the two major
food items in the diet (40 and 25 %, respectively). This feeding strategy minimizes
the impact of urchin grazing on the structure and the viability of T. testudinum and
the algal-seagrass community. Over 68.5 % of the volume of the gut contents of
L. variegatus consisted of particle sizes in the 0.5–3.0 mm category. Intestinal
tract pH ranged from 6.0 in the esophagus and 6.5–7.0 in mid-stomach. The
average dry weight of the gut contents was 113 mg cm-3. Of this 68 % was
CaCO3. Food passage in the gut averaged 0.22 cm3 ind-1 h-1 for the first 14.5 h,
then values decreased rapidly. The consumption rates of L. variegatus that was
taken from T. testudinum was 0.6 g dry wt day-1.
Hay (1984) compared the pattern of fish and urchin grazing on overfished and
less fished reefs in Haiti, Saint Croix, Saint Thomas, Belize, Honduras and Pan-
ama. In Belize he worked at Carrie Bow Cay and Lighthouse reefs and in
Honduras on Media Luna and Becerro reefs. On the overfished reefs, the rate of
T. testudinum removal increased significantly with depth, urchin densities were
high, and urchin grazing was equal to, or greater than, fish grazing in shallow
habitats. At the overfished reefs (Haiti, Saint Croix and Saint Thomas) the percent
of T. testdinum eaten per hour by sea urchins was between 0 and 10.8 % with a
D. antillarum density range between 5-20 ind m-2. At the less fished reefs
(Honduras and Belize) it was between 0 and 1 % with D. antillarum densities
of \ 1 ind m-2 for the Honduras sites and around 0.2 and 8 ind m-2 for the Belize
sites. He concluded that on the reef unaffected by humans, the overall impact of
urchins on macrophytes was minor relative to the impact of grazing fishes.
The combined effects of a reduction in herbivory due to D. antillarum mass
mortality in 1983–1984 (Lessios et al. 1984) and the increase in vacant substratum
because of coral death (Aronson et al. 2000) have caused dramatic changes in the
Caribbean. This has led to several attempts to understand the ecosystem dynamics
after these events. McClanahan (1999) examined the ecology of a grazer living
sympatrically with D. antillarum and Echinometra viridis. He assessed the role
that finfish and invertebrate predators play in controlling the distribution of
E. viridis as well the ability of E. viridis to control exposed fleshy algae on patch
reefs on the Glovers Reef Atoll lagoon. McClanahan’s (1999) results showed that
the abundance of E. viridis is constrained by predation, which restricts E. viridis to
cryptic locations. He proposed that this sea urchin is unable to control exposed
erect fleshy algae as well as D. antillarum (McClanahan 1999).
In despite of McClanahan’s (1999) results, Aronson et al. (2002a) reported that
high densities of the sea urchin E. viridis kept the cover of fleshy and filamentous
macroalgae to low levels. They concluded that herbivorous activity of E. viridis
could help re-establishment of the coral Agaricia tenuifolia. This trend has been
observed by Aronson and his colleagues throughout the Caribbean as well in
Belizean reefs (Aronson 2002b; Aronson et al. 2005). Carpenter and Edmunds
(2006) reported that dense populations of Diadema now occur over a multi-
kilometer-wide scale across the entire Caribbean. They conclude that population
recovery of D. antillarum is occurring at both local and regional scales and that
grazing by this echinoid is creating conditions that favor the recruitment of corals
(Carpenter and Edmunds 2006).
Brown-Saracino et al. (2007) examined urchin population characteristics,
bioerosion rates, their fish predators, and potential competitors on three unpro-
tected reefs and one reef within Hol Chan Marine Reserve in the lagoonal regions
of the reef system off Belize. Their report presents densities of Balistidae and
Labridae (as predator fish of urchins) and Labridae (potential competitors of
urchins). Population data on coral reefs and algae were collected. They recorded
urchin wet weight and test diameter. They also analyzed gut contents to estimate
yearly bioerosion rates per square meter of reefs. Six species of sea urchin were
found at the study sites, D. antillarum, E. lucunter, E. viridis, E. tribuloides,
L. variegatus and L. williamsi. Echinometra viridis was the dominant species at all
but one of the patch reef sites surveyed. Furthermore, D. antillarum was found in
low abundances and restricted to northern sites. Balistids were rare at all sites,
although labrids were relatively abundant. A significant inverse relationship
between the abundance of Labridae and urchin densities was found. Parrotfish and
urchin densities were not related. Calcium carbonate comprised approximately
90 % of the urchin gut content, and followed by organic material. Urchin bioe-
rosion rates ranged from lowest at Hol Chan Reserve (*0.2 kg CaCO3 m-2 yr-1)
to highest at Coral Gardens (*1.4 kg CaCO3 m-2 yr-1), an unprotected area close
to Hol Chan. Small urchins and low proportion of organic material in gut contents
were found at low algae covered sites, suggesting that the herbivors were food
limited. Taken together their results indicate that both top-down and bottom-up
factors contribute to the control of sea urchin populations in the lagoons off Belize.
They also suggested that Marine Protected Areas (MPA) may provide the means
for limiting sea urchin densities and bioerosion of coral reefs and result in reef
recovery, returning the calcium carbonate cycle to a more balanced state.
Pacific
Fischer (1990) described the importance of D. mexicanum as a bioeroder in Costa

Rica. This species is found in very shallow waters (down to 8 m), and produces
stronger bioerosion on the limestone. At Caño Island, Guzman (1988) found low
densities of this sea urchin, but nevertheless it caused bioerosion at the base of
massive corals. Also, at Cocos Island grazing activities of this sea urchin caused
high levels of bioerosion and reduced coral recruitment (Guzman and Cortés
1992). After the 1982-83 El Niño event, Guzman and Cortés (1992) stated that
coral recovery would take centuries due the high densities of bioeroders and low
coral reproduction rates. Guzman and Cortés (1992, 2007) indicated that part of
the deterioration of the reef structures at Cocos Island was due to the bioerosive
action of D. mexicanum.
In 2002, there was a five-fold increase of coral cover and a notable sea urchin
reduction (Guzman and Cortés 2007). Guzman and Cortés (2007) determined that
D. mexicanum was not playing a important role on the reef bioerosion. However,
the urchin could still fulfill its key herbivorous role in assisting the recruitment of
corals, as has been observed in other reefs on the Caribbean (Sammarco et al.
1974; Sammarco 1980, 1982a, b; Mumby et al. 2006). Lessios et al. (1996)
reported the presence of an Indo-Pacific echinoid (Echinothrix diadema and
Echinothrix calamaris; Fig. 3.5) at Cocos Island, and that probably arrived to the
island during the 1982–1983 El Niño (Lessios et al. 1996, 1998). These two
species are well known bioeroders in the Indo-Pacific (Bak 1990; Peyrot-Clausade
et al. 2000; Carreiro-Silva and McClanahan 2001). However, their impact in
Cocos Island is unknown. But due to their current densities (0.03–0.02 ind m-2;
Alvarado and Chiriboga 2008) it could be predicted that they would be less
compared to that of D. mexicanum.
3.2.3 Reproduction and Ontogenetic Development
Caribbean
Hendler (1988b) analyzed the ontogenesis of the oral papillae in representatives of

the four major groups of Amphiuridae in order to ascertain if comparisons of
growth series during postlarval ontogenesis can be used to distinguish homologous
structures in closely related taxa. Based on the development of the oral papillae,
Hendler (1988b) concluded that the post-larval ontogenesis of the amphiurids is
more reliable than larval morphology as an indicator of phylogenetic affinity.
Byrne (1991) studied reproduction and development of the ophiuroid Ophionereis
olivacea, a protandric hermaphrodite species that brood its young. She established
that sex reversal occurs at disk diameters ranging between 2.2 and 4.0 mm.
Byrne (1988) examined the oogenesis of Ophiolepis paucispina to determine
the vitellogenic mechanism involved with the production of the large oocytes and
the role of the genital haemal sinus. Anatomical examination of the gonads showed
that the ovary wall consists of two parts, the inner and outer sac, which is separated
by the genital coelom. The inner sac is composed of the coelomic epithelium,
which is a connective tissue layer that contains the haemal sinus, and the germinal
epithelium. Experimental evidence revealed that each oocyte is surrounded by the
haemal sinus during oogenesis. It appears that that the oocytes bulge into the
haemal sinus as they enlarge, and thereby, develop an evagination in the genital
sinus. The Golgi complex and endoplasmic reticulum are probably involved in the
elaboration of yolk bodies. Endocytotic activity is ubiquitous during oogenesis,
and indicated the uptake of exogenous material.
Byrne (1988) concluded that the large oocytes of O. paucispina are produced
through encytotic incorporation of yolk precursors derived from a somatic source,
and that the haemal sinus functions as the proximate source and store of these
precursors. Byrne’s (1988) results together with the results of other studies (see
Byrne 1988 for references) suggests that extraovarian vitellogenesis may be
common in echinoderms. Additionally, Byrne (1989) described the ultrastructure
of the ovary and oogenesis of O. paucispina. She documented the developmental
of oocyte growth from the early proliferative stage through the previtellogenic and
vitellogenic stages. Byrne (1988, 1989) contributed evidence that the echinoderm
genital haemal sinus is an intragonadal nutrient store. Byrne (1989) concluded that
her experimental evidence, taken together with the biochemical evidence for the
somatic and ovarian synthesis of vitellogenin, indicate that entirely autosynthetic
yolk formation may be rare, whereas mixed auto- and heterosyntheticy yolk for-
mation may be common in Echinodermata.
Pacific
The only reference to reproduction in echinoderms in the Pacific is that of Pearse

(1968) with D. mexicanum at Cocos Island. He reported that all the individuals
studied showed all gametogenic stages in the gonads, and suggested that the
reproduction occurs year-round. Furthermore, there is no research presently
available on crinoid, asteroid or holothuroid reproduction from the Caribbean or
Pacific coasts.
3.2.4 Pollution
Rojas et al. (1998) studied the effect of coastal pollution at Cahuita coral reef on
the sea cucumber Holothuria mexicana. They determined metal concentrations in
the sea cucumbers and suggested their use as bio-indicators in the future. The
respiratory tree was the structure that showed the highest levels of metals, with the
exception of Cu and Pb which were respectively higher in the muscles and the
body wall, respectively (Rojas-Guitart 1990; Rojas et al. 1998).
3.2.5 Physiology and Anatomy
Lawrence (1967) studied the gut’s lipid reserves of three sea urchins (E. lucunter,
T. ventricosus, and L. variegatus) from Portete (Costa Rica). He calculated the
ratio of the amount of total and neutral lipid (in mg) in the gut with respect to size
of the individuals. Echinometra lucunter had the greatest amounts of total lipid and
neutral lipid. The ratio of total lipid to body size was 1.00 for E. lucunter, while it
was only 0.68 for T. ventricosus and 0.40 for L. variegatus. Also, E lucunter had
the highest gonad index, test index, and lantern index. Tripneustes ventricosus
appears to be the most fragile of the three urchins because its test and lantern
indices were the lowest. The high levels of neutral lipid found are comparable to
that in the gut of temperate urichins, such as Strongylocentrotus purpuratus
purpuratus, and higher than that in the gut of Stomopneustes variolaris.
Hendler (1984b) made the first report of color change in brittlestars. He studied
individuals of all the Caribbean species of the genus Ophiocoma: O. echinata,
O. paucigranulata, O. pumila and O. wentii. All four species were found to change
color in natural light regimes. Color change was observed in situ at Panamá, while
experimental data was recorded from individuals at Carrie Bow Cay, Belize.
He observed five individuals of each Ophicoma species in order to document diel
color-change. He also analyzed severed arms of O. wendtii and other individuals
with broken arms in order to find out if isolated arms would maintain the same
pattern of color-change as intact individuals. Additionally, portions of dorsal arm
plates chipped from O. wendtii arms were examined to find whether the chips
change color at the same time as whole brittlestars and severed arms. His results
showed that the day-night color change involves a replacement of brown shades by
gray and black. Individuals of all four species change color in the late afternoon.
Experimental evidence demonstrated that transformation occurs over a period of
three to four hours and it is effected by chromatophores. Furthermore, as seen with
the separated body parts, they appear to respond to light independently of the
nervous central system.
During the day O. paucigranulata was brownish-black. At night the overall
color fades to dark gray. In O. echinata regions that are brown during the day turn
into gray at night. Ophiocoma pumila undergoes a comparatively subtle color-
change during the day from yellowish-brown to brownish-gray. At night, the
brown shades turn to gray and the gray ones turn black. The color-change of
O. wendtii is striking. During the day, specimens had a homogeneous brownish-
gray color-phase, including the spines that sometimes have brownish-orange tips.
At night, the disk turns grayish–brown and black and the dorsal side of the arm
develops conspicuous gray or white and black bands while the ventral side of the
arm remains unbanded. At night the arm spines often turn white basally, gray
along the shaft, and orange.
Hendler (1984b) demonstrated that the four species are not equally sensible to
light. Three species withdrew their arms into crevices in response to illumination,
while O. pumila showed little reaction to illumination. Hendler (1984b) tested
whether O. paucigranulata was more sensitive than O. echinata by measuring the
speed of reaction. Ophiocoma wendtii was the most sensitive species, reacting
rapidly to the lights even when the light was attenuated with two layers of red
cellophane. The other three species did not react to dim red light. Ophiocoma
wendtii reacts to lower levels of illumination at night more than during the day,
and exhibits negative phototaxis in moonlight as well as in sunlight. He argued that
color-change is either an endogenous rhythm, or a response to the light regime and
that the color-change may expose photosensitive tissues that control the ophiu-
roid’s detection of shaded places in the reef. Finally, he concluded that negative
phototaxis, coordinated with color-change, may facilitate the defensive shadow-
seeking response under varying illumination levels. Color change also confers
pigmentation patterns which may camouflage ophiuroids from predatory fish.
Hendler and Byrne (1987) described for the first time a presumed photoreceptor
system in the body wall of the ophiocomid brittlestar O. wendtii, and it was based
on examination of the fine structure of the dorsal arm plate (DAP). This led Cobb
and Hendler (1990) to test the photoreceptor hypothesis, that microscopic, trans-
parent structures of the skeletal plates, and that their associated chromatophores
and nerve bundles comprise a photoreceptor system in O. wendtii as porposed by
Hendler and Byrne (1987).
In order to examine the functional, systematic, and paleontological importance
of their distinctive integument, Byrne and Hendler (1988) undertook a compara-
tive examination of the arm structure of shallow water and bathyal ophiomyxid
brittlestars. Additionally, they used the behavior and arm morphology to evaluate
the specialization of Paleozoic ophiuroids. Specimens of Ophiomyxa flaccida,
Ophiomyxa tumida and Ophiogeron supinus, from the subfamily Ophiomyxinae;

and Ophiobyrsa serpens, from the subfamily Ophiobyrsidae, were collected at
several Caribbean locations. Ophiomyxa flaccida was collected using SCUBA in
Belize, Panamá and Bahamas Islands at depths between 5-25 m. Ophiobyrsa
serpens was collected at a depth of 24 m on the Belize Barrier Reef. Ophiomyxa
tumida and O. supinus were collected with the Johnson-Sea-Link submersible in
the Bahamas Islands at depths between 300–600 m. Their analyses showed that the
arms of O. supinus were largely feltwork connective tissue fibers with the
predominance of interfibrillar matrix, while the arm integument of O. flaccida,
O. tumida and O. serpens was composed of a thin layer of feltwork connective
tissue and a thick layer of fibrillar connective tissue. The authors’ observations on
the locomotion behavior and morphology of the ophiomyxid brittlestar argued
against the sedentary nature of Paleozoic ophiuroids. They noted that limited arm
mobility, rather than integumentary structure, is evidence of functional similarity
between O.supinus and some Paleozoic ophiuroids. They suggestsed Paleozoic
ophiuroids may have been epifaunal forms with a pattern of locomotion more like
some recent astropectinid asteroids than amphiurid ophiuroids.
3.3 Diversity and Distribution
Maluf (1988a, b) synthesized echinoderm distribution and composition from

California to Peru. She reported 630 species, and peaks of richness in the Gulf of
California, Panama and the Galapagos Archipelago. The last two are places
involved a high sampling effort. Then Maluf (1988b) compared species similarity
and found that CA, from Guatemala to Costa Rica, is a separate unit. She also
indicated that the zone between Panama and Costa Rica possess a high degree of
endemism and is a transition point, while Panama, Colombia and Ecuador had a
greater affinity.
Alvarado and Cortés (2004) considered both the Caribbean and Pacific echi-
noderms of Central American. They found 315 species, with Panama and Costa
Rica having the greatest number of species. However, their calculations of simi-
larity were done without any coast differentiation or discrimination of the presence
of the oceanic island, Cocos Island that belongs to Costa Rica. This fact prevents a
detailed evaluation of the richness of the Central American Pacific coast. In later
investigations, Alvarado et al. (2008), Miloslavich et al. (2010) and Alvarado
(2011) reported 253 species of echinoderm fauna for the Caribbean of CA,
including the Archipelago of San Andres and Providencia (Colombia). In com-
parisons with the other countries of the Caribbean, such as Costa Rica and Gua-
temala, they are the less diverse countries, but probably the CA coast is the least
studied (with the exceptions of Panama and Belize) (Alvarado 2011). For the
Pacific side of CA, Alvarado et al. (2010) reported 287 echinoderm species.
In this chapter, excluding Panama (Table 3.1, Appendix) we report a total of
416 species of echinoderms for both the Pacific and the Caribbean coast of CA.
Table 3.1 Number of echinoderm species per class in each country by coast (from Alvarado et al. 2008, 2010, Alvarado and Cortés 2010, Alvarado 2011,
and Appendix)
Coast Country Crinoidea Asteroidea Ophuiroidea Echinoidea Holothuroidea Total
Pacific Guatemala 0 0 7 5 4 16
El Salvador 0 10 17 13 18 58
Honduras 0 0 3 4 4 11
Nicaragua 0 6 12 14 7 39
Costa Rica 0 12 38 28 28 106
Cocos Island 2 33 30 31 31 127
Caribbean Belize 5 18 58 27 26 134
Guatemala 2 5 4 8 4 23
Honduras 11 30 14 29 11 95
Nicaragua 9 30 6 19 2 66
Costa Rica 4 7 16 13 4 44
Caribbean coasts 16 54 66 47 31 214
Pacific coasts 2 46 60 48 53 207
Total 18 100 123 94 80 416
3 Central America Echinoderms: Diversity, Ecology and Future Perspectives
93
According to Pawson (2007) 7,000 echinoderm species have been reported

globaly. CA is a very diverse area, with 5.9 % of the world species. The Caribbean
and Pacific coasts are very similar in the number of total species (214 and 207
species, respectively, with seven in common; Table 3.1), and differing more
notably in the numbers of holothuroids and crinoids. The first group is more
diverse in the Pacific and the second group is more diverse in the Caribbean. It is
important to note that the number of species from the Pacific coast without con-
sidering Cocos Island and the Cocos Ridge, CA possesses only 126 species. Cocos
Island and the Cocos Ridge contributed with 39 % of the echinoderm species of
the Pacific of CA. Even without Cocos Island, the Pacific coast of Costa Rica is the
richest in CA with 106 species (Table 3.1). Honduras is the least diverse
(11 species). However, Cocos Island is the richest location on the Pacific with 127
species. Belize (134 species) is the richest country on the Caribbean, while
Guatemala (23 species) is the least diverse. Ophiuroidea is the most diverse class
on both coasts, while Crinoidea is the least diverse (Table 3.1).
Seven species of echinoderms are found on both coasts: three ophiuroids
(Amphiodia violacea, Amphipholis squamata and Ophiactis savignyi), one echi-
noid (Clypeaster speciosus) and three holothuroids (Holothuria arenicola,
Holothuria impatiens and Benthodytes sanguinolenta). Amphiodia violacea is
present on the Pacific coast of Costa Rica and Cocos Island, and in the Caribbean
coast of Belize. Amphipholis squamata is present on the Pacific coasts of Costa
Rica and Nicaragua, and on the Caribbean coast of Belize, Guatemala and Hon-
duras. Ophiactis savignyi is present on the Pacific coast of El Salvador, Nicaragua,
Costa Rica and at Cocos Island, and on the Caribbean coast of Belize, Honduras
and Costa Rica. Clypeaster speciosus is present on the Pacific side at Cocos Island,
and on the Caribbean coast of Costa Rica. Holothuria arenicola is present on the
Pacific coast of Costa Rica and at Cocos Island, and on the Caribbean coasts of
Belize, Guatemala and Honduras. Holothuria impatiens is present on the Pacific
coast of El Salvador, Costa Rica and at Cocos Island, and on the Caribbean coast
of Belize. Benthodytes sanguinolenta is present on the Pacific side at Cocos Island
and on the Caribbean coast of Honduras (see Appendix).
Six species are distributed on all the countries coasts of the Pacific side
and Cocos Island: Ophiocoma aethiops, Ophiocoma alexandri, Echinometra
vanbrunti, Clypeaster rotundus, Holothuria difficilis and Isostichopus fuscus. Ten
species are found on the Caribbean coasts of all the CA countries: Davidaster
discoidea, Astropecten articulatus, Oreaster reticulatus, Astrophyton muricatum,
Ophiocoma echinata, Eucidaris tribuloides, Diadema antillarum, Echinometra
lucunter, Meoma ventricosa and Holothuria mexicana. One hundred and twenty-
eight species are reported for the Pacific coast of only one country, 43 species for
two countries, 14 species for three countries, three species for five countries and
six species for six countries. On the Caribbean coasts of CA 130 species are
reported for only one country, 43 species for two countries, 25 species for three
countries, five species for four countries and ten species for five countries (see
Appendix).
3.4 Paleontology
Donovan et al. (2005) indicated that the knowledge of echinoid fossils from Belize
is poorly documented. In their paper they reported the presence of two echinoids:
Echinolampas sp. cf. aldrichi from the Mid Miocene to Quaternary, and
Echinocorys sp. cf. scutata from the Mid Eocene. Gischler (2010) analyzed five
Holocene sediment cores from Glovers and Lighthouse reefs, and Turneffe islands,
Belize. He picked out echinoid spines, plates, and teeth. These were identified to
genus based on comparison with these pieces of echinoids collected live and
treated with dilute sodium hypochlorite and dried. A few fragments of the massive
spines of E. tribuloides and the burrowing echinoid Clypeaster were found in cores
from Turneffe. Diadema and Echinometra pieces were the most commonly
encountered. Gischler (2010) reported that three significant echinoid mass mor-
tality events occurred during 6.3–6.1 kyr BP (Glovers reef) as well as 4 and 1.3 kyr
BP (Turneffe Islands). The first mass mortality event apparently only affected
Diadema. The second and third events affected both Diadema and Echinometra.
Alvarado et al. (2006) reviewed echinoderm paleontological research that
occurred from 1880 until 2001 in Costa Rica, and listed 46 species. The fauna is
composed of Crinoidea with one genus and Echinoidea with seven orders, 18
families, 28 genera and 45 species. Of the echinoids, the order Spatangoida was
the most diverse, with seven families, 15 genera and 26 species. Within this order,
the most diverse family was Brissidae, with eight genera. The genus Schizaster had
seven species. They indicated that the majority of species are Cretaceous and
Miocene, from 11 geological formations. The fossil echinoderm fauna of Costa
Rica is similar to other areas of CA and México, with a good record from the
Miocene and a notable decline to the Pliocene (Buitrón 1978). Alvarado et al.
(2006) pointed out that a greater effort is needed in the investigation of fossil
echinoderms to improve its understanding, and to have a real appreciation of its
composition. Specimen identification, due to lack of specialists and literature, is
one of the greatest problems. There is also a need to explore more areas in CA.
3.5 Threats to Echinoderms
Echinoderms are vulnerable to a number of threats from natural and anthropogenic

origin. The threats can also be classified as directly or indirectly related to echi-
noderms. Any deterioration of the environment where the echinoderms live can be
classified as an indirect threat. Because the majority of echinoderms in CA live in
shallow coastal waters, the most important habitats are coral reefs, rocky reefs and
seagrass meadows. All threated due to human activity.
Cortés (2007) indicated the main natural disturbances to coral reefs in CA are
the following: (1) extensive mortality of corals due to bleaching associated with
extremes in water temperature or salinity, sediment or bacterial infection;
Fig. 3.6 Portete reef platforms uplifted after a 7.5 earthquake on the Southern Caribbean of
Costa Rica in 1991 (Photograph by J. Cortés)
(2) hurricanes; and (3) tectonic activity. On April 22, 1991, a 7.5 magnitude
earthquake in Southern Caribbean of Costa Rica resulted in an uplift of the coast of
between 50 and 190 cm (Fig. 3.6), and caused mass mortalities among the inter-
tidal and reef communities (Cortés et al. 1993, 1994). Among those populations
most affected were the corals, algae, sea urchins and chitons. Dense populations
found in a narrow band just below sea level before the uplift of the sea urchins
E. lucunter, L. variegatus and M. ventricosa were killed by the exposure in Portete
reef platform; the most impacted was E. lucunter (Cortés et al. 1993; Alvarado
et al. in prep.).
Among the main anthropogenic impacts to coral reefs in CA, Cortés (2007)
emphasized pollution, including garbage, oil, heavy metals from farming and
industry, and sedimentation. Cortés (2007) also mentions tourism as an important
source of disturbance induced by curio collection, construction of tourism facili-
ties, anchor and boat-related damage and damage caused by divers and snorkelers.
In this sense, tourism can also be viewed as a direct threat to echinoderms, and
especially to regular and irregular intertidal sea urchins that are collected as a
personal souvenir or for aquaria. Even if there is no official register of these
collections, anecdotal comments from local residents indicate that the populations
of some sand dollars have diminished over time. Inside Marine Protected Areas
(MPA) this activity is banned, but due to lack of patrolling and conscience this
practice continues.
The most important direct threat to echinoderms in CA is illegal fisheries and
by-catch of longline and trawling fisheries. Sea cucumbers fisheries is probably
one of the main illegal fisheries in CA. In El Salvador, this activity is strictly
regulated since holothurians are considered threatened species due to unauthorized

fisheries for overseas smuggling (Decree No. 36 Ministry of the Environment and
Natural Resources; Diario Oficial No. 383 (103) 5 June 2009, p. 75). The activity
stopped after legal prosecutions and holothurians are recolonizing depleted areas.
In Costa Rica sea cucumber fisheries are forbidden from any MPA, but an official
decree (N823.138/MAG; La Gaceta N880, 27th April 1994, p. 4) of the Ministry of
Agriculture establishes a maximum of ten permits for artisanal fishers for fisheries
of sea cucumbers of the species Holothuria inornata and Isostichopus fuscus. The
decree established a maximum of 1,200 sea cucumbers per fisher per month per
defined area. The permit is for two months between October and April and the sea
cucumbers can be disembarked only at official ports. They are subject to inspection
and fishermen have to present a monthly report. We have contacted the Costa
Rican authority (Instituto Costarricense de Pesca y Acuicultura-INCOPESCA) to
get more information about the decree and its implementation, because there was
no baseline investigation to establish the quotas or the target species, or any
research about the status of the populations, or reproduction seasons for the
establishment of that decree. The answer from the national authority is contra-
dictory. Some officials tell us that there is no sea cucumbers fishery at the present.
Other officials said that there are Japanese and Mexican carrying out this activity
on the North Pacific coast of the country. There are no records of any permit
granted. According to Toral-Granda (2008), in a Global FAO report of sea
cucumber fishery, Hong Kong imported from Costa Rica 1,268 kg dry weight of
sea cucumbers from 1999 to 2004 (1999: 108 kg; 2000: 664 kg; 2001: 325 kg;
2003: 7 kg; 2004: 164 kg). One threat to sea cucumbers in Costa Rica, and
probably for the rest of CA, is the lack of coordination, register, control and
patrolling by the national authorities, INCOPESCA and the ministry of Agricul-
ture, the Environment and Foreign Trade.
Toral-Granda (2008) indicated that sea cucumber fishing activities in Nicaragua
dated back to 1994 where the first exports are recorded. In 2005, oriental traders
requested a permit to exploit this resource. A one-year permit was granted while
scientific research was undertaken on the Pacific coast. No taxonomic information
exists on the species harvested during that year. Currently three species are known
to be commercially harvested. At least another seven unidentified sea cucumbers
are harvested. In the Pacific coast, the most important species is I. fuscus. In the
Caribbean, the most important species are H. mexicana, H. arenicola and
I. badionotus. All products are exported to China and Taiwan as there is no
consumption in Nicaragua. Toral-Granda (2008) indicated that Hong Kong
reported imports from Nicaragua involving 6,078 kg dry weight from 2003
to 2005 (2003: 252 kg; 2005: 5,826 kg). Currently, no sea cucumber fishing
activities are regulated. However, there is a plan to start applying total allowable
catches and fishing seasons to both Pacific and Caribbean species in Nicaragua
(Toral-Granda 2008).
By-catch is one of the most significant issues affecting fisheries management
today, due to the incidental mortality of non-target species. This affects biodi-
versity and trophic food webs (Hall et al. 2000). According to Allan Bolaños
(Programa de Restauración de Tortugas Marinas-PRETOMA, Costa Rica, pers.

comm. 2011), a recent study of artisanal fishery on the Pacific coast of Costa Rica
showed that every 1,000 hooks captured 15–20 sea stars (Astropecten aramatus,
Luidia tessella, L. superba, and Tethyaster canaliculatus, this last two new reports
for Costa Rica). Ingo Wehrtmann (Universidad de Costa Rica, pers.comm 2011)
pointed out that deep shrimp trawling (depth of 150–200 m) did not capture any
echinoderms. However, in shallow waters (depth of 20–50 m) there was a by-catch
of echinoderms (Allan Bolaños, pers. comm. 2011). In El Salvador asteroids are
captured during shrimp trawling activities (10–50 m). There are no studies on the
possible disturbances associated with this activity.
The natural direct threats to echinoderm populations are less numerous or
common in CA. The only well documented one has been over the mass mortality
of D. antillarum by a waterborne pathogen in the Caribbean (see ecology section).
For the Pacific side, there are no reports of disease or mass mortalities, or research
focused on test abnormalities, or parasites.
CA is a region with high echinoderm diversity, a products of its rich evolutionary

and geological history and its particular geographical location. This fact has
attracted the attention of researchers for the last 150 years, as indicated in the
number of publications that we have reported. Nevertheless, these 150 years of
investigations have not been equal for all countries of the region. The scientific
history of the region cannot be separated from the political history. The various
armed conflicts and the political instability were factors that resulted in a low
priority given scientific research. Panama, Belize and Costa Rica have been rel-
atively more stable compared to Guatemala, Honduras, El Salvador and Nicaragua
and they lead in research and consequently in greater numbers of publications, and
hence, more species reported. The case of the Caribbean of Honduras and Nica-
ragua is interesting. For example, the results of the RV John Elliot Pillsbury
expedition reported a high diversity of asteroids, however, the sampling was
limited to a few stations. Comparing the richness of species in the Caribbean,
Alvarado (2011) indicates that the number of species from Central America would
increase considerably if research increases. Moreover, Alvarado (2011) explains
that the richness of species in Belize, Costa Rica and Panama is a reflection of the
presence of important research centers like CIMAR, STRI and the station at Carrie
Bow. Nevertheless, the richness of environments that these countries possess also
contributes to the high numbers of species. Guatemala is a special case, since its
Caribbean coastline is very short and, its Pacific coast consists of sandy beaches.
This results in a smaller number of species.
This chapter has provided evidence that the great majority of investigations
have been focused on the Ophuiroidea and Echinoidea, but few efforts on Aster-
oidea and Holothuroidea and practically none on Crinoidea. Most of the research
has been on the Pacific, and indicates that strong efforts are needed on the
Caribbean coast. There are few studies focused directly on Asteroidea and
Holothuroidea since they generally only mentioned as part of the echinoderm
fauna of a particular region.
Ecology, taxonomy and evolution have been the main lines of investigations in
CA. These research efforts should continue, and at the same time, studies focused
on reproduction and physiology should be encouraged. Moreover, it is necessary to
start research in fisheries, aquaculture and management that help stakeholders and
authorities make decisions for more sustainable extractions. To be able to conduct
research in CA, it is necessary to support and to promote the exchange of infor-
mation among the countries, as well as to fund the formation of new researchers
for this group.
Acknowledgments We would like to thank A. Bolaños, I. Wehrtmann, C. Fernández,

M. Springer, R. Vargas, F.A. Solís-Marín and the late C.G. Ahearn for their help with literature,
comments and museum access. M. Chavez for the elaboration of the figures. JJAB thanks
CONICIT, MICIT and CONACYT. We appreciate the comments made by J. Cortés, J. Lawrence
and T. Hopkins that improved this paper.
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Chapter 4
Echinoderm Diversity in Panama:
144 Years of Research Across the Isthmus
Simon E. Coppard and Juan José Alvarado
4.1 Introduction
The Republic of Panama is the southernmost country in Central America between

07°000 –10°000 north and 77°000 –83°000 west and is located between Costa Rica to
the north and Colombia to the south. The country runs from east to west with a
length of 772 km and a width that varies from 60 to 177 km (Meditz and Hanratty
1987). Panama has both Caribbean and Pacific coastlines that total some 2,857 km
and which incorporate a vast diversity of marine habitats and ecosystems. The
80 km Panama Canal bisects the country and although this is theoretically a
freshwater barrier, it provides a link between the Atlantic and Pacific Oceans.
Approximately one hundred and fifty rivers flow into the Caribbean, while over
three hundred rivers flow into the eastern Pacific.
Panama’s weather is governed by the Inter-Tropical Convergence Zone which
dictates the seasonal patterns of wind and rainfall in the region. The dry season
(mid-December to mid-April) is characterised by low precipitation but strong
winds as a result of the northeast trade winds (Forsbergh 1969; Amador et al.
2006). The rainy season (May to November) has light winds with heavy rainfall
S. E. Coppard (&)
Smithsonian Tropical Research Institute, PO BOX 0843-03092Balboa, Ancón,
Panamá
e-mail: CoppardS@si.edu
J. J. Alvarado
Universidad de Costa Rica, San Pedro, 11501-2060, San José, Costa Rica
e-mail: juanalva76@yahoo.com
J. J. Alvarado
Posgrado en Ciencias Marinas y Costeras,
Universidad Autónoma de Baja California Sur, La Paz, México

108 S. E. Coppard and J. J. Alvarado
which varies regionally from less than 1.3 m to more than 3 m per year, with
rainfall on the Caribbean side typically being heavier than on the Pacific side of the
continental divide (Meditz and Hanratty 1987).
The formation of the Isthmus of Panama is considered to be one of the most
important geologic events of the past 60 million years as it had an enormous
impact on Earth’s climate and environment. It occurred due to the gradual collision
of the Pacific, Cocos, Nazca and Caribbean plates which led to uplift of the sea
floor and the formation of volcanic islands. Over the next 12 million years vast
quantities of sediment from North and South America were deposited in between
the newly emerging islands, with final closure of the straits occurring in the late
Pliocene, 2.8 million years ago (mya) (Coates et al. 2005). This formation blocked
the North Equatorial Current which had previously flowed westward through the
Central American Seaway and forced the current northwards forming the Gulf
Stream of today (Berggren and Hollister 1974; Burton et al. 1997). With the
Pacific and Atlantic no longer mixing, there were significant changes in temper-
ature, salinity and primary productivity. This resulted in both species extinctions
and species radiations as environments changed (Jackson et al. 1993). Sister
species that were isolated on either side of the isthmus and that subsequently
diverged are referred to as geminates (Jordan 1908). These represent initially
similar genomes that were isolated and diverged over the last 3 million years
(Lessios 2008) and provide insights into evolutionary adaptations in response to
the changing environments.
Today the Caribbean coast of Panama incorporates coral reefs, seagrass beds,
rocky shores, mud-flats and mangroves and has a narrow continental shelf.
It continues to be influenced by the Atlantic North Equatorial Current which flows
through the Lesser Antilles (Kinder et al. 1985). The coast of the south-western
Caribbean including Panama is also affected by a cyclonic gyre that develops from
this current (Gordon 1967; Kinder et al. 1985; Gómez et al. 2005). For an in-depth
review of the Caribbean habitats of Panama, see Gómez et al. (2005). The
Caribbean experiences very small tidal ranges (less than 0.5 m) and has relatively
stable conditions. Its waters are clear and nutrient poor (D’Croz and Robertson
1997; D’Croz et al. 2005; Collin et al. 2009) and unlike the north coast of
Colombia there is no upwelling. These conditions are suitable for growth of
extensive coral reefs which date back to the early Pleistocene (Glynn 1982; Cortés
1993). Surface waters of the Caribbean have an upper 200 m layer of warm water
heated by solar radiation, which has a correspondingly high salinity. Surface water
temperatures vary from 27 °C in the dry season to 30 °C in the rainy season, while
salinity varies from 36 psu in the dry season to 32 psu in the rainy season (D’Croz
and Robertson 1997). Two archipelagos are present off the Caribbean coast of
Panama, the San Blas (Kuna Yala) Archipelago which has a very narrow conti-
nental shelf and is subject to strong ocean influence with an oligotrophic coastal
area (D’Croz and Robertson 1997) and the Bocas del Toro Archipelago, with a
larger continental shelf, and greater influence from continental run-off (D’Croz
et al. 2005). Five national parks have been created along the Caribbean coast of
4 Echinoderm Diversity in Panama 109
Fig. 4.1 Distribution of coral reefs and mangrove forests along the coasts of Panama (adapted
from Gómez et al. 2005). Black stars: coral reefs; grey shading: mangroves
Panama: Bastimentos Island National Marine Park, Portobelo, Soberania, Galeta,

Chagres, and the wetlands of San San Pond Sank (Gómez et al. 2005).
The Pacific coast of Panama is comprised of two gulfs; the Gulf of Chiriquí and
the Gulf of Panama which are separated by the Azuero Peninsula (Fig. 4.1). These
regions experience broad semi-diurnal tides with a range of up to 6 m (Glynn 1972).
Despite their close proximity, they experience very different hydrological condi-
tions. The Gulf of Panama includes the Gulf of San Miguel, the Gulf of Parita and
Panama Bay, as well as the Pearl Islands Archipelago which is a Marine Special
Management Zone (Guzman et al. 2008). The gulf is broad and shallow, with much
of it being less than 40 m deep due to the extension of the continental shelf. The
coastal environments consist of muddy and sandy beaches with extensive mangrove
forests, which receive large amounts of freshwater runoff from rivers such as the Rio
Chepo and Rio Chico as well as the discharge from the Panama Canal. Coral reefs
are only moderately developed and are associated with the Pearl Islands Archi-
pelago and the inshore island of Taboga. Throughout the rainy season the surface
water temperature ranges from 26–28 °C, while salinity is typically below 30 psu
(D’Croz et al. 1991). During the dry season (December to April) the Gulf of Panama
experiences wind-driven, seasonal upwelling of colder, more saline, nutrient-rich
water (Forsberg 1963; Forsbergh 1969; D’Croz et al. 1991; D’Croz and Robertson
1997; Gómez et al. 2005; Pennington et al. 2006), which results in significantly
increased primary productivity. This occurs due to the displacement of the surface
water by northern winds and can result in a surface water temperature decrease of
12 °C within 24 h (Glynn 1972). The Gulf of Panama is also subjected to the effects
of the El Niño-Southern Oscillation (ENSO) every two to seven years (Guzman and
Cortes 2007) resulting in the warming of the water up to 31 °C (Glynn 1985a).
Fig. 4.2 Pocillopora spp. framework on Uva Reef, Gulf of Chiriquí, 2010
In contrast, the Gulf of Chiriquí has a narrower continental shelf and less
freshwater input from continental runoff. It contains Chiriquí National Park and
Coiba Nacional Park which has World Heritage Site status. Several island com-
plexes are present, including the islands of Boca Brava, Los Ladrones, Islas Secas,
Parilla, Isla Palenque, Montuosa, and Isla Jicaron, with Isla Jicarita being the
island furthest south of the Panama mainland. In the northwest of the gulf, water
depth gradually increases down the continental shelf. However, in the east, water
depth sharply drops to 160 m before plunging down the continental slope and
reaching a depth of 1,000 m less than a 1 km south of the Azuero Peninsula. The
Gulf of Chiriquí is shielded from the trade winds out of the North by the Central
Cordillera and therefore experiences no seasonal upwelling. The Gulf of Chiriquí
contains some of the most developed coral reefs on the continental shelf in the
eastern Pacific. These are predominantly formed of a loose Pocillopora spp.
framework (Fig. 4.2) that originated in the Holocene (Glynn 1976; Cortés 1993).
4.2 Research
The first records of the echinoderm fauna of Panama date back to the late 19th and
early 20th century and are the result of the expeditions of the U.S. Fish Commission
Steamer Albatross. Addison Emery Verrill (Fig. 4.3a) wrote the first paper in 1867
that included echinoderms from Panama. It contained descriptions of many new
genera and species. In 1891 Alexander Agassiz (Fig. 4.3b), onboard the Albatross
collected specimens from the intermediate depths of the eastern Pacific and the
Caribbean Sea to study the relationship between forms of marine life on either side
Fig. 4.3 a Addison Emery Verrill, b Alexander Agassiz, c Christian Frederik Lütken,
d Theodore Mortensen (photos courtesy of the Smithsonian National Museum of Natural History)
of the Isthmus of Panama. The collections from this expedition were used by many
authors to describe new echinoderm species (e.g. Agassiz 1892, 1898, 1904;
Ludwig 1894, 1905; Lütken and Mortensen 1899, (Fig. 4.3c, d), Clark 1917).
In the 1930’s and 1940’s expeditions to the eastern Pacific were undertaken by
the New York Zoological Society onboard the Arcturus 1925 and Zaca
1937–1938, and by the Allan Hancock Foundation onboard the Velero III
1937–1941. As a result of these expeditions, a significant amount of echinoderm
research was published (e.g. Clark 1939; Clark 1940, 1948; Ziesenhenne 1940,
1942, 1955; Deichmann 1941, 1958). In 1915 Mortensen spent several months
collecting echinoderms and other invertebrates in the Gulf of Panama, including
from St. Elmo in the Pearl Islands Archipelago and the islands of Taboga and
Taboguilla in Panama Bay. His findings were later included in a broader work on
larval development (Mortensen 1921) and as part of his monumental monograph
on echinoids (Mortensen 1928, 1935, 1940, 1943a, b, 1948a, b, 1950, 1951). Other
researchers in the 1920’s, 30’s and 40’s made collection trips to specific regions in
Panama (e.g. Boone 1928, Clark 1946) focusing more on specific taxa. In later
years echinoderms were collected from the Caribbean Sea off Panama as part of
general benthic sampling expeditions undertaken by the RV Oregon in 1962, the
RV Pelican in 1963, the RV Oregon II in 1970, the RV Pillsbury in 1971 and the
RV Alpha Helix in 1977.
The Smithsonian Tropical Research Institute (STRI) established its first field
station on Barro Colorado Island in the Panama Canal Zone in 1923. The marine
program at STRI began fifty-one years ago, in 1961. Today STRI runs two field
stations on the Caribbean Coast (Galeta Marine Laboratory in Colón and Bocas del
Toro Research Station) and two in the Pacific (Naos Island Laboratories complex and
a Field Station on Ranchería Island, Coibita). The first research vessel was acquired
by STRI in 1970, the 65-ft RV Tethys (1970–1972), followed by the 45-foot RV
Stenella in 1972 (1972–1978) and the 63-foot RV Benjamin in 1978 (1978–1994).
These research vessels allowed STRI scientists to collect and document the shallow
water echinoderm fauna. In 1994 STRI purchased the 96-foot RV Urraca
(1994–2008) that had an A-frame and oceanographic winch which finally provided
the means to trawl and dredge to depths of 250 m. In 1995 the submersible DSRV
Johnson Sea Link was used to collected samples from the Gulf of Chiriquí at depths
over 1,000 m, providing a further insight into the deep water species present off
Panama’s Pacific coast. STRI researchers who worked on echinoderms in the 70’s
and 80’s included Peter Glynn, Gordon Hendler and Harilaos Lessios. For just
over 30 years Lessios has worked at STRI on the life histories and evolution of
echinoderms in Panama. His research has focused primarily on echinoids, including
work on speciation, molecular biogeography and the evolution of reproductive
isolation. Lessios (2005a) published a review of the echinoids of the Pacific waters of
Panama, providing information on the species and new records of their distributions.
4.3 Ecology
Ecological research in Panama has included work on all classes of echinoderms,

but with fewer studies on crinoids. Eleven species of echinoderms were recorded
in the seagrass meadows of Thalassia testudinum Banks and Sol. ex K.D. Koenig
(1805) in the Canal Zone off the Caribbean coast of Panama (Heck 1977). Species
richness of echinoderms correlated with proximity to well developed reefs and was
found to be stable throughout the study period (July 1974 to May 1975), without
any major decrease in population numbers during periods of reduced salinity.
Fig. 4.4 a Echinometra lucunter, b Echinometra viridis at Galeta Point Panama
Mortality of sea urchins on the fringing reef-flat off Galeta Island has been
reported to occur twice yearly as the result of physical stress generated by
extended aerial exposure of the reef platform (Glynn 1968; Hendler 1977; Cubit
et al. 1986). Tripneustes ventricosus (Lamarck, 1816), Lytechinus variegatus
(Lamarck, 1816) and Diadema antillarum Philippi, 1845 were found to be less
tolerant to high temperature than either Echinometra lucunter (Linnaeus, 1758)
(Fig. 4.4a) or Echinometra viridis Agassiz, 1863 (Fig. 4.4b), and are therefore
more likely to die during the periods of exposure.
In the Caribbean D. antillarum has been shown to be the dominant herbivore on
coral reefs that have been subjected to heavy human fishing pressure, while
herbivorous fishes dominate in more pristine conditions where urchin populations
are kept in check by fish predators (Hay et al. 1983; Harborne et al. 2009). Off
Galeta Island, where human fishing pressure is reported to be low (Hay 1984),
almost all grazing on bioassays of T. testudinum was attributed to herbivorous
fishes (Hay 1984). Off the San Blas Islands of Panama, the grazing activity of
D. antillarum has been shown to be the primary determinant of fleshy algal
biomass on shallow reefs (\3 m) (Foster 1987a). At the time of Foster’s study
(pre-mass mortality of D. antillarum, see Sect. 4.6) this was particularly true in the
back reef environment, where D. antillarum occurred at densities three times
higher (*6 ind m-2) than in Elkhorn coral habitats (\2 ind m-2) (Foster 1987a).
Chemical defence by coral reef macroalgae has been shown to have an import role
in deterring grazers (Hay et al. 1987). Extracted terpenoid compounds (naturally
occurring in macroalgal species of Dictyota, Dilophus, Laurencia and Stypopodium)
were applied to assays of T. testudinum and shown to deter grazing by herbivorous
fishes and D. antillarum at Galeta (Hay et al. 1987). However, Cymopol, a terpenoid
bromohydroquinone from the green alga Cymopolia barbata (Linnaeus) Lamouroux,
1816, significantly reduced feeding by reef fishes but significantly stimulated feeding
in D. antillarum.
The ability of the damselfish Stegastes dorsopunicans (Poey, 1868) to exclude
and therefore reduce the grazing impacts of D. antillarum from their territories was
studied by Foster (1987b) in San Blas. Despite the major negative impact of
D. antillarum on the algal biomass in damselfish territories, this damselfish species

was found to rarely defend its feeding areas against D. antillarum and therefore
had little effect on its distribution.
On the Pacific coast of Panama grazing by D. mexicanum results in the
bioerosion of the predominantly Pocillopora spp. coral reefs (Glynn 1988).
This species contributes 78 % of the overall bioerosion 13 kg m-2 y-1 produced
by benthic eroders (Glynn 1988; Eakin 1991). Analysis of faecal pellets has
revealed that the diet of D. mexicanum consists of 75.2 ± 11.03 % coral and
only 23.0 ± 11.4 % of crustose coralline algae (CCA). In contrast the diet of
Toxopneustes roseus (A. Agassiz, 1863) is composed of approximately 20 %
coral and 80 % of CCA (Glynn 1988). This highlights the potential damage
D. mexicanum poses to reef structures, particularly when present in large numbers.
Such a threat was realised following the 1982–83 El Niño Southern Oscillation
event (ENSO). This resulted in mass coral bleaching and a 50 % reduction of live
coral cover on Uva Reef (Glynn 1985a), which was followed by a dramatic
increase in the population size of D. mexicanum from 2–5 ind m-2 pre 1983 to
60–100 ind m-2 post 1982-83 ENSO (Glynn 1988, 1990; Eakin 1991, 1992). Such
large numbers of D. mexicanum resulted in high rates of reef framework erosion,
shifting the reef environment into a state of near accretionary stasis (Eakin 1992;
Glynn 1997). Low predation pressure (with the removal of potentially important
fish predators as a result of over-fishing), increased habitat availability, and the
increased abundance of benthic algae as a result of the newly dead coral sub-
stratum allowed D. mexicanum to attain high population densities (Glynn 1988).
However, such a large population of D. mexicanum could not be sustained and
in the year 2000 the population of D. mexicanum dropped to near 1974 levels
([10 ind m-2) (Eakin 2001).
The ability of damselfish in the eastern Pacific to exclude Diadema from their
territories appears to be a very different situation to that reported in the Caribbean.
The territorial action of Stegastes acapulcoensis (Fowler, 1944) on Uva Reef
results in few sea urchins being present in their territories (Glynn 1990), with
consequently lower levels of erosion (6.3 mm y-1) than elsewhere on the reef
(21.8 mm y-1) (Eakin 1992).
Ecological studies on irregular echinoids in Panama have focused on sand
dollars. Seilacher (1979) studied the distribution of Mellita quinquiesperforata
(Leske, 1778) (as Mellita lata Clark, 1940), a subjective junior synonym of M.
quinquiesperforata) at Maria Chiquita on the Caribbean coast of Panama. This
species lives partially buried in the sand (with some of the apical surface exposed)
and is restricted to a 4 m wide region of the surf-zone that runs parallel to the
beach (Seilacher 1979). This species was observed feeding along this zone, but
migrated to deeper water when the intensity of the wave action increased.
The behavioural ecology of a population of Mellitella stokesii (L. Agassiz,
1841) (as Encope stokesi L. Agassiz, 1841 [sic]) was studied over a year at Venado
Beach, on the eastern Pacific coast of Panama by Dexter (1977). Densities of this
species were reported to vary from a low of 19 ind m-2 in the rainy season to a
Fig. 4.5 Mellitella stokesii

at Venado beach, a aboral
view, b feeding during low
tide
high of 65 ind m-2 in the dry season, with numbers concentrated in the mid-
intertidal zone where the median sediment grain size was between 170 and
200 lm. No sand dollars were found in the muddy sand bar which had a high silt
and clay fraction. Migrations towards the substrate surface were observed at low
tide which correlated with the appearance of an organic film on the substrate,
composed of a combination of diatoms and fine detrital material. Observations by
one of the authors (SEC) at Venado Beach (see Fig. 4.5a, b) agree with Dexter’s
findings. During high tide this species typically remains buried in the substrate,
perhaps to avoid predation, and only migrates to the surface at low tide, where it
moves through the upper sediment layer in the shallow tide pools ingesting small
particulate mater (Fig. 4.3b). Water temperatures in these pools regularly gets in
excess of 40 °C (Coppard unpublished data), demonstrating that M. stokesii has
a tolerance for high water temperatures. Dexter (1977) reports that this species has
a high growth rate, in conjunction with a small maximum size (she recorded a
maximum horizontal diameter of 57 mm) and a short life expectancy of typically
Fig. 4.6 Mellitella stokesii

(oral views) on Venado beach
with Dissodactylus spp.,
a multiple wounds, b a single
wound
less than 1 year. Recruitment of the population was reported to occur throughout
the year, but with a peak in settlement during February, in the middle of the
Panamanian dry season. This is the season of increased food availability for
planktotrophic larvae, due to the upwelling of nutrients and subsequent increase in
phytoplankton (Martin et al. 1970).
Two species of pinnotherid crab (pea crab), Dissodactylus nitidis Smith, 1870
and Dissodactylus xantusi Glassell, 1936 were found associated with M. stokesii
(Dexter 1977). These species remove the spines near the lunules or marginal slits
on the oral surface, creating a wound in which they live (see Fig. 4.6a, b). Dexter
questioned whether this relationship is really commensal, as she found that these
crabs in aquaria removed larger numbers of spines on the oral surface, which
decreased the ability of the sand dollar to feed and move, ultimately resulting in
the death of the host.
Telford (1982) described the feeding habits of four species of Dissodactylus and
found that all of them feed extensively upon their sand dollar hosts, with
50–100 % of their food intake coming from their host’s tissues. This is supported
by the fact that some M. stokesii often have multiple wounds with only a single
crab present (Fig. 4.6a).
Ecological work on ophiuroids in Panama is primarily the result of research by
Gordon Hendler. Hendler and Meyer (1982) reported the presence of the poly-
chaete Branchiosyllis exilis (Gravier, 1900) associated with Ophiocoma echinata
(Lamarck, 1816) on the Caribbean coast of Panama. This association has only been
reported from Panama, despite the fact that both species’ distributions are over-
lapping and widespread. The polychaete displays active recognition of its host and
seeks out O. echinata in preference to other species of the same genus. They
consistently found one polychaete per host, indicating that B. exilis is aggressive to
member of its own species, and concluded that B. exilis is parasitic on O. echinata.
An interesting association between juvenile and adult Ophiocoma aethiops
Lütken, 1859, from Punta Paitilla in Panama was documented by Hendler et al.
(1999). Juveniles were found in the bursae (respiratory structures that in brooding
and viviparous ophiuroids also serve as brood chambers) of adults that live
intertidally at Punta Paitilla but not those that live subtidally at Isla Taboguilla.
They proposed this association helped protect juveniles against desiccation and
predators. No similar association was found in other species of Ophiocoma form
either the Pacific (O. alexandri Lyman, 1860) or from the Caribbean (O. echinata,
O. wendtii Müller and Troschel, 1842) of Panama.
Much of the ecological work on starfish in Panama has focused on Acanthaster
planci (Linnaeus, 1758) (e.g. Glynn 1973, 1974, 1976, 1977, 1981– 1985a,b,
1990). Glynn (1973) assessed the possible effect of A. planci on the coral species
on Uva Reef in the Gulf of Chiriquí. He reported an average density of 25–36 ind
ha-1, with a disc diameter of 17–19 cm and a size frequency that had a unimodal
distribution. Population densities of A. planci remained fairly stable on Uva Reef
from 1970 to 1980, ranging from 7 to 30 ind ha-1 (Glynn 1981). At that time these
densities were comparable to population sizes in the Indo-Pacific that were not
considered to have a serious impact on coral communities (Glynn 1974). However,
prey preference data (Glynn 1974) indicate that A. planci selectively eats rarer
non-branching corals (e.g. species of Pavona, Gardineroseris, Porites, Millepora,
Fig. 4.7), which are replaced by faster-growing species (e.g. Pocillopora spp.),
resulting in A. planci having a negative effect on species diversity (Glynn 1976).
Such a preference can partly be explained by the presence of crustacean symbionts
(Trapezia sp. and Alpheus sp.) in large branching pocilloporid colonies which
repulse A. planci by nipping at its vulnerable oral surface (particularly the sensory
tube feet) as it attempts to mount the coral to feed (Glynn 1976).
Between 1982 and 1983 a strong El Niño-Southern Oscillation had a dramatic
impact on the coral reefs of the Eastern Tropical Pacific, causing a die off of
between 70–95 % of coral on Pacific Panamanian reefs (Glynn 1985a, b). How-
ever, population densities of A. planci before and after the 1983 event were not
statistically different (Glynn 1990), with A. planci continuing to feed on the
Fig. 4.7 Acanthaster planci

feeding on Millepora
intricata Milne-Edwards and
Haime, 1860, Coiba National
Park, Gulf of Chiriquí (photo
courtesy of Angel Chiriboga)
dispersed, surviving coral. As the reefs recovered the relative effect of predation by
A. planci intensified due to the loss of the crustacean guards (Trapezia and
Alpheus), therefore reducing local coral species diversity (Glynn 1985b).
Fong and Glynn (1998) modelled the impacts of predation by A. planci and that
of the El Niño-Southern Oscillation (ENSO) on the structure of a population of the
massive coral Gardineroseris planulata (Dana, 1846) on Uva Reef. The results of
the simulations suggested that predation was far more important than the ENSO in
controlling the size structure of the coral G. planulata in the eastern Pacific.
Although A. planci has few predators, it has been observed in Panama being
attacked by the Harlequin shrimp Hymenocera picta Dana, 1852 and by the
polychaete Pherecardia striata (Kingberg, 1857) (Glynn 1981). The shrimp
attacks the aboral surface of the starfish, eventually amputating the arms and using
its chelae to expose and feed on the internal soft parts. However, the harlequin
shrimp typically prefers other starfish species such as Phataria unifacialis (Gray,
1840), Pharia piramidatus (Gray, 1840) and Nidorellia armata (Gray, 1840)
(Glynn 1977). The wounds created by the harlequin shrimp are used also by the
polychaete P. striata to enter A. planci and feed on the internal tissues. Glynn
(1981) calculated that 5–6 % of the A. planci population is under attack by
H. picta and about half of these starfish are killed over a 3-week period.
Guzman and Guevara (2002a) studied the spatial distribution, abundance, and
size structure of Oreaster reticulatus (Linnaeus, 1758) (Fig. 4.8) in the Bocas del
Toro Archipelago. They found a mean density of 149.7 ind ha-1 and estimated the
population size to be over seven million. Approximately 45 % of the population
was found at a density of 308.3 ± 50.9 ind ha-1, in areas dominated by the
Fig. 4.8 Oreaster reticulatus

in the Thalassia testudinum
seagrass bed, Bocas del Toro
(photo courtesy of Edgardo
Ochoa)
seagrass T. testudinum, with a substratum that consisted of coarse, calcareous sand.

The lowest density (5.6 ind ha-1) was observed in coral reef habitats.
Feeding patterns of O. reticulatus off the San Blas Islands were studied by
Wulff (1995). She found that 61.4 % of the starfish were eating sponges; however,
this depended on their availability. Oreaster reticulatus fed upon sixteen of twenty
species that normally grow only on the reefs, but surprisingly only one of fourteen
species that live in the seagrass meadows and rubble flats surrounding the reefs.
Therefore O. reticulatus actively searches for and consumes edible sponges that
wash into the seagrass beds from reefs, or off mangrove roots (Wulff 1995, 2006).
Meyer (1973) described the feeding behaviour, microgeographic distribution,
population size, bathymetric distribution and adaptive skeletal morphology of
eight species of Caribbean crinoids from four main study areas, including San Blas
and Galeta, in Panama. Based on his findings he divided the species into two
categories: 1) those forming a filtration fan (Nemaster grandis Clark, 1909,
Tropiometra carinata (Lamarck, 1816), Comactinia echinoptera (Müller, 1840),
Analcidometra caribbea (A. H. Clark, 1908) and Neocomatella pulchella (Pour-
talès, 1878)) and 2) those with a radian posture (Davidaster rubiginosus (Pourtalès,
1869), Davidaster discoidea (Carpenter, 1888) and Ctenantedon kinziei Meyer,
1972). Species diversity and population size of shallow-water comatulids was
found to be greatest in Panama (seven species). Meyer (1973) proposed that the
diversity and abundance of comatulid crinoids can be correlated to increased
primary productivity which occurs close to continental land masses.
Ecological studies on holothuroids in Panama have been carried out by Guzman
and Guevara (2002b). They studied the population structure, distribution and
abundance of the sea cucumbers Isostichopus badionotus (Selenka, 1867),
Holothuria (Halodeima) mexicana Ludwig, 1875 and Astichopus multifidus
(Sluter, 1910) in the Bocas del Toro Archipelago. Holothuria (Halodeima)
mexicana was found to be more abundant in seagrass, I. baniodotus occurred at
higher densities in interspersed areas, while A. multifudus showed no specific
habitat preference. For the Bocas del Toro Archipelago Guzman and Guevara
(2002b) estimated populations of 7,630,164 individuals for H. (H.) mexicana,

5,536,349 for I. badionotus, and 231,074 for A. multifidus. However, the densities
of these sea cucumbers in Panama were 50 times lower than those measured at
other Caribbean sites (Guzman and Guevara 2002b).
4.3.1 Physiology
Lawrence and Glynn (1984) studied the diet of the sea urchin Eucidaris thouarsii
(Valenciennes, 1846) through the absorption of nutrients from the predation on the
coral Pocillopora damicornis (Linnaeus, 1758) in the Gulf of Panama. The
absorption efficiency was determined to be 44.7 % for carbohydrates, 85.3 % for
protein and 69.3 % for lipids. The urchin absorbs 0.402 mg carbohydrate,
2.814 mg protein and 1.108 mg lipid for each gram of P. damicornis ingested;
0.201 mg carbohydrate, 1.407 mg protein and 0.554 mg lipid per individual per
day; and absorbs 0.824 9 10-3 kcal carbohydrate, 7.950 9 10-3 kcal protein and
235 9 10-3 kcal lipid per individual per day. The authors concluded that despite
the high efficiency of digestive absorption of organic constituents of the coral
ingested, E. thouarsii needs to consume large quantities to meet its nutritional
requirements.
Work by Hendler and Byrne (1987) on Ophiocoma wendtii collected in Galeta,
Panama Caribbean, Belize and Florida, resulted in the identification and descrip-
tion of three structures in the dorsal arm plate (DAP) that comprise a photoreceptor
system. The upper surface of the DAP bears transparent microscopic structures
which are expanded peripheral trabeculae of the calcite stereom. These structure
are part of the photoreceptor system and can facilitate light transmission through
the DAP by decreasing light refraction, reflection and absorption that occur at
stereom/stroma interfaces. Bundles of nerve fibres located below the expanded
peripheral trabeculae function as primary photoreceptors. The third elements of the
system are the chromatophores that regulate the intensity of light impinging on the
sensory tissue are regulated by the diurnal activity cycle. During the day
the chromatophores cover the expanded peripheral trabeculae and thereby shade
the nerve fibres. At night they retract into inter-trabecular channels, uncovering the
expanded peripheral trabeculae and thereby exposing the nerve fibres to trans-
mitted light. Thus, the transparent stereom may play a role in photoreception, in
addition to its generally recognized skeletal function. Although ciliated cells that
may be sensory are present in the epidermis of O. wendti, they do not appear to be
photoreceptors (Hendler and Byrne 1987).
4.3.2 Reproduction
Reproductive cycles of echinoids on both sides of the Panama Isthmus have been
studied by Lessios (1981a, 1984, 1985a, 1991). Populations of D. mexicanum and
E. vanbrunti from Punta Paitilla, Isla Urabá and Culebra Island in the seasonal
Gulf of Panama have well-defined, synchronous, reproductive cycles (Lessios
1981a). The majority of individuals reached a peak gonad size in September, with
spawning predicted to occur from September to October. This would allow the
newly metamorphosed sea urchins, rather than the larvae, to benefit from the
increased food production as a result of the dry season upwelling (Lessios 1981a).
Populations of D. antillarum and E. lucunter from Maria Chiquita and Fort
Randolph in the less seasonal Caribbean demonstrated only slight, if any syn-
chrony between individuals. However, E. viridis which is rarer at these locations
had a well defined, population wide cycle. Based on these data Lessios (1981a)
proposed that in a constant environment the intensity of selection for synchrony
between individual gametogenic cycles may be inversely proportional to popula-
tion density. Later work by Lessios (1985a) on Caribbean echinoids did not
directly support this hypothesis. However, Lessios (1985a) suggested that popu-
lation density may be one of the factors determining the degree of synchronicity.
This study (1985a) revealed that E. viridis, L. williamsi, Clypeaster rosaceus
(Linnaeus, 1758) and Leodia sexiesperforata (Leske, 1778) are reproductively
active during Panama’s rainy season from May to November, but quiescent during
the dry season (Lessios 1985a), while L. variegatus, T. ventricosus, and Clypeaster
subdepressus (Gray, 1825) have ripe gonads throughout the year. Only E. lucunter
showed non-periodic fluctuations in readiness to spawn.
Lessios (1984, 1991) also worked on lunar spawning periodicity in Panamanian
echinoids. Diadema mexicanum was found to spawn on the third quarter of the
lunar cycle, with a peak three to four days after full moon, while its allopatric
congener D. antillarum, spawned during the first lunar quarter, one to two days
after new moon. Lessios (1984) wrote that it is therefore possible that geminate
species of Diadema have acquired a prezygotic isolating mechanism while in
allopatry and therefore reproductive isolation need not be the product of natural
selection against hybridization, thus casting doubt on the speciation by rein-
forcement theory.
Lunar spawning rhythms were not found in T. ventricosus, E. viridis,
L. sexiesperforata, C. rosaceus or L. williamsi (Lessios 1991). Eucidaris tribu-
loides and D. antillarum were found to have a distinct lunar rhythm spawning at
different lunar phases, while L. variegatus has a semilunar rhythm spawning
every new and full moon (Lessios 1991). Such spawning rhythms are unlikely to
be cued by tides as the Atlantic coast experiences only small tidal ranges
(\0.5 m) where water level is determined as much by meteorological conditions
as by tides. Lunar periodicity was therefore shown to not be a lineage specific
trait inherited from a common ancestor, but has evolved independently in dif-
ferent echinoid taxa (Lessios 1991).
Temporal and spatial variation in egg size of thirteen Panamanian echinoids was
studied by Lessios (1987). His results showed that mean egg size from different
females collected from the same locality on the same day were significantly different
in all species studied. However, daily (within-month) variation in egg size was only
significant in E. tribuloides, D. antillarum, and L. variegatus, while no significant
differences were found in the daily means of egg volume in D. mexicanum and E.
viridis. Monthly means of egg volume were different in L. variegatus, L. williamsi,
T. ventricosus, E. viridis, E. vanbrunti, C. rosaceus and C. subdepressus. Between-
years variation was also significant in all of these species except L. variegatus. All
Caribbean species showed a decline in egg size after September, however, this could
not be explained by any obvious environmental fluctuation. Echinometra lucunter
showed no significant variation between months of the same year but exhibited
differences between years. No significant monthly or annual variation was observed
in E. tribuloides, E. thousarii (Valenciennes, 1846), D. antillarum, D. mexicanum or
Leodia sexiesperforata. Correlations between size of eggs collected at a particular
time and the intensity of spawning by the population at that time was not significant,
suggesting that size of mature eggs is not determined by the reproductive state of the
parental population (Lessios 1987).
Lessios (1990) looked at adaptation and phylogeny as determinants of egg size
in twenty-two species of echinoids and two species of asteroids from the Carib-
bean and Pacific coasts of Panama. Lessios (1990) identified four trends: (1)
species have eggs that are significantly different from those of other congeneric
species; (2) species evolving for three million years in separate environments have
accumulated differences in their egg sizes and the direction of these differences is
not random; (3) egg size of congeneric species is similar where developmental
modes are similar (n.b. the volume of Clypeaster rosaceus eggs is eight times
greater than that of its congener C. subdepressus, an adaptation that reflects
developmental mode as the larvae of C. rosaceus are facultative planktotrophs,
while those of C. subdepressus are obligate planktotrophs); (4) there is no simi-
larity between the egg sizes of species belonging to the same family or higher
taxonomic category, even if these species inhabit the same ocean. He concluded
that because the phenomena of smaller eggs in the Pacific (a pattern attributed to
higher levels of primary productivity in the eastern Pacific) holds for lineages that
have not shared a common ancestor for a long time, it is likely that independent
adaptation by the members of each pair has occurred due to the environments of
the two oceans during the last three million years.
McAlister (2008) studied the plasticity of larval feeding arms for geminate
species pairs to determine how food availability can influence growth and the arm
length of the larvae. No plasticity in arm length was found in either Caribbean or
Pacific species, however, arm length was significantly longer in Caribbbean spe-
cies (including in D. antillarum). McAlister (2008) suggests that historical changes
in food levels led to the development of longer feeding arms in Caribbean species,
while plasticity may be limited by egg size or the timing of reproduction.
The evolution of gametic incompatibility in neotropical Echinometra was
investigated by Lessios and Cunningham (1993). They used fertilization
experiments to determine the degree of reproductive isolation both between the

sympatric E. lucunter and E. viridis, in the Caribbean, and with the allopatric
E. vanbrunti from the eastern Pacific. They reported that, contrary to the predic-
tions of the ‘speciation by reinforcement hypothesis’, the degree of incompatibility
between the allopatric E. lucunter and E. vanbrunti is higher than between the
sympatric E. lucunter and E. viridis. Crosses between E. viridis and E. vanbrunti
produced rates of fertilization almost equal to those of homogamic crosses, while
sperm of E. lucunter were found to be able to fertilize eggs of the other two
species. However, few E. lucunter eggs are fertilized by heterospecific sperm
(Lessios and Cunningham 1993). In the wild E. lucunter and E. viridis maintain
their genetic integrities despite having this unidirectional gamete isolation.
Allozyme data placed the speciation event of the sympatric species after the rise of
the isthmus. Thus there is no correlation between genetic divergence and strength
of reproductive isolation (Lessios and Cunningham 1993).
Later work by McCartney and Lessios (2002) confirmed these findings and
demonstrated a much stronger block to cross-species fertilization of E. lucunter
eggs than was previously shown. They reported that gamete incompatibility in
these species is weaker in sympatry than in allopatry, and based on the mtDNA
data in McCartney et al. (2000), such gamete incompatibility in these species arose
in the last 1.5 million years.
Zigler et al. (2008) looked at egg energetics, fertilization kinetics, and popu-
lation structure in echinoids with facultatively feeding larvae. This work included
C. rosaceus, one of only two echinoid species known to have facultatively
planktotrophic larvae. The eggs of C. rosaceus resemble those of echinoid species
with obligately feeding larvae in egg energetic density, egg buoyancy and fertil-
ization kinetics, but resemble non feeding larvae in time to metamorphosis, the
early formation of the coelom and the reduced allocation to larval feeding struc-
tures (Zigler et al. 2008).
Cross-fertilization experiments between C. rosaceus and C. subdepressus by
Zigler et al. (2008) revealed that bidirectional gametic incompatibility has evolved
between the two species. These species are commonly found together or within close
proximity to one another in the shallow waters of the Caribbean in Panama (both
species live in the sand, but C. rosaceus is typically more common in turtle grass
beds). These species have no temporal reproductive isolation, as both have ripe
gonads for much of the year, but with peaks from July to October (Lessios 1984).
The results of Zigler et al. (2008) indicate that the small eggs of C. subdepressus
are particularly resistant to fertilization by C. rosaceus sperm, while the larger
C. rosaceus eggs are slightly more vulnerable to fertilization by C. subdepressus
sperm, but at levels that are unlikely to results in conspecific fertilizations in nature.
Research on the sea urchin sperm protein bindin in sea urchin species found off
the coasts of Panama has been conducted by Zigler and Lessios (2003a, b, 2004),
McCartney and Lessios (2004), Zigler et al. (2005), and Geyer and Lessios (2009).
Bindin is a sea urchin sperm protein that mediates sperm-egg attachment and
membrane fusion. Across echinoid genera, its divergence has been shown to be
correlated with heterospecific incompatibility in fertilization (Zigler et al. 2005).
In sympatric species, a region of the bindin molecule reportedly evolves at high

rates under strong positive selection (Metz and Palumbi 1996), whereas in allo-
patric species (e.g. T. ventricosus in the Caribbean and T. depressus in the eastern
Pacific), bindin variation was reported to be low, with no evidence of positive
selection. However, this does not appear to be the case for all sea urchin species.
Lytechinus williamsi and L. variegatus, which have overlapping distributions in
the Caribbean, show no clear evidence of selection (Zigler and Lessios 2004).
While in spatially overlapping E. lucunter and E. viridis, positive selection was
only found to have accumulated along the bindin lineage of E. lucunter
(McCartney and Lessios 2004). Further, Geyer and Lessios (2009) found no evi-
dence to suggest that the source of selection on bindin in E. lucunter is
reinforcement, as bindin showed no evidence of stronger selection in areas of
sympatry relative to areas of allopatry. Geyer and Lessios (2009) propose that the
evolution of this molecule is a result of the processes acting within species, such as
sexual selection and sperm competition.
The reproductive cycle of the starfish O. reticulatus in Isla Solarte, Bocas del
Toro Archipelago, was studied from February 2000 to February 2001 by Guzman
and Guevara (2002a). Males and females with ripe gonads were observed almost
every month, however, the gonad index revealed peaks in reproductive activity,
during July, November, and January. Gonad maturation was not triggered by
changes in water temperature (these ranged from 27 °C to 30 °C), but may have
responded to chemical cues in the water (e.g. nutrients, salinity) (Guzman and
Guevara 2002a).
Guzman et al. (2003) assessed the reproductive status of two commercial
species of sea cucumber, I. badionotus and H. (H.) mexicana in the Caribbean of
Panama. They reported that for both species the minimum reproductive length was
13–20 cm with a weight of 150 g. Both species were gonochoric with a 1:1 sex
ratio. Gametogenesis and spawning occurred throughout the year; however there
were periods of peak reproductive activity, between July and November for
I. badionotus and from February to July for H. (H.) mexicana (Guzman et al.
2003).
4.3.3 Evolution
Molecular work on echinoids in Panama started with Lessios (1979), who used
geminate species of Panama (E. tribuloides and E. thouarsii, D. antillarum and D.
mexicanum, E. viridis, E. lucunter and E. vanbrunti) to test the molecular clock
hypothesis. This hypothesis predicts that each protein changes at a constant rate, so
that the degree of divergence between two species is linearly related to the time for
which their lineages have remained separate (Wilson et al. 1977). Lessios (1979)
found that this does not hold for Panamanian echinoids, as very different rates of
divergence were found in the enzymatic proteins between geminates in the
different genera. Eucidaris and Echinometra exhibited transisthmian distances
sixteen and thirty-seven times greater than intraspecific ones respectively, while
the trans-isthmian distance in Diadema was twenty times smaller than for the
Echinometra. Lessios (1979) proposed that the most plausible explanation was that
the molecules used in this investigation were evolving under the influence of
natural selection.
In 1981(b) Lessios combined this molecular work with morphometric data
using twenty morphological measurements. The morphometric data showed that
differentiation between members of each geminate pair are not significantly dif-
ferent from local variation within each species. However, the ratio of inter to
intraspecific Mahalanobis distance was congruent with the molecular data, being
lowest in Diadema, intermediate in Eucidaris, and highest in Echinometra.
Morphological rates of divergence were found to be more rapid in sympatric
species, possibly due to habitat separation between congeners (Lessios 1981b).
Bermingham and Lessios (1993) used mitochondrial DNA (mtDNA) and
isozymes in these sea urchin species to measure the degree of divergence. Trans-
isthmian isozyme divergence in the geminate pairs differed by an order of mag-
nitute, while mtDNA divergence was equivalent in all pairs demonstrating that
mtDNA can provide accurate estimates of time since separation (Bermingham and
Lessios 1993).
After the mass mortality of D. antillarum in the Caribbean (see Sect. 4.6.
Threats to echinoderms), Lessios (1985b) looked at the genetic consequences of
the restriction in the genetic structure of populations. His results showed no loss of
genetic variability to that recorded in the same populations of D. antillarum in
1977. Lessios suggested that organisms with planktonic larvae are likely to recover
from local extinction events without significant losses of their genetic variability,
as larval exchange allows re-seeding of affected areas by recruits not exposed to
the bottleneck.
A phylogenetic survey of sea urchin retroviral-like (SURL) retrotransposable
elements in 33 species of echinoid (regular sea urchins, sand dollars, and heart
urchins) was conducted by Gonzalez and Lessios (1999). High ratios of synony-
mous to nonsynonymous substitutions suggest that the reverse transcriptase of the
elements is under strong purifying selection (Gonzalez and Lessios 1999). They
report that despite the predominance of vertical transmission, sequence similarity
of 83–94 % for SURL elements from hosts that have been separated for 200 Myr
suggests four cases of apparent horizontal transfer between the ancestors of the
extant echinoid species. In three additional cases, elements with identical RT
sequences were found in sea urchin species separated for a minimum of 3 Myr.
Thus, horizontal transfer plays a role in the evolution of this retrotransposon
family (Gonzalez and Lessios 1999).
Cytochrome oxidase I (COI) divergence was assessed in Atlantic and eastern
Pacific Echinometra by McCartney et al. (2000), using the closure of the Isthmus
of Panama to date cladogenic events. They reported that the Atlantic species
E. lucunter and E. viridis diverged 1.27–1.62 mya at a time in the Pleistocene
when sea levels fell and Caribbean coral speciation and extinction rates were high.
The fact that these species split so recently, yet do not hybridize, demonstrates that
reproductive barriers between marine species can occur in less than 1.6 million
years (McCartney et al. 2000).
A global molecular phylogeny of Diadema (including D. antillarum and
D. mexicanum from Panama) was published by Lessios (2001) and Lessios et al.
(2001). This molecular study showed that Diadema is composed of reciprocally
monophyletic mtDNA clades, and that the eastern Atlantic D. antillarum
(D. antillarum-b) from Madeira, Canary Islands, Cape Verde and Sao Tome is
genetically distinct from the western Atlantic D. antillarum. These species are
separated by the long stretch of deep water separating the eastern from the western
Atlantic. Diadema antillarum ascensionis Mortensen, 1909 from the central
Atlantic islands of Ascension and St. Helena was found to be genetically isolated,
but nested within the Brazilian clade of the western Atlantic D. antillarum. Lessios
suggests that the biogeographic barrier between the Caribbean and Brazil may be
caused by the outflow from the Amazon and Orinoco Rivers. High levels of gene
flow were reported among populations of D. mexicanum in the eastern Pacific,
with D. mexicanum from the Galapagos and Cocos Island belonging to the same
population as sea urchins from Panama and Mexico (Lessios et al. 2001).
A study of the phylogeography of the pantropical sea urchin Tripneustes by
Lessios et al. (2003) showed that based on morphology, COI, and bindin data,
T. depressus A. Agassiz, 1863 from the eastern Pacific is in fact the same species
as T. gratilla (Linnaeus, 1758) from the western Pacific. The formation of the
Isthmus of Panama, the deep water separating the eastern and western Atlantic,
and the freshwater plume of the Orinoco and Amazon rivers between the
Carribbean and the coast of Brazil were shown to be important barriers to the
evolution of Tripneustes. However, the Eastern Pacific Barrier (5,000 km of deep
water) between the central and eastern Pacific was unimportant in the subdivision
in the genus.
A phylogeographic study of the genus Lytechinus by Zigler and Lessios (2004)
using mitochondrial COI, and the entire molecule of nuclear bindin shows that the
genus Lytechinus is paraphyletic (using Toxopneustes and Tripneustes as out-
groups), with Sphaerechinus granularis (Lamarck, 1816) coming out as the sister
species to L. euerces Clark, 1912 . The authors propose that L. euerces should be
moved into the monotypic Sphaerechinus, rather than placing Sphaerechinus in
Lytechinus which would increase the range of the genus to the temperate eastern
Atlantic and Mediterranean (Mortensen 1943a). Lytechinus semituberculatus
(Valenciennes in L. Agassiz, 1846) and L. panamensis formed a clade with no
distinction between the two species, while L. anamesus and L. pictus were also
phylogenetically indistinguishable. A well supported Atlantic clade was formed of
L. williamsi and the three subspecies of L. variegatus (L. variegatus variegatus,
L. variegatus atlanticus, L. variegatus carolinus). Lytechinus williamsi from the
Caribbean was found to share mtDNA haplotypes with L. variegatus variegatus,
however, their bindin was shown to be distinct and to coalesce within each
morphospecies. A small private clade of mtDNA was found in some L. williamsi
from Panama and Belize. Zigler and Lessios (2004) suggest that this may be
indicative of former differentiation in the process of being swamped by intro-

gression, or of recent speciation.
Molecular work on the starfish A. planci, including specimens from Panama,
revealed that it is not in fact a single species, but a pan-Indo-Pacific species
complex consisting of four deeply diverged clades (Pacific, Red Sea, Northern
Indian Ocean, and Southern Indian Ocean) (Vogler et al. 2008). These clades
reportedly diverged between 1.95 and 3.65 mya (Pliocene to early Pleistocene)
and have genetic distances (8.8–10.6 %) equivalent to the distances between other
sibling species of starfish (Waters et al. 2004). Vogler et al. (2008) propose that
this speciation process was driven by sea level changes (Pillans et al. 1998) that
isolated populations.
Zulliger and Lessios (2010) published a global phylogeny of the genus
Astropecten based on 117 specimens belonging to 40 species. This genus is one of
the most species-rich genera among starfish containing over 150 described species
with six species recorded from Panama: A. armatus Gray, 1840, A. articulatus (Say,
1825), A. cingulatus Sladen, 1883, A. exiguus Ludwig, 1905, A. fragilis Verrill,
1870 and A. regalis Gray, 1840. Such diversity is remarkable, because most species
of Astropecten have a long-lived planktotrophic larval stage. Consequently one
would expect this to lead to a low speciation rate (Zulliger and Lessios 2010).
Zulliger and Lessios (2010) compared their molecular phylogeny to Döderlein’s
(1917) morphological phylogeny. They found high levels of congruence on the
whole, but many discrepancies on a local scale. Phylogentic inference from their
data reveals that that morphological and ecological convergence has taken place in
Astropecten, resulting in allopatric non-sister taxa with similar morphologies and
habitat preferences. Varieties of several species were also shown to exhibit genetic
distances large enough to justify recognizing them as separate species.
Reviews on speciation in organisms separated by the Isthmus of Panama have
been written by Lessios (1998, 2008). Lessios (1998) wrote a chapter in ‘‘Endless
Forms: Species and Speciation’’ on the first stage of speciation as seen in organ-
isms separated by the Isthmus of Panama. Lessios points out that although gem-
inate species have had a long history of contributing evidence relevant to
speciation, they remain an underutilised tool for understanding vicariant specia-
tion. Lessios’s (2008) review showed that a total of 38 regions of DNA have been
sequenced in nine clades of echinoids, 38 clades of crustaceans, 42 clades of fishes,
and 26 clades of molluscs with amphi-isthmian subclades. Of these, 34 clades are
likely to have been separated at the final stages of Isthmus completion, 73 clades
split earlier and eight clades maintained post-closure genetic contact (e.g. via a
circumglobal route or through the Panama Canal). As no vicariant event is better
dated than that of the Isthmus of Panama, molecular divergence between species
pairs remains to be of great interest, particularly as adaptive divergence can be
seen in species life history parameters.
4.4 Diversity and Distribution of Echinoderms
Chesher (1972) published a list of Pacific and Caribbean echinoids and identified
potential geminate species pairs. Maluf (1988) synthesised and tabulated the
composition and distribution of central eastern Pacific echinoderms from the
literature, which has proved to be a valuable resource for researchers. Lessios
(2005a) updated the records of echinoids found in the Pacific waters of Panama,
and distinguished species that have been documented for Panama from those that
have collection records that straddle Panama, but with no published records of
actual presence in Panama. In the latter category Lessios (2005a) lists Kamptosoma
asterias (Agassiz, 1881), Plesiodiadema horridum (Agassiz, 1898), Cystocrepis
setigera (Agassiz, 1898), Pourtalesia tanneri Agassiz, 1898, Homolampas fulva
Agassiz, 1879, Echinometra oblonga (Blainville, 1825) and Encope perspectiva
Agassiz, 1841. The inclusion of E. perspectiva in this category is a mistake as it
was collected in Panama from Playa Grande, on San Jose Island, Pearl Islands
Archipelago by Clark (1946) and more recently by Coppard (2010) from the same
location. Plesiodiadema horridum is listed in the Smithsonian, National Museum of
Natural History collections as collected off Panama (USNM 21050). However, the
coordinates of the collection site (3°09.000 N, 82°08.000 W) indicate that it was
collected off Colombia. Lessios (2005a) gives details of why the subtropical
Centrostephanus coronatus (Verrill, 1867) and central and western Pacific species
E. oblonga are unlikely to occur in Panama, and that the record of Caenocentrotus
gibbosus (Agassiz, in Agassiz and Desor, 1846) in Panama is a mistake in the
literature.
Alvarado et al. (2008) assessed the diversity of echinoderms in the Caribbean of
Central America and gave faunal lists for each country. Panama had the greatest
species richness, with 154 species listed, comprised of 15 species of crinoids, 23
species of asteroids, 56 species of ophiuroids, 30 species of echinoids and 30
species of holothuroids. Among the crinoids they included both Davidaster
insolitus Clark 1917 and Davidaster discoideus (Carpenter, 1888). The number of
species of crinoids should be reduced to 14 because D. insolitus is currently
considered a junior synonym of D. discoideus (Messing 2010). Among the
asteroids, two species are missing from their list for Panama. These are Echinaster
(Othilia) echinophorus (Lamarck, 1816) collected from Limon Bay (Smithsonian
National Museum of Natural History, USNM 38235) and Fort Randolph (USNM E
26417, USNM E 26418), and Persephonaster patagiatus (Sladen, 1889) collected
from the Gulf of Mosquitos (USNM E 31501). The number of ophiuroid species is
incorrect as Amphiodia riisei (Lütken, 1869) [sic] is a junior synonym of
Ophiophragmus riisei (Lütken in: Lyman 1860) (Stöhr 2010). Among the holo-
thuroids they list Thyone deichmannae Madsen, 1941 and Thyone inermis Heller,
1868 which are junior synonyms of Havelockia inermis (Heller, 1868) (Hansson
2011). Although Alvarado et al. (2008) state that there are 30 echinoid species
recorded from off the Caribbean coast of Panama they only list 29 species which
agrees with more recent work by Coppard (2010).
Table 4.1 Number of echinoderm species recorded from Panamanian waters

Pacific Caribbean Both pacific and caribbean Total
Crinoidea 7 14 0 21
Total 260 144 10 414
Alvarado et al. (2010) documented the echinoderm diversity in the Pacific coast
of Central America, again providing faunal lists for each country. Although
Alvarado et al. (2010) included the paper by Lessios (2005a) in their bibliography
they unfortunately ignored Lessios’s comments and tables. These indicate echinoid
species that have distributions in the eastern Pacific that straddle Panama (but have
not been recorded from Panama), those unlikely to occur in Panama, and errors in
the literature. Alvarado et al. (2010) also incorrectly listed Encope grandis
Agassiz, 1841 in Guatemala, El Salvador, Honduras, Nicaragua and Panama. This
species is endemic to the Gulf of Mexico; Encope michelini Agassiz, 1841 in
Costa Rica (this is an Atlantic species that occurs off Florida), Dendraster
excentricus (Eschscholtz, 1829) in Panama (this temperate species has a distri-
bution range from Alaska to Baja California), as well as including many junior and
senior synonyms as separate taxa (e.g. Ophionereis annulata (Le Conte, 1851) and
Ophionereis dictyota Ziesenhenne 1940; Sclerasterias alexandri (Ludwig 1905)
and Hydrasterias diomediae Ludwig 1905; Lissothuria ornata Verrill 1867 and
Thyonepsolus beebei Deichmann, 1937). Alvarado et al. (2010) list 253 species of
echinoderms from Panama, the country with the highest species richness in Central
America. This is explained by the long coastline (1,690 km) of Panama which
incorporates high levels of coastal heterogeneity and reflects the greater echino-
derm research effort than in other Central America countries (Alvarado et al.
2010).
A web-based project by Coppard (2010) (‘‘The Echinoderms of Panama’’) has
documented 414 species of echinoderms that occur off the Caribbean and Pacific
coasts of Panama (Table 4.1 and Appendix). This work was carried out in con-
junction with the Encyclopaedia of Life and contains images (including type
material), diagnostic descriptions and references with links to nomenclators,
GenBank, Barcode of Life, and depth, distribution and specimen data. All species
names were checked with the World Register of Marine Species and lists of junior
synonyms are given. Where physical specimens were not found in museum col-
lections, records from the literature were used.
Since the final closure of the Panama Isthmus unequal rates of speciation,
extinction and migration have occurred in the Caribbean and Pacific, resulting in
very different levels of biodiversity seen on either side of the isthmus today
(Vermeij 1978; Jackson et al. 1996; Budd 2000). Among invertebrates, greater
levels of diversity have been reported in the Caribbean than in the Pacific for
Fig. 4.9 Nemaster grandis

with arms spread for feeding,
Bocas del Toro, Caribbean
Sea (photo courtesy of
Edgardo Ochoa)
corals (Glynn 1982), sponges (Van Soest 1994), benthic foraminiferans (Collins
1999), cheilostome bryozoans (Cheetham et al. 2001) and cupuladriid bryozoans
(O’Dea et al. 2004). With the exception of crinoids, the diversity of echinoderms
off Panama shows a very different trend, with a far greater diversity in the Pacific
(270 species) than in the Caribbean (154 species). The high diversity of crinoids in
the Caribbean is most likely a consequence of greater coral reef development,
which provides a broad range of suitable habitats for species such as Nemaster
grandis (Fig. 4.9).
For the other classes of echinoderms, the greater number of species in the
Pacific of Panama can partially be explained by a number of species having
crossed the Eastern Pacific Barrier (e.g. Toxopneustes roseus, Tripneustes
depressus, Acanthaster planci) and thus have a West Pacific origin, with no sister
species in the Caribbean. However, in many genera, particularly those that contain
mud and sand dwelling species, a genus in the Caribbean often contain one or two
species, while in the Pacific the same genus will often have more than double the
number of species (e.g. in Panama the genus Luidia has three species recorded
from the Caribbean, but nine species from the eastern Pacific; Astropecten, two
species from the Caribbean, four species from the Pacific; Encope, one species
from the Caribbean, five species from the Pacific). The different hydrological
conditions in the Gulf of Chiriquí and the Gulf of Panama have created high levels
of coastal heterogeneity providing a diverse range of habitats. However, such
morphological variation that has given rise to authors erecting many species names
in the Pacific may simply reflect ecophenotypic varriation within a species.
Sampling the sharp drop down the continental slope to a depth over 1,000 m
just south of the Azuero Peninsula has resulted in a number of deep water species
being recorded from the Pacific of Panama, such as sea lilies (e.g. Calamocrinus
diomedae Agassiz, 1890, depth 1,430 m), brittle stars (e.g. Astrodia plana (Lütken
and Mortensen 1899), depth 3,058 m; Asteronyx loveni Müller and Troschel, 1842,
Fig. 4.10 Deep water ophiuroids, a Astrodia plana and b Asteronyx loveni
depth 1,866 m, Fig. 4.10), starfish (e.g. Dytaster gilberti Fisher, 1905, depth
2,690 m; Pectinaster agassizi Ludwig 1905, depth 2,323 m), sea urchins (e.g.
Salenocidaris miliaris (Agassiz 1898), depth 3,058 m; Caenopedina diomedeae
Mortensen, 1939, depth 850 m; Araeosoma leptaleum Agassiz and Clark, 1909,
1,062 m) and sea cucumbers (e.g. Molpadia spinosa (Ludwig, 1893), depth
3,279 m; Protankyra brychia (Verrill, 1885), depth 4,000 m).
Ten species have collection records from both sides of the Panama Isthmus.
These are the ophiuroid species Hemipholis elongata (Say, 1825), Ophiactis
savignyi (Müller and Troschel, 1842), Ophioderma panamensis Lütken, 1859 and
Amphipholis squamata (Delle Chiaje, 1828); the asteroid species Linckia guil-
dingii Gray, 1840, Ctenodiscus crispatus (Retzius, 1805); the echinoid species
Moira atropos (Lamarck, 1816) (the subspecies Moira atropos clotho Michelin,
1855 is recorded from the Pacific), Meoma ventricosa (Lamarck, 1816) (the
subspecies Meoma ventricosa grandis Gray, 1851 is recorded from the Pacific);
and the holothuroids Holothuria (Thymiosycia) arenicola Semper, 1868 and
Holothuria (Thymiosycia) impatiens (Forskål, 1775). With the final closure of the
isthmus (2.8 mya) there has been no gene flow between populations in the Pacific
and Caribbean. However, the opening of the freshwater Panama Canal in 1914
provided a means for echinoderms to cross the isthmus, either as larvae in the
ballast tanks of ships or associated with fouling organisms on the hulls of ships.
Roy and Sponer (2002) found identical mtDNA haplotypes of Ophiactis savignyi
in the Caribbean and Pacific. This led them to suggest that these sponge-dwelling
brittle stars have been recently transported between oceans on the hulls of ships.
Global phylogenies have also shown that genes of certain species have travelled
around the world during the last 3 million years providing a connection between
tropical species on the two sides of tropical America (Bowen et al. 2001, 2006;
Lessios et al. 2001; Rocha et al. 2005).
Some echinoderm species have records in Panama that may represent transient
populations. Arbacia spatuligera (Valenciennes, 1846) is recorded (specimen not
examined) from the reef off French Fort (USNM 1017377), in Panama Bay. This
may be a misidentification or represent a transient population of this species, with
larvae carried up from Ecuador into the Gulf of Panama but with insufficient
recruitment to establish a viable population. This species has therefore not been
included in the Pacific data for Panama, since to the best of our knowledge it is the
only record of this species in Panama. Records of Lytechinus pictus in Panama are
probably erroneous. Mortensen (1921) described L. panamensis based on speci-
mens he obtained from Taboga Island. Clark (1948) collected one specimen from
Bahia Honda as well as one specimen from Gorgona Island, Colombia. However,
Clark (1948) states that these may be specimens of L. panamensis, a view sup-
ported by Zigler and Lessios (2004) and Lessios (2005a). Lessios (2005a) reports
that he has only seen two specimens of this species collected off Isla Saboga in the
Perlas Archipelago (8°380 N, 79°030 W). Clark (1925) and Mortensen (1928–1951)
speculated that Lytechinus from Peru and Ecuador may not be L. semituberculatus
but L. panamensis, with the larvae of this species occasionally settling in the Gulf
of Panama.
Panama is the type locality of 82 species of echinoderms (nine Caribbean, 73
eastern Pacific, one of these species occurs in both the Caribbean and Pacific).
However, four of these species (starfish) are now considered subjective junior
synonyms of other species. The majority of the Pacific species were established
from Panamanian specimens in the late 1800s and early 1900s by authors such as
Lütken and Mortensen (1899) and Ludwig (1894, 1905), as a result of the material
collected on expeditions by the Albatross. The majority of new Caribbean species
were discovered much later in the 1960s, 1970s and 1980s by authors such as
Chesher (1968, 1970, 1972) and Hendler (1995 2005), as a result of the labora-
tories of the Smithsonian Tropical Research Institute in Panama. In recognition of
this, Hendler (2005) named the ophiuroid species Ophiothrix stri Hendler 2005.
The sustainable aquaculture of echinoderms in Panama has not been developed. In

1997 the Panamanian government granted a permit for the harvesting and
processing of bêche-de-mer (sea cucumbers) in Bocas del Toro. However, this
exploitation was short lived, with the permit being revoked 30 days later.
Nevertheless, in that short time period Guzman and Guevara (2002b) estimate that
750,000 sea cucumbers were harvested (based on 25 fishermen extracting an
average of 1,000 and a maximum of up to 1,500 cucumbers day-1). This consisted
primarily of three species Holothuria (Halodeima) mexicana, Isostichopus
badionotus and Astichopus multifidus. This quantity represents a biomass of ca.
180 tons (based on average body wet weight of 240 g). If such fishing pressure
was permitted, Guzman and Guevara (2002b) estimate that stocks of the three
Fig. 4.11 Astichopus

multifidus photographed in
Bocas del Toro (photo
courtesy of Juan Sanchez)
species of holothuroids would collapse within a very short period of time

(305 days for H. (H.) mexicana, 221 days I. badionotus and only 9 days for
A. multifidus), based on a catch effort of 25,000 ind day-1 (Fig. 4.11)
Commercial bêche-de-mer fishing currently is banned in all Panamanian waters
under Decree 157–2003. However, there are reports of illegal fishing activities,
with five species being commercially exploited in Panama: Actinopyga agassizi
(Selenka, 1867) H. (H.) mexicana, A. multifidus, I. badionatus (Caribbean) and
I. fuscus (eastern Pacific) (Toral-Granda 2008). In Parque Nacional Marino Isla
Bastimentos, H. (H.) mexicana showed a maximum density of 956 ind ha-1 in
480 ha representing only 10 % of the shallow water protected area, while
I. badionotus and A. multifidus were strikingly absent in 95 % of the protected
insular area around Cayos Zapatillas. This indicates that that illegal fishing has
taken place inside the park (Guzman and Guevara 2002b). Data from the statistics
department in Hong Kong, revealed that 281 kg (dry weight) of bêche-de-mer
were imported from Panama in 2004 and 408 kg (dry weight) in 2005 (Toral-
Granda 2008).
4.6 Threats to Echinoderms
Between 1983 and 1984, D. antillarum suffered one of the most extensive and
severe mass mortality events ever recorded for a marine animal. This was probably
caused by a waterborne pathogen transported by ocean currents, since plots of
surface currents in the Caribbean Sea coincided with the spread of the die-off
(Lessios 1988; Miller et al. 2003). The mass mortality of D. antillarum has been a
major factor leading to a shift from coral-dominated to algae-dominated com-
munities on many Caribbean reefs during the past 20 years (Lessios 1988). The
die-off was first observed in January 1983 at Galeta Point, near the entrance of the
Panama Canal. In approximately three months it had extended to the San Blas
Archipelago, before reaching the Panama-Colombia border by the end of June. By
Fig. 4.12 Diadema

antillarum on a reef in San
Blas, depth 2 m, June 2010
January 1984 the die-off had spread throughout the rest of the Caribbean with
population densities in Panama reduced to 5.8 % of their previous levels (Lessios
et al. 1984a). The first symptom of the presumed disease was accumulation of
sediment on the spines and sloughing off of the spine ectoderm. This was followed
by loss of pigmentation in the skin covering the spine muscles, the peristome, and
in the periproctal cone. Spines became brittle and broke, while others fell off
completely, exposing areoles and tubercules. Tube feet became flaccid, no longer
retracted fully upon stimulation, and were unable to cling to the substrate. Many
individuals at this stage were found on their sides or being moved to and fro by the
surge. In advanced stages large patches of skin pealed off the test, the peristome
and the anal cone lost practically all pigmentation, and the spines could be pulled
off with little force (Lessios et al. 1984b).
Before the mass mortality, mean densities of D. antillarum on reefs at Punta
Galeta and San Blas where 1.82 and 1.91 ind m-2, respectively (Lessios et al.
1984b). After the mass mortality the densities dropped to an average of 0.05 ind
m-2 in San Blas, and to 0.004 ind m-2 in Punta Galeta. This density reduction was
not observed in other species of echinoids (E. lucunter, E. viridis, Echinoneus
cyclostomus (Leske, 1758), E. tribuloides, L. variegatus and T. ventricosus (La-
marck 1816)) on the same reefs. In San Blas, the mean sizes were reduced from
48.6 mm to 25.0 mm (Lessios et al. 1984b). However, this mass reduction on the
population did not have any consequence on its genetic variability (Lessios
1985b).
Five years after the event, Lessios (1988) did not observe any recovery of the
populations on the San Blas reefs, with little influx of juveniles and no increase in
population densities of other sea urchins species (E. viridis, E. lucunter, E. tribuloides,
T. ventricosus, or L. variegatus). He reported no alteration in the reproductive
behaviour of D. antillarum, proposing that the low recruitment success could be an
effect of low populations densities and low adult numbers.
Fig. 4.13 Mellitid sand dollars showing varying degrees of predation and secondary infection,
a M. longifissa, collected alive (sublethal predation), b E. laevis (lethal predation, no secondary
infection), c M. kanakoffi, found dead (probable death from secondary infection due to the large
amount of necrotised tissue)
Ten years after the mass mortality densities remained at less than 3.5 % of their
pre-mortality levels, with D. antillarum being absent in its preferred habits in San
Blas (Lessios 1995). Moreover, the mean size of individuals remained low
(17.3–35.2 mm). This was thought to be a direct result of the disease, as food
availability was not a limiting factor (Lessios 1995).
Twenty years after the mass mortality populations remained at less than 6.5 %
of their pre-mortality levels in San Blas (Lessios 2005b). Some successful
recruitment events have been observed on a few reefs. However, these have not
been sustained over time (Lessios 2005b). The lack of apparent recruitment could
have been caused by too few adults in upstream areas to reproduce at a rate that
would overcome the normal sources of mortality (the Allee effect) or by the
continuing presence of the pathogen that devastated the populations in 1983–1984.
Recent observations (2010) on some San Blas reefs indicate a slight recovery
(Fig. 4.12), however, it remains to be seen if these populations will be sustained.
On 27 April 1986 more than 8 million liters of crude oil spilled into a complex
region of mangroves, seagrassses, and coral reefs east of the Caribbean entrance to
the Panama Canal (Jackson et al. 1989). At the seaward edge, numbers of the most
abundant sea urchin E. lucunter was reduced by 80 % within a few days of the
spill, and the reef flat was littered with its skeletons (Jackson et al. 1989). How-
ever, further inshore no such mortality of E. lucunter was reported (Jackson et al.
1989).
During 2008–2009 Coppard (unpub. obs.) found large numbers of dead and
dying sand dollars of the genera Encope and Mellita washed up on beaches from
Punta Abreojos, Baja California Sur, Mexico, to Santa Elena Bay, La Libertad in
Ecuador in the eastern Pacific and off Costa Rica and Belize in the Caribbean. In
Panama dead mellitid sand dollars were found at Playa Las Lajas in the Gulf of
Chiriqui, but not on beaches in the Gulf of Panama. These dead sand dollars were
unusual in that they appeared to have an infection that originated at one or more
circular denuded regions on the aboral surface of the test (Fig. 4.13a). The cause of
such scars is most likely the result of attacks by gastropods from species such as
Northia pristis (Deshayes in Lamarck, 1844) and Polinicies sp. in the Pacific and
Cassis tuberosa (Linneaus, 1758) in the Caribbean which have been reported to
attack, and bore into the tests of mellitid sand dollars (McClintock and Marion
1993; Sonnenholzner and Lawrence 1998). Attacks always occurred on the aboral
surface, and in the majority of cases centred over one or more petals. When the
gastropods managed to bore through the test into the coelom (Fig. 4.13b), death of
the sand dollar resulted. However, the presence of multiple scars of varying size
indicates that such attacks are not always lethal. Following such attacks bacteria
infected the majority of wounds which became dark green or black as a result of
necrosis (Fig. 4.13c).
In many instances where the coelom had not been penetrated, the bacteria in the
wound spread and caused the death of the sand dollar. Such pathogenic bacteria
may be similar to that reported in Meoma ventricosa in Curaçao by Nagelkerken
et al. (1999). In this instance a tetradotoxin-producing bacterial strain of the genus
Pseudoalteromonas was found in the sediment and was ingested by M. ventricosa.
The density of such strains of bacteria may have increased as a result of the
outflow of polluted water from the harbour.
In Bocas del Toro, only 2.8 % of the shallow habitats in the archipelago are
within the protected area (Guzman and Guevara 2002a). Since 1997, coastal
environments in Bocas del Toro have changed, as infrastructure such as roads,
marinas, and hotels have developed. Destruction of these seagrass habitats is
resulting in an increase in runoff and sedimentation in the coastal zone (Guzman
and Guevara 1998). This has already started to affect the coral reefs (Guzman and
Guevara 2001), and could potentially modify the function of the remaining coastal
ecosystems.
The illegal harvesting of sea cucumbers continues to put pressure on the
remaining populations particularly on the Caribbean coast of Panama, along the
coastal perimeter of Laguna de Chiriquí, from the point of Península Valiente
through the southern section of Isla Popa and Cayo Agua (Cruz 2000; Guzman and
Guevara 2002a).
4.7 Conclusions and Recommendations
Echinoderm diversity in Panama is reported to be greater on both the Caribbean

(Alvarado et al. 2008) and eastern Pacific coasts (Alvarado et al. 2010) than in
other Central American countries. The length of coastline, with high levels of
coastal heterogeneity may explain high levels of species richness. However, this
diversity relative to other Central American countries is probably more of a
reflection of the research effort that has taken place in Panama. The first phase of
echinoderm research in Panama began in the late 1800’s with the collections made
by the U.S. Fish Commission Steamer Albatross in the eastern Pacific. This period
of exploration resulted in many new species being described. The second phase of
echinoderm research in Panama in the 1930’s and 1940’s continued to explore the
diversity of species and was the result of expedition by the New York Zoological
Society and the Allan Hancock Foundation. These expeditions resulted in the
collection of a vast number of specimens (some 28,835 echinoids alone from
eastern Pacific of the Americas) with the description of many new species. The
third phase of echinoderm research in Panama began with the inception of the
marine program at STRI in 1961. This has resulted in extensive ecological and
evolutionary echinoderm research, which has increased knowledge of echinoderms
not only in Panama, but in many countries in tropical America, and other regions
in the Atlantic and Pacific. Most of this knowledge concerns the ecology, repro-
duction and evolution in echinoids and certain species of asteroids (e.g. A. planci
and O. reticulatus). However, less is known about holothuroids, ophiuroids and
crinoids, particularly on a molecular and ecological level, highlighting a need for
more research in these classes of echinoderm.
The continued monitoring of populations of Diadema antillarum needs to be
maintained to determine whether the recently observed population increase is
sustained, while the monitoring of sea cucumber populations, particularly in Bocas
del Toro is needed to protect against illegal fishing. Guzman and Guevara (2002b)
have proposed that protected areas should be established allowing the monitoring
of natural populations. These could be part of the Parque Nacional Marino Isla
Bastimentos (PNMIB) or may be located close to fishing communities willing to
protect the resource.
Acknowledgments Simon Coppard thanks SENACYT (La Secretaría Nacional de Educación

Superior Ciencia Tecnología e Innovación) (Panama) who helped fund this project as part of
project COL08-002 and David M. Rubenstein who funded Simon through a Smithsonian
Rubenstein fellowship. Juan José Alvarado thanks Consejo Nacional de Ciencia y Tecnología
(CONACYT; Mexico) and Consejo Nacional para Investigaciones Científicas y Tecnológicas
(CONICIT; Costa Rica). We are grateful to Edgardo Ochoa, Juan Sánchez, Angel Chiriboga and
Dave Pawson for providing us with photographs. We appreciate the comments made by J. Cortés,
J. Lawrence and H.A. Lessios that improved this paper.
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Chapter 5
Echinoderms of Colombia
Milena Benavides-Serrato, Giomar Helena Borrero-Pérez,

Jaime R. Cantera K, Mishal Cohen-Rengifo and Raúl Neira
5.1 Introduction
Colombia, together with Indonesia, Brazil, and Mexico, shares a privileged

position at the top of the list of countries with the highest species richness in the
world, the so called megadiverse countries (IAvH 1998a, b). However, this rep-
utation is based upon the relatively well documented diversity of terrestrial biota
such as orchids, amphibians, birds, mammals, and butterflies (Andrade et al.
1992). Biodiversity in Colombia is related to the high diversity of habitats that
occur on the mainland. Close to half of the country is covered by the Amazon
River system and the Orinoco savannas. The Andes Mountains are separated into
three ranges, and drainage from the Magdalena and Cauca River basins forms the
wetlands of the Caribbean and feeds the tropical dry forest ecosystems.
M. Benavides-Serrato (&) G. H. Borrero-Pérez

Instituto de Investigaciones Marinas y Costeras—INVEMAR, A.A 1016,
Santa Marta, Colombia
e-mail: milena_benavides@invemar.org.co
G. H. Borrero-Pérez
e-mail: giomar_borrero@invemar.org.co
J. R. Cantera K R. Neira
Departamento de Biología, Universidad del Valle, A.A. 25360, Cali, Colombia
e-mail: jaime.cantera@correounivalle.edu.co
R. Neira
e-mail: rneira56@yahoo.com
M. Cohen-Rengifo
Centre d’Oceanologie de Marseille, Université de la Méditerranée,
Marseille Cedex, France
e-mail: ablocsberg@gmail.com

146 M. Benavides-Serrato et al.
Fig. 5.1 Location of the land and sea areas of Colombia (System Information Laboratory
LABSIS-INVEMAR)
The watershed of the western Andes and the Pacific lowlands is an extremely rich
region, isolated for many years, constituting the Chocó biodiversity hotspot. The
Sierra Nevada de Santa Marta, the highest coastal mountain in the world, is an
isolated mountain in the Caribbean, rising 5,500 m above the hot deserts of the
Guajira Peninsula (Díaz and Acero 2003).
However, with nearly 1,000,000 km2 of territorial waters, nearly half the ter-
ritory of Colombia is maritime. It is the only South American country with coasts
on both the tropical Pacific Ocean and the Caribbean Sea, with a total shoreline of
nearly 3,000 km. Because Colombia possesses oceanic islands located far off-
shore, wide sections of both the tropical eastern Pacific and the southern-central
Caribbean are included within the boundaries of its Economic Exclusive Zone
(Fig. 5.1).
5 Echinoderms of Colombia 147
There are remarkable differences between the two coasts in terms of climatic,
geological, oceanographic, ecological, and biological features (Botero and Alvarez-
León 2000; Marrugo-González et al. 2000). In addition, coastal morphology, cli-
mate, and physical oceanographic processes vary greatly along spatial gradients as
well as seasonally within both the Caribbean and the Pacific. These features allow
the great majority of tropical marine habitats to be well represented in Colombia.
Considering that each habitat harbors its own particular biota, the total diversity of
marine organisms in such a variety of environments is likely to be very high. Their
study represents a huge challenge for future biological research.
The information presented in this chapter is derived from a compilation and
review of the knowledge of Colombian echinoderms, based on diverse sources of
information such as published echinoderm studies, undergraduate thesis, technical
reports and extracted information from databases of museums as National Museum
of Natural History (NMNH) and other databases available such as Biodiversity
Marine Information System (SIBM) of Colombia, Ocean Biogeographic Infor-
mation System (www.iobis.org) and Inter-American Biodiversity Information
Network (www.iabin.net). This chapter represents a baseline study of echinoderms
in Colombia, useful for continuing and developing further research. Because of the
physical and biological differences for the Caribbean and Pacific coasts of
Colombia, as well as historical differences in the research carried out on these two
regions, the information is presented separately for the two coasts.
5.1.1 Colombian Caribbean Sea
The Colombian Caribbean coast extends from 8°400 N and 77°210 W at the Panama-
Colombia border, northeastwards to 11°550 N and 71°180 W at the Colombia-
Venezuela border (Fig. 5.2). San Andrés and Providencia archipelago (between
12°–15° N and 79°–82° W) are two high-standing oceanic islands and a series of
atolls and coral banks which are aligned in a NNE direction along the lower
Nicaragua Rise. The total coastline of the Colombian Caribbean is 1,937 km, with
a terrestrial area of 6,958 km2, insular area of 79 km2, and territorial sea of
532,162 km2. It includes eight departments: Guajira, Magdalena, Atlántico,
Bolivar, Sucre, Cordoba, Antioquia and San Andres, Providencia and Santa Cat-
alina (INVEMAR 2010).
There is a remarkable climatic gradient along the Caribbean mainland coast.
The northernmost part of the Guajira peninsula is extremely dry and hot and the
land vegetation is very sparse. In contrast, annual precipitation in the area around
the gulf of Urabá, close to the Panama boundary is continue and is over
3,000 mm year-1, and coastal plains and mountains there are covered by dense
tropical rain forest. The Urabá region represents a northern outpost of the Chocó
diversity hot spot for the terrestrial biota (Gentry 1986).
On the other hand, geomorphologic, physical oceanographic and ecological
processes vary greatly along spatial gradients as well as seasonally within the
Fig. 5.2 Ecoregions and coastal departments of the Colombian Caribbean Sea: GUA: Guajira,
PAL: Palomino, TAY: Tayrona, MAG: Magdalena, ARCO: Coralline Archipelagos, MOR:
Morrosquillo, DAR: Darién, SAN: San Andrés and Providencia Archipelago, COC: Oceanic
Caribbean. (System Information Laboratory LABSIS-INVEMAR)
Colombian Caribbean. This results in an irregular or heterogeneous distribution of

major habitats and ecosystems in the region. Upwelling in La Guajira peninsula
promotes the abundance of marine vegetation (seagrass and algal beds) but limits
the development of coral reefs. The water conditions of a large section of the
middle part of the coast are influenced by heavy discharge of freshwaters and
sediments by major rivers (Magdalena, Canal del Dique, Sinú) which form alluvial
plains, and deltaic formations with coastal lagoons. Estuarine conditions prevail in
these lagoons and mangrove forests are extensive. In contrast, warm waters and
high isolation around the offshore islands on the continental shelf and at the
oceanic archipelago of San Andrés and Providencia foster the development of
coral reefs. Finally, the northern ranges of the Sierra Nevada de Santa Marta sink
steeply into the sea. This result in an irregular shoreline with cliffs and pocket
beaches creating a diverse mosaic of rocky and sandy shores, fringing reefs, and
seagrass meadows (INVEMAR 2000).
Insular Formations of Colombian Caribbean Sea
Two main island groups are present in the Colombian Caribbean Sea, the archi-
pelago of San Andrés, Providencia, Santa Catalina and its cays, and the archi-
pelago of Rosario, San Bernardo and Isla Fuerte. The former was declared a World
Biosphere Reserve in the year 2000 and is characterized by volcanic elevations
covered by carbonate platforms produced by the reef biota. These emerge to form
islands and cays, surrounded by coral reefs and sedimentary plains, vegetated by
algae or seagrasses that form a shallow island shelf (down to 40 m deep),
surrounded by seafloor exceeding 600 m in depth. The second archipelago is part
of the Marine Protected Area Corales del Rosario, San Bernardo and Isla Fuerte,
the only marine protected area of Colombia, enclosing the most developed coral
reefs of the Colombian north coast and a high biodiversity of other ecosystems
(INVEMAR 2010).
Caribbean Ecoregions
The Colombian Caribbean Sea has been divided into nine ecological regions
according to natural characteristics like geomorphology, hydrography, sedimen-
tology, and ecosystems. These will be used to present the diversity and distribution
of echinoderms. They are: Guajira (GUA), Palomino (PAL), Tayrona (TAY),
Magdalena (MAG), coralline archipelagos (ARCO), Morrosquillo (MOR), Darién
(DAR), San Andrés and Providencia Archipelago (SAN) and Oceanic Caribbean
(COC) (Fig. 5.2). Six of these regions are coastal and limited by the 200 m isobath
that defines the continental shelf platform. SAN is an insular sector that comprises
the atolls, coral reefs and islands of the San Andres and Providencia archipelago.
The Oceanic Caribbean sector (COC) includes the continental slope extending
from 200 to 2,700 m in the Colombian Basin (Tabares et al. 1996; INVEMAR
2000). Table 5.1 summarizes the main physical, oceanographic and ecological
features of these ecoregions.
5.1.2 Colombian Pacific Ocean
The Pacific coast of Colombia is located in the northwest region of South America,
between 7°120 N–77°530 W and 1°260 N–78°490 W, bordering Panama in the north
and Ecuador in the south. Four departments are in this area: Chocó, Valle del
Cauca, Cauca and Nariño, and includes the islands of Gorgona, Gorgonilla and
Malpelo. The coastline extends for 1,576 km. The departments have a terrestrial
area of 8,181 km2, an insular area of 14 km2, and a territorial sea area of
359,995 km2 (Steer et al. 1997; INVEMAR 2002) (Fig. 5.3).
The Pacific coast of Colombia can be divided into two areas based on phys-
iographic and hydrographic characteristics. The northern coast, from Panama to
Table 5.1 Characteristics of Colombian Caribbean and Pacific ecoregions (Modified from Díaz and Acero 2003)
150
Ecoregion Climate and oceanography Coastal, shelf and seafloor morphology Main ecosystems represented
Caribbean
Guajira (GUA) Highly influenced by strong winds, desert. Low coastal plains with scattered rocky cliffs. Seagrass beds, sandy beaches, salt
Seasonal upwelling causing high primary and Wide continental shelf marshes, scattered fringing
secondary marine production mangrove forests, sandy bottoms
Palomino (PAL) Dry to semi humid; heavy coastal swell; several Highly exposed coast dominated by coarse Beaches, coastal lagoons and
small rivers sand beaches. Continental shelf narrow swamps, sandy bottoms
Tayrona (TAY) Semi-desert to dry; seasonal upwelling High coastal mountains form a series of small Rocky shores, coral reefs, seagrass
bays and inlets with pocket beaches and beds, beaches, rocky and sandy
cliffs. Continental shelf very narrow or bottoms
absent
Magdalena Dry. Coastal waters highly influenced by Coastal morphology very heterogeneous and Coastal lagoons and estuaries,
(MAG) discharges of the Magdalena drainage system dynamic. Continental shelf rather narrow mangrove forests, sandy beaches,
with heavy sediment loads (turbid waters) made of terrigenous sand and silts muddy bottoms
Coralline- Dry climate. Transparent, oceanic waters Offshore shallow carbonate platforms and Coral reefs, seagrass beds, bioclastic
Archipelagos alternate seasonally with turbid waters from coral shoals, scattered coralline island cays sandy bottoms
(ARCO) continental discharges
Gulf of Dry to semi-humid. Coastal waters influenced by Coastal morphology dominated by alluvial Seagrass beds, mangrove forests,
Morrosquillo continental discharges (mostly turbid waters) plains and deltaic systems; low exposure to estuaries, beaches, muddy
(MOR) wave action. Continental shelf wide and bottoms
muddy
Darién (DAR) Semi-humid to very humid. High influence of Coastal morphology very variable (alluvial Estuaries, mangrove forests,
continental discharges (predominantly turbid plains, terraces, rocky cliffs). Continental scattered seagrass and corals,
waters) shelf wide muddy bottoms
San Andrés and Dry to semi-humid, frequently exposed to storms Two major high standing islands, sandy cays, Coral reefs, sandy beaches and
Providencia and hurricanes. Oceanic waters submerged carbonate platforms submarine bottoms, scattered
Archipelago seagrass beds and mangroves
(SAN)
(continued)
M. Benavides-Serrato et al.
Table 5.1 (continued)
Oceanic Oceanic waters and seafloor beyond continental Continental and insular shelf slopes and Pelagic realm; bathyal and abyssal
Caribbean and island shelves abyssal plains, scattered guyots and sea ecosystems, water depths
(COC) mounts between 150 to more than
3,000 m
Pacific
Northern Pacific Very rainy; rather clear waters; seasonally Very narrow continental shelf; high tectonic Scattered mangrove forests and coral
(PAN) affected by upwelling activity; coastal morphology defined by formations
5 Echinoderms of Colombia
mountains with a dense canopy of rain

forest; steep rocky (volcanic) cliffs
alternating with pocket beaches
Baudó (BAU) Marine waters moderately influenced by Coastal morphology defined by a large alluvial Sandy beaches, intertidal sand flats
terrestrial runoff plain with extensive sandy beaches and
intertidal sand flats; rather narrow
continental shelf
Buenaventura Turbid waters Coastal morphology heterogeneous. There are Alluvial and deltaic plains with
(BUE) terraces of soft rocks with steep cliffs mangroves, intertidal mud plains,
subject to marine erosion; narrow sandy beaches and littoral
continental shelf terraces
Naya (NAY) Influenced by many rivers and littoral barrier Coastal morphology defined by an ample Highly dynamic estuarine system;
islands; turbid waters alluvial plain; rather wide continental shelf well developed mangrove forests
behind the barrier islands
Sanquianga Large deltaic system formed by several Wide continental shelf Mangrove forests and intertidal sand
(SAQ) meandering rivers which form a complex plains (nesting habitats for sea
system of estuaries, channels and open turtles). A protected area
lagoons; turbid waters (Sanquianga National Park)
Tumaco (TUM) Influenced by rivers, such as the Mira River, Narrow continental shelf Alluvial plains; scattered cliffs;
adjacent to the Ecuadorian border; turbid fringing mangrove forests
waters
(continued)
151
152
Gorgona Island Relatively clear waters Mountain island covered by rain forest Rocky and sandy shores, fringing
(GOR) coral carpets and reefs,
submerged rocky and soft
bottoms
Malpelo Island Oceanic island surrounded by waters up to Steep basaltic cliffs, insular shelf absent Rocky bottoms; scattered coral
(MAL) 1,000 m depth carpets; breeding area for oceanic
birds
Oceanic Pacific Moderately high pelagic productivity enhanced Territorial and Economic Exclusive Oceanic It is one of the most important
(PAO) by nutrients from terrestrial runoff Zone from 200 to 3,000 m depth commercial tuna fishing areas in
American Pacific
M. Benavides-Serrato et al.
Cabo Corrientes, has uplifted shores dominated by rocky cliffs and rocky shores
with intrusions of sand and silt, forming a high volcanic coastline (Galvis and
Mojica 1993; INGEOMINAS 1998; INVEMAR 2008). The continental shelf is
narrow and the 200 m isobaths is located about 15 km from the coastline (Cantera
and Contreras 1993). Despite the high rainfall in the area, the proximity of the
coast to the Serranía de Baudó does not permit the formation of rivers. The few
rivers that exist have a relatively low discharge.
The southern area, from Cabo Corrientes to the border with Ecuador, has
lowline coasts with tidal plains. It is mostly covered by mangroves, with cliffs in
Málaga, Buenaventura and Tumaco Bay. This area has several rivers with high
flow. The main ones are the Baudó and San Juan (INGEOMINAS 1998;
INVEMAR 2008). Sediments deposited by these rivers have formed a wide
continental shelf, extending about 55 km to Gorgona Island (Cantera and Con-
treras 1993) (Fig. 5.3).
Insular Formations of Pacific Colombia
The two main islands or systems of islands in the Colombian Pacific, Gorgona and
Malpelo, have been designated as protected areas (Fig. 5.3). The first is located
35 km from the continent in the Department of Cauca, between 2°550 N–78°090 W
and 3°000 N–78°140 W. It is the largest island territory on the continental shelf of the
Pacific, with a total area of 14.1 km2 and surrounding an area of 610 km2 (Fig. 5.3).
Sixty-one thousand ha of the waters surrounding Gorgona and Gorgonilla islands
were designated as a National Natural Park (PNN) in 1983 (UAESPNN 1998;
Giraldo et al. 2008). Although the coral reefs of the PNN Gorgona have a relatively
small area, they are one of the most developed of the Eastern Tropical Pacific (ETP)
(Zapata and Vargas-Ángel 2003).
Malpelo is an oceanic island 490 km west of the Department of Valle del Cauca
between 3°510 N and 81°350 W, with a terrestrial area of 3.5 km2. It was declared a
Fauna and Flora Sanctuary (SFF) in 1995. Because of its especially sensitive
marine nature, the marine protected area was expanded from 651 to 8,574 km2 in
2002. In 2006, UNESCO classified it as a World Heritage Site (UAESPNN 1998;
Díaz et al. 2000; UNESCO 2006). The island is separated from the mainland by a
depth of more than 3,000 m (Graham 1975). It is the only one that emerges from
the Malpelo Ridge, a submarine volcanic cordillera 241.4 km long and 80.5 km
wide (Chase 1968). The SFF is surrounded by clear oceanic waters that allow the
presence of corals to a depth of 30 m (Graham 1975).
Pacific Ecoregions
The Pacific region is subdivided into nine ecoregions based on environmental,

social, cultural and political factors: Pacific North (PAN), Baudó (BAU),
Buenaventura (BUE), Naya (NAY), Sanquianga (SAQ), Tumaco (TUM), Gorgona
(GOR), Malpelo (MAL), and Pacific Ocean (PAO) (Fig. 5.5). These ecoregions
will be used to present the diversity and distribution of echinoderms. Six of these
sectors are coastal and limited by the 200 m isobath, which in turn defines the
continental shelf. In the cases of GOR, geographical boundaries are the 40 m
isobath, on the eastern side and the 200 m isobath on the western side, while MAL
emerges from 500 m deep sea floor isobath, beyond the continental shelf. The
ecoregion PAO is represented by all jurisdictional Colombian marine areas on the
Pacific, adjacent to the conventional boundary of the continental shelf below a
depth of 200 m (INVEMAR 2000). Table 5.1 summarizes the main physical,
oceanographic, and ecological features of these ecoregions.
5.2.1 Historical Account of Scientific Studies About Echinoderms

on the Caribbean Coast of Colombia
Studies of the echinoderms in the Colombian Caribbean Sea are recent, dating
from the 1970s. They have been focused on shallow waters (0–50 m). The studies,
which were mostly taxonomic, were published by Allain (1976), Meyer and
Macurda (1976), Caycedo (1978, 1979), Álvarez (1981), Quiñones (1981), Gallo
(1985, 1988a, b), and Aguilar-Duarte (2005). These authors published taxonomic
lists and ecological and biological notes of echinoderms of different ecosystems
along the Colombian Caribbean coast, particularly of Tayrona National Natural
Park (PNNT), other areas near Santa Marta and Rosario Corals, and San Bernardo
Islands National Natural Park (PNNCRSB). Three new species of echinoderms
were described from Colombian material, Ctenantedon kinziei Meyer, 1972,
Holothuria (Thymiosycia) thomasi Pawson and Caycedo, 1980, and Ophiothrix
synoecina Schoppe, 1996.
Some investigations have been focused on the ecology of the echinoderm
community in different kinds of habitats, such as sandy beaches and rocky shores
(Arevalo 1978; Schoppe 1993). This kind of research has increased during the last
decade including soft bottom, coral reefs, and other ecosystems (Gárces and
Romero 2003; Vélez 2003; Bejarano et al. 2004; Aguilar-Duarte 2005; Monroy
and Solano 2005; Gómez and Rodriguez 2009; Romero 2009; González-Gaviria
2010). On the other hand, echinoderms have been incorporated into the Coral
Reefs Monitoring National System (SIMAC) in Colombia. On the Caribbean
Colombian coast, echinoderms have been observed from 1998 to 2011. Studies
have focused on seven species of sea urchins: Diadema antillarum, Echinometra
viridis, E. lucunter, Eucidaris tribuloides, Lytechinus variegatus, L. williamsi, and
Tripneustes ventricosus (Navas-Camacho et al. 2010).
Some studies have been conducted, such as Allain (1978), who studied
the deformations of tests of Lytechinus variegatus collected in Cartagena Bay
Fig. 5.3 Ecoregions and coastal departments of the Colombian Pacific Sea: PAN: Northern
Pacific; BAU: Baudó; BUE: Buenaventura; NAY: Naya; GOR: Gorgona Island; SAQ:
Sanquianga; TUM: Tumaco; MAL: Malpelo Island; PAO: Oceanic Pacific (System Information
Laboratory LABSIS-INVEMAR)
(MAG ecoregion). Ortiz (2006) studied the change in the ossicle density and shape
during growth of some species of the genus Holothuria from the Caribbean
and Pacific coasts.
With respect to deep waters, there are few records from the decade of the 1960s
from samples collected by international research cruises in Colombian Caribbean
waters. Among them were five biological voyages by the research vessel Oregon
in 1963: Oregon I: 87, 92, and 104 and Oregon II: 4 and 22 (Palacio 1974); the
research vessel of the University of Miami (R/V John Elliot Pillsbury) in 1966 and
1968, which sampled the coast of Panama, Colombia, Venezuela, Surinam and
French Guyana (Voss 1966; Bayer et al. 1970).
In 1998, a new era of research into of colombian echinoderms began, focusing
especially on mid-water (depth of 200 m) and deep-water (up to a depth of
1,000 m) echinoderms, as results of explorations carried out in the Colombian
Caribbean Sea: CIOH-INVEMAR-SMITHSONIAN, Macrofauna I-II, Macrofa-
una-CORPOGUAJIRA, Macrofauna-MARCORAL y ANH I-II. These expeditions
were conducted entirely by the Marine and Coastal Research Institute ‘‘José Benito
Vives de Andreis’’ (INVEMAR) with support from several institutions in
Colombia and other countries. These included the National Museum of Natural
History (NMNH-Smithsonian Institution), the Ministry of Environment, Housing
and Territorial Development (MAVDT), the Administrative Department of
Science, Technology and Innovation (COLCIENCIAS), the Autonomous Regional
Corporation of La Guajira (CORPOGUAJIRA), and the National Hydrocarbons
Agency (ANH). An important result of these projects was the discovery and
subsequent description of the azooxanthellate deep coral banks in some localities
of the ARCO, TAY and PAL ecoregions (Reyes et al. 2005).
Result from the last 13 years of research include five new records of the class
Crinoidea (Benavides-Serrato et al. 2011), 30 Asteroidea (González et al. 2002;
Benavides-Serrato et al. 2005, 2011), 46 Ophiuroidea (González et al. 2002;
Borrero-Pérez and Benavides-Serrato 2004a; Borrero-Pérez et al. 2008; Bena-
vides-Serrato et al. 2011), eight of Echinoidea (Borrero-Pérez et al. 2002a, b;
González et al. 2002; Borrero-Pérez and Benavides-Serrato 2004b), and 11 of
Holothuroidea (González et al. 2002; Borrero-Pérez et al. 2003). Recently,
Benavides-Serrato et al. (2011) created a catalogue of the classes Crinoidea,
Asteroidea and Ophiuroidea, in which they described 169 species distributed in the
Colombian Caribbean Sea.
Based on this information, some studies about the patterns of distribution and
abundance, ecological aspects and preliminary biogeography of echinoderms
present in the continental shelf and slope of the Colombian Caribbean Sea between
depths of 20 and 940 m have been published (Benavides-Serrato et al. 2001;
Benavides-Serrato and Borrero-Pérez 2004, 2010; Borrero-Pérez et al. 2004).
Part of this important process was the consolidation of the echinoderm col-
lection of the Colombian Museum of Marine Natural History (MHNMC) into
3,804 lots, principally collected from regions of the Colombian Caribbean Sea.
The evolution of the collection coincides with information retrieval by the
INVEMAR projects and also with the activities developed in these projects aimed
at improving the preservation and management of the collections. At the same
time, the Marine Biodiversity Information System (SIBM) was improved and is
now available through the INVEMAR web page (www.invemar.org.co), where a
special section is designated for the echinoderm collection.
5.2.2 Historical Account of Scientific Studies About Echinoderms

in the Pacific Coast of Colombia
Echinoderms from the Eastern Tropical Pacific (ETP), particularly on the Pacific
coast of Colombia, are not well known. The first records were collections made by
European explorers from the eighteenth century, when some descriptions of
species of this area were published.
To find works specifically devoted to the understanding and study of the fauna
of echinoderms of this area, it is necessary to go to the mid-nineteenth century.
At this time, several scientists from the United States and European institutions
published new species descriptions and reported new distributional data. The most
important contributions to the lists of species of echinoderms at that time in this
area are the works of Gray (1840), who described a large number of species of
asteroids, and Verrill (1867, 1868, 1869, 1870 and 1871), who published many
descriptions and lists of common species of shallow waters and described a large
part of the regular echinoids of the ETP. Lütken (1859) and Lyman (1879, 1882)
contributed descriptions of new species of ophiuroids. Selenka (1867) contributed
to knowledge of the holothurians. The Challenger Expedition (1873–1876) took
samples near the Galapagos Islands and the Gulf of Panama (Sladen 1889; Théel
1886), including species found in Colombian waters. At the end of the nineteenth
and the beginning of the twentieth century, as a result of the Albatross expeditions,
Agassiz (1891–1905) described a large number of species, mainly sea stars and sea
urchins of the Panamic province (Baja California to Peru) (Agassiz 1881, 1888).
H.L Clark (1916) published a study of asteroids of this region. Some major
expeditions visited the shores of the eastern Pacific Ocean in search of tropical
marine fauna, visiting specific places along the Colombian Pacific Coast, and gave
some records of echinoderm species. In particular, the ship ‘‘Saint George,’’ which
visited Gorgona Island in 1924–1925, the ‘‘Askoy Expedition,’’ which visited
Panama, Colombia and Ecuador in 1941, and Velero III (1931–1941) of the Allan
Hancock Foundation, which also visited Gorgona and elsewhere, contributed to
the knowledge of the marine fauna of the Colombian Pacific. These expeditions
resulted in the descriptions of other echinoderm species and important collections
of irregular sea urchins (Clark 1940, 1948) and holothurians (Deichmann 1938).
In the mid-twentieth century, a series of surveys undertaken by the Zoological
Society of New York and led by W. Beebe resulted in publications of echinoderms
of shallow waters from northern Baja California to Panama. In 1943, A.G. Fraser
made a compilation of the history of exploration, with sampling areas, maps, and
information on the cruises of ‘‘Velero III,’’ which focused on the echinoderm
species of rocky habitats.
Taxonomic knowledge of the species of the ETP was also increased by the
work done in Mexico by M.E. Caso between 1950 and 1987 (Caso 1953, 1971,
1974, 1976, 1978a, b, 1979, 1980, 1983, 1987), and by the expedition of the
Smithsonian Institution in 1972, which visited Malpelo Island and listed five
species of ophiuroids and four of echinoids in a paper about sublittoral
communities (Birkeland et al. 1975) and reported the presence of six species of
asteroids (Downey 1975). Hendler (1979) contributed to knowledge of ophiuroid
biology. The synthetic works of Clark (1989), Clark and Downey (1992),
and Maluf (1988) on echinoderms of Central America included published and
unpublished reports of the ETP, with new reports and ecological information on
this group in the area. More recent works on echinoderms of ETP are those of
Pawson (1995), Solis-Marin (1997), and Lessios (2005).
The taxonomic and ecological investigation of Colombian Pacific echinoderms
was done almost entirely after the 1980s. In this decade, a research group from the
Universidad del Valle, Cali, published papers, mainly species lists or descriptive
ecology, for some coastal localities between 1985 and 1999. These include
the papers of Neira and Von Prahl (1986), who listed 20 species for the island of
Gorgona, including 11 new records; Cantera et al. (1987), who reported two
species of ophiuroids (Ophiactis savingny and Ophiothela mirabilis) associated
with the gorgonian Leptogorgia alba; and Cantera and Neira (1987) who collected
for the first time a parasitic mollusk (genus Echineulima) from the carapace and
spines of the sea urchin Centrostephanus coronatus on Gorgona Island.
Neira and Cantera (1988) reported 17 species of echinoderms of Bahía Malaga:
six asteroids, two echinoids, eight ophiuroids, and one holothurian, of which six
were new records for the Colombian Pacific. Pardo et al. (1988) recorded the
presence in the Colombian Pacific of 70 species, with the first analysis of the
descriptive ecology of echinoderms, including distribution and abundance patterns
of species of several habitats of the Pacific coast of Colombia. Von Prahl (1990)
reported 17 species of echinoderms from Malpelo Island. Neira et al. (1992)
reported the presence of 15 species of echinoderms of the PNN Ensenada Utría
(five asteroids, five ophiuroids and five echinoids) and a short description of two
new records for the Colombian Pacific (Mithrodia bradleyi and Encope ecua-
dorensis). Brando et al. (1992) increased the number from Malpelo Island to 18
species. Furthermore, Schoppe and Werding (1996) studied the effect of the
activity of the boring urchin Echinometra vanbrunti upon rocky shores. Maluf
(1991) increased the list of echinoderm species in a paper mentioning 200 species
of the Galapagos archipelago, including 50 species shared with Malpelo Island.
Cantera et al. (2001) reported the echinoderms associated with bioerosion of
coral reefs of Gorgona Island, finding three species of sea urchins: Diadema
mexicanum and Centrostephanus coronatus that were abundant in the reefs, and
Astropyga pulvinata that was very occasional. Arboleda (2002) listed 118 species
of echinoderms in a paper about the status of knowledge of fishes and some
invertebrates of the Pacific coast of Colombia. Toro-Farmer (1998) and Toro-
Farmer et al. (2004) studied the patterns of distribution and bioerosion rate of the
sea urchin Centrostephanus coronatus on Gorgona Island, finding a rate of
bioerosion of ca. 0.19 kg m-2 year-1. Neira and Cantera (2005), in a review of
published and unpublished literature and unpublished data of zoological collec-
tions on the taxonomy and ecology of Pacific Colombian echinoderms, reported 66
species found in shallow bottoms at 10 localities on different substrates confirmed
by presence in collections or by valid scientific data. Ortiz (2006), whose work was
based on material in national collections, studied the variations of some ossicles of
holothurians of both Caribbean and Pacific coasts of Colombia and contributed to
an improvement of the descriptions of species.
In the Management Plan of the Marine Protected Area of Malpelo (2005–2009),
109 species of echinoderms were included. These species had been recorded since
1975 by various institutions and unofficial sources from shallow and deep water
(up to a depth of 1000 m) near the island. Cohen-Rengifo (2007) and Cohen-
Rengifo et al. (2007) increased the number of species of Malpelo Island to 84.
Fifty-six percent of these species occur at depths of more than 200 m, Defined as
such, we can consider that the number of species surrounding Malpelo Island are
44, the other are of the Oceanic Pacific Ecoregion (PAO) defined as oceanic zones
with depths of more of 200 m.
5.3 Diversity of Echinoderms and Distribution
5.3.1 Geographical Distribution of Species of Echinoderms

in the Caribbean Sea
In total, 259 species of echinoderms have been reported from the Colombian
Caribbean Sea. These species have been collected on the coasts from depths of 0 to
940 m in all ecoregions that comprise the Colombian Caribbean Sea that were
defined earlier. The species that have been studied belong to five classes of
echinoderms. Ophiuroidea is the most diverse with 80 species, followed by
Asteroidea (67 species), Echinoidea (49 species), Holothuroidea (44 species) and
Crinoidea (22 species).
Number of Species Per Class Per Caribbean Ecoregion
In the Colombian Caribbean Sea, ARCO and TAY are the ecoregions with the
highest number of recorded echinoderms, 172 and 158 species respectively. This
may be the result of the greater diversity of ecosystems present and the greater
number of studies carried out in these ecoregions. Furthermore, MAG (134 spe-
cies), GUA (122 species) and DAR (115 species) are also represented by a high
number of species, whereas PAL (82 species), MOR (54 species), and SAN
(45 species) have a lower number of recorded species. This is consistent with the
smaller number of studies in these ecoregions (Fig. 5.4).
Ophiuroidea has more species in PAL, TAY, MAG, ARCO and DAR than the
other classes. In some zones, Asteroidea is better represented, such as GUA and
MOR, where a high number of species of this class is present.
Fig. 5.4 Total number of echinoderms and number of species per class in each ecoregion of the
Caribbean Colombian coast. The total number of echinoderms in each ecoregion is presented in
the white square and is reflected in the size of the graphic. Details of the number of species per
class in each ecorregion are in Table 5.2 (System Information Laboratory LABSIS-INVEMAR)
Table 5.2 Total number of species and species per ecoregion of the Colombian Caribbean Sea
Ecoregion Crinoidea Asteroidea Ophiuroidea Echinoidea Holothuroidea Total
GUA 15 41 34 20 12 122
PAL 8 27 29 11 7 82
TAY 16 40 54 29 19 158
MAG 9 32 43 29 21 134
ARCO 17 30 56 44 25 172
MOR 7 22 15 8 2 54
DAR 10 28 33 26 18 115
SAN 2 9 9 10 15 45
Total 22 67 80 51 44 264
GUA Guajira, PAL Palomino, TAY Tayrona, MAG Magdalena, ARCO Coralline MORArchi-
pelagos, DARMorrosquillo, Darién, SAN San Andrés and Providencia Archipelago
Distribution of Species by Habitat
The ecoregions of the Colombian Caribbean Sea include different kinds of habitat,
such as seagrass beds, sandy beaches, salt marshes, mangroves, coastal lagoons
and swamps, sandy bottoms, rocky shores, coral reefs, estuaries, and soft bottoms.
Echinoderms could inhabit each of these ecosystems. However, only some of them
have been studied and species of echinoderms have been reported in only some of
them. Below are descriptions of the habitats and the research that includes
information on echinoderms.
Intertidal and Shallow Water Ecosystems
Most studies conducted on shallow water echinoderms of the Colombian Carib-

bean Sea have been developed in the ecoregion TAY, specifically in Tayrona
National Natural Park (PNNT), Punta Betín, and Santa Marta Bay, and in the
ecoregion ARCO in Rosario Corals and San Bernardo Islands National Natural
Park (PNNCRSB). Among these studies are those by Gallo (1985, 1988a, b), who
collected 43 species of Asteroidea, Ophiuroidea and Echinoidea at depths from 0
to 30 m at PNNT; Meyer (1972), who described the new crinoid species Cten-
antedon kinziei from Punta Betín and Santa Marta Bay; and Caycedo (1978, 1979),
who published lists of echinoderms found in PNNT and PNNCRSB from at depths
from 0 to 38 m. There are also some lists of echinoderms published by Álvarez
(1981), who recorded 70 species of all echinoderm classes and Allain (1976), who
recorded 38 species of echinoids of all ecoregions the Colombian Caribbean Sea.
It is important to note the study made by Quiñones (1981), who listed 23 species of
echinoderms from the SAN ecoregion, specifically in Providencia and Santa
Catalina Islands.
Rocky Shores
In the Colombian Caribbean Sea, most of the rocky shores are distributed in a
discontinuous line along the continental coast and also around some islands.
Generally they are interrupted by beaches, coastal lagoons, and estuaries (Batista-
Morales and Diaz-Sanchez 2010). Rocky shores in the Colombian Caribbean Sea
have been little studied, so studies focusing on the echinoderms are also scarce.
Gárces and Romero (2003) carried out research with the aim of characterizing the
structure of the ophiuroid community associated with the mesolittoral rocky shore
of Gaira bay (TAY ecoregion), determining spatial-temporary changes of the
community between August, 2001, and May, 2002. As a result, 12 species were
identified. The most frequent and abundant were Ophiactis savignyi and Ophiothrix
angulata, and the least frequent and abundant was Ophiocoma wendtii. Monroy
(2003) and Monroy and Solano (2005, 2008) developed an ecological study
focused on evaluating the distribution of populations of the Colombian endemic
species Ophiothrix synoecina and its host Echinometra lucunter and other
accompanying fauna of this association. They found a broad distribution of
E. lucunter throughout the Caribbean coast (DAR, ARCO, TAY, GUA), different
from the restricted distribution of O. synoecina, which is found only on northern
Colombian rocky shores (TAY and GUA ecorregions). The results of this research
showed a significant reduction in the density of the sea urchin and its associated
fauna, including O. synoecina. Specifically, the frequency of occurrence of
O. synoecina has declined about 50 % in recent years in the Santa Marta area
(TAY), in contrast to population studies made by Schoppe (1993) in the same area.
Recently, Romero (2009) developed a study in Bahía Concha (PNNT) in the
ecoregion TAY, where was established the population dynamics of E. lucunter in
the rocky shore ecosystem and was measured the spatial–temporal variation with
respect to size and density. The results indicated that environmental conditions
could be affecting population density rocky shores.
Coral Reefs
Coral reefs have been intensively studied in Colombia, focusing on mapping,

zonation, health status, and the associated fauna and flora. Twenty-one coral areas
of the Caribbean are broadly distributed. 80 % of them are in the SAN ecoregion,
where they are also more developed and complex. The other areas are scattered
along the continental coast, mostly within ARCO and TAY (Díaz et al. 2000). As
mentioned above, the populations of seven species of sea urchins have been
monitored in the Colombian Caribbean Sea through the National System of
Monitoring of Coral Reefs (SIMAC). It was found that changes in the populations
of sea urchins could be determined by the genus Echinometra, especially E. viridis.
Although is not possible to define the causes of the annual variation of the density
of sea urchins recorded, the patterns could be the result of factors such as predation,
recruitment, inter-specific interactions, and environmental and weather changes.
Although there is no specific information about Colombian Diadema antillarum
populations before their massive mortality (Lessios et al. 1984; Murillo and Cortés
1984; Liddell and Ohlhorst 1986; Hunte et al. 1986; Carpenter 1988; Carpenter
1990; Hendler et al. 1995), González-Gaviria (2010) showed that in the PNNT and
Playa Blanca-Rodadero (Santa Marta) there has been a slow and irregular recovery
in comparison with other Caribbean populations.
Among echinoderms studies associated with Colombian Caribbean coral reefs
it’s important to mention Bejarano et al. (2004), who characterized the ophiuroid
community in the microhabitats provided by sponges and other reef components at
San Bernardo Archipelago (ARCO). The community was composed of 22 mor-
photypes, including: Ophiothrix angulata, O. oerstedii and O. suensoni. On the
other hand, Gómez and Rodriguez (2009) studied the ophiuroid community
associated with sponge populations in Cispata Bay (MOR). They described the
different kinds of associations between ophiuroids and sponges and identified 13
and 12 species of echinoderms and poriferans respectively.
Sandy Beaches
Nearly 60 % of the 1,650 km of the Colombian Caribbean coast consists of sandy

beaches. Studies of these beaches have focused mainly on the zonation patterns of
the faunal communities at a few specific sites. In the Caribbean, beaches with
coarse sand exposed to wave action, which occur largely in the northern half of the
coast, have a more diverse community dominated by bivalves of the genus Donax
and ‘‘sand dollars’’ such as Mellita sp. A specific echinoderm study was published
by Arevalo (1978), who observed the biological and ecological aspects of Mellita
quinquiesperforata populations on public beaches from Bocagrande to La Boquilla,
Cartagena (MAG ecoregion). He noted that there are morphometric and behavioral
variations with respect to other coastal populations of the Atlantic, considering the
species’ distribution, which is from the coast of Massachusetts to southern of
Brazil.
Mangrove Areas
Mangroves are among the most biologically productive ecosystems in the world
and play an important role in the sustainability of fisheries. They protect the
shoreline against erosion, and provide an important resource of wood. Five species
of mangrove have been reported in the Caribbean; Avicennia germinans and
Rhizophora mangle are the most common and abundant (Orjuela et al. 2010).
Unlike along the Pacific coast, the Caribbean mangroves are limited to narrow
sections within lagoons, swamps, estuaries, and river mouths. The major coverage
of mangroves is in Ciénaga Grande de Santa Marta (CGSM) in the ecoregion
MAG (Orjuela et al. 2010). Echinoderm studies of this habitat are few. Báez
(2001) characterized a community associated with roots of R. mangle in the
Cispatá bay (ecoregion MOR), where two species of ophiuroids were found:
Ophiothrix oerstedii and Ophiactis savignyi.
Seagrass Beds
Seagrass beds are distinct and unique ecosystems with relatively high rates of
primary production which support detritus-based food chains. In Colombia, they
occur solely in the Caribbean. The seagrass ecosystems provide habitats for a wide
variety of marine organisms, promote sediment stabilization, and maintain a
dynamic environment for nutrient cycling (Díaz et al. 2003). Seagrass beds and
their associated communities were rarely studied in Colombia until very recently.
Díaz et al. (2003) developed an integrative study of Colombian seagrass beds,
including composition and distribution, community structure, and environmental
problems. As a result of this research, more than 550 species of fishes and
invertebrates and about 35 species of macroalgae were recorded in the seagrass
beds of Colombia (Montoya-Maya 2002). Seventeen species of echinoderms have
been found in these beds. Tripneustes ventricosus and Lytechinus variegatus have
been frequently observed, especially in the ecorregions SAN, GUA, TAY and
ARCO. In some areas particularly in the ecoregions GUA and ARCO, populations
of Oreaster reticulatus are frequent. Specific studies focused on echinoderms
associated with this ecosystem are few. Moreno-Forero and Hernández-Torres
(1992) published an ecological study of size composition of L. variegatus asso-
ciated with sea grass beds of Thalassia testudinum in Isla Grande–Islas Rosario
(ARCO ecoregion). The size composition and some morphometric relations are
described, and some ecological notes on the population are given.
Soft Bottoms
Soft bottoms are extensive marine ecosystems, at depths from 0 to more than
10,000 m. They predominate in the territorial sea of the Colombian Caribbean and
Pacific Oceans, with an extension of 889.400 km2, comprising 99.5 % of under-
water ecosystems (Guzmán-Alvis and Solano 2002). Soft bottoms can include
sand, clay, silt and mud particles, are not stable substrates, and only form low
topographic features (Montoya-Cadavid 2010). Data collected from 1998 to 2011
show increasing of knowledge of this ecosystem in Colombia. Taxonomic studies
have concentrated on the Guajira (GUA), Tayrona (TAY), Magdalena (MAG), and
Oceanic Caribbean (COC; depth [ 200 m) ecosystems (INVEMAR 2010). Most
of the specific echinoderm studies on this ecosystem, has been developed at depths
between 20 and 1,000 m, which are mentioned in the next section on deep water
ecosystems. The only study in shallow soft bottoms was published by Vélez
(2003), who studied some structural aspects of the ophiuroid community associ-
ated with infaunal bottoms in shallow waters of the San Andrés and Providencia
Archipelago (SAN ecoregion). During this research, that identified 15 species of
ophiuroids, two climatic periods were included during 1 year (1999–2000).
Deep Water Ecosystems
Research in deep water ecosystems has been conducted throughout the Colombian
Caribbean Sea and not confined to sites like most studies in shallow water. Most
studies have been carried out in the soft bottom ecosystem, which is dominant at
depths greater than 200 m. As a result of exploration in soft bottoms, banks of
deep azooxanthellate coral communities have been found, which are mentioned
below.
Soft Bottoms
For this habitat in deep waters, it is important to consider the marine explorations
made before 1998 by international and national investigations focused especially
on depths between 200 and 1,000 m. Among them are the explorations made by
the R/V Pillsbury in 1966–1968, whose results were presented by Bayer et al.
(1970). These authors listed more than 100 species of echinoderms collected
between Cartagena and the gulf of Urabá along the ecoregions DAR, MOR,
ARCO and MAG from 70 to 3,000 m depth. The results of other explorations,
carried out by the INVEMAR-CIOH-SMITHSONIAN in 1995, were published by
González et al. (2002), who listed 22 echinoderm species collected in shallow and
mid waters (30–380 m), between Cartagena and the gulf of Urabá in the same
ecoregions, but including some localities in the PNNCRSB (ARCO). However, the
most complete information about echinoderm communities of soft bottoms is from
explorations carried out between 1998 and 2009, such as Macrofauna I-II, Mac-
rofauna-CORPOGUAJIRA, Macrofauna-MARCORAL, and ANH I-II. In these
explorations, 188 species of echinoderms were collected, of which 81 were
recorded for the first time in Colombian Caribbean waters; Ophiuroidea was the
most abundant and richest class. In addition, an ecological analysis using multi-
variate and univariate statistics was undertaken, including all echinoderm records
collected during those biological expeditions. The results of this analysis showed
four assemblages defined by depth: the first, in shallow waters (depth of 20–70 m),
was characterized by Astropecten antillensis and Ophiura acervata. Intermediate
waters (depth of 150 m) were characterized by the clypeasteroid Clypeaster eu-
clastus. The third, to a depth of 300 m, had Caryometra atlantidis and Amphiura
rathbuni as characteristic species. The last conglomerate, between depths of 500
and 940 m, with the brittle star Ophiernus adspersus adspersus and the sea star
Cheiraster (Chiraster) planus as the characteristic species. According to the
results, the first change in faunal composition of the Colombian Caribbean Sea
occurs between depths of 150 and 300 m. An interesting result is that in the
conglomerates from depths of 300 and 500 m, a different faunal composition was
observed between the samples located in the northeastern Colombian Caribbean
Sea and those to the southwest with respect to the Magdalena River (Borrero-Pérez
et al. 2004; Benavides-Serrato and Borrero-Pérez 2010).
Deep Azooxanthellate Coral Community Banks

Based on results of the Macrofauna I and II explorations (1998–2002) carried out
by INVEMAR, there are deep azooxanthellate coral banks in three localities of the
Colombian Caribbean Sea: off La Guajira Peninsula (PAL ecoregion), off Santa
Marta (TAY ecoregion), and off San Bernardo Islands (ARCO ecoregion)
(Fig. 5.5, Table 5.3). Cladocora debilis (thin tube coral) was the main coral matrix
builder in the bank formation off La Guajira, at a depth of 70 m. Echinoderms
were the most abundant taxon collected in these beds, with 1,367 specimens. The
crinoid Analcidometra armata was the dominant echinoderm, which has been
reported as a common species attached to gorgonians in the Caribbean Sea (Meyer
1973; Hendler et al. 1995). Ophiothrix angulata was found in several stations
along the sampling area. It has also been reported on Oculina varicosa banks off
Fig. 5.5 Representation of the total number of echinoderms and number of species per class
within each deep azooxanthellate coral bank from north to south along La Guajira Peninsula
(PAL), Santa Marta (TAY) and San Bernardo Islands (ARCO) (Reyes et al. 2005). The total
number of echinoderms in each bank is presented in the white square and is reflected in the size of
the graphic; details of the number of species per class in each bank are in Table 5.3 (System
Information Laboratory LABSIS-INVEMAR)
Table 5.3 Number of species per class on each deep coral setting of the Colombian Caribbean
Sea
Class Deep coral bank
GUA SMA SBE
Crinoidea 5 1 5
Asteroidea 1 7 1
Ophiuroidea 11 12 16
Echinoidea 6 5 8
Holothuroidea – 2 1
Total 23 27 31
GUAi Guajira, SMA Santa Marta, SBE San Bernardo
Florida (Reed and Mikkelsen 1987). Species found in the bank formation off Santa
Marta, at a depth of 200 m, included Ophiothrix suensoni, commonly found on
gorgonias (Hendler et al. 1995) and the sea cucumber Holothuria lentiginosa
enodis, previously recorded off Florida from Oculina varicosa banks (Pawson
et al. 1982). Another 14 species were found exclusively in this area. Three of them,
Stephanasterias albula, Trigonocidaris albida, and Palaeobrissus hilgardi, were
the first records for the southern Caribbean. The bank formation off San Bernardo,
at a depth between 155 and 160 m, was the most diverse sector for echinoderms,
with 38 species. Brittle stars such as Ophioderma appressa, Astrocnida isidis,
Asteroporpa (Asteroporpa) annulata, and Asteroschema oligactes were found
attached to stony corals and gorgonians. Also, some rare species, such as Oph-
iosyzygus disacanthus, previously recorded as dwelling in hard bottoms (Turner
and Heyman 1995; Borrero-Pérez and Benavides-Serrato 2004b), the stalked cri-
noid Democrinus conifer, and the reef dwelling comatulid crinoid Davidaster
rubiginosus were present. These three unique azooxanthellate coral communities
sustain one of the highest species diversities found along the Colombian conti-
nental shelf (Reyes et al. 2005).
5.3.2 Geographical Distribution of Species of Echinoderms

in the Colombian Pacific Ocean
One hundred and twenty-four species of echinoderms have been reported from the
Pacific coast of Colombia. These species have been reported from the coast of
Chocó (Punta Ardita, Cabo Marzo and Gulf of Cupica, Utria Bight), Valle del
Cauca (Malaga bay, Buenaventura bay, Gulf of Tortugas), Cauca (mouth of the
Guapi River), Nariño (Mulatos Beach, Tumaco Bight), Gorgona, and Malpelo
Island. The species reported are Asteroidea (34 species), Ophiuoroidea (30),
Echinoidea (28), and Holothuroidea (32). There are no reports of Crinoidea.
Echinoids and asteroids are the classes with highest species richness. Ophiuroids
have the highest number of individuals.
Number of Species Per Class Versus Ecoregion in the Colombian Pacific

Ocean
The localities with most species are the deeper parts of the oceanic Pacific (52),
Malpelo (44), and Gorgona (41 species), and deeper parts of the oceanic Pacific
(52) (Neira and Cantera 2005; Cohen-Rengifo 2007). The other area with a high
number of species is Buenaventura (31) (Fig. 5.6, Table 5.4). As in the Caribbean,
areas with a high number of studies have more species reported. Some studies have
shown that deep species can be very abundant.
Fig. 5.6 Representation of the number of echinoderms and number of species per class in each
ecoregion of the Pacific Colombian coast. The total number of echinoderms in each ecoregion is
presented in the white square and is reflected in the size of the graphic; details of the number of
species per class in each bank are in Table 5.4 (System Information Laboratory LABSIS-
INVEMAR)
Distribution of Species by Habitat
Because of the abundance estuaries and river discharges, echinoderms are not
abundant along a large part of the Pacific coast of Colombia coast. Malpelo Island
has been studied intensively (Cohen-Rengifo 2007; Cohen-Rengifo et al. 2007).
These authors reported 84 species of echinoderms occurring around this island,
including species of deeper adjacent areas attributed here to the PAO region.
Table 5.4 Number of species per class in each ecoregion of the Colombian Pacific Ocean
Ecoregion Asteroidea Ophiuroidea Echinoidea Holothuroidea Total
PAN 6 4 6 1 17
BAU 5 5 4 2 16
BUE 9 12 8 2 31
NAY 8 1 4 1 14
SAQ 2 0 4 0 6
TUM 0 4 2 2 8
GOR 7 13 10 11 41
PAO 11 11 14 16 52
MAL 15 9 10 10 44
Total 34 30 28 32 124
Pacific North (PAN), Baudó (BAU), Buenaventura (BUE), Naya (NAY), Sanquianga (SAQ),
Tumaco (TUM), Gorgona (GOR), Malpelo (MAL) and Pacific Ocean (PAO)
Gorgona island (GOR ecoregion) also has a high number of records (41 species),
although it has been relatively less sampled. Among the coastal localities, the
tectonic estuary of Málaga Bay (BUE ecoregion) has the highest number of species
(25), followed by the localities along the north Pacific coast (e. g. Utría Bight,
19 species). All these localities are characterized by rocky areas as cliffs, shores,
and shallow bottoms. All localities also have high salinity (30 PSU or more).
In contrast, estuarine areas of the southern Pacific coast of Buenaventura Bay
(BUE), Tumaco Bay (TUM), the mouth of the Guapi River (NAY), and Mulatos
Beach (SAN), have a low number of species. But these species are always asso-
ciated with the external zones of bays or estuaries. The low number of species in
these areas is the result of estuarine conditions (low salinity, high turbidity, and
sedimentation) and rarity of suitable rocky substrate.
Most species of echinoderms were found in more than one habitat. Rocky
substrates and coral reefs share some species of the cryptofauna, because of the
similarity of cavity structure of both rocky shores and coral framework. It is also
possible that physically contiguous habitats, such as cliffs and rocky shores, or
sandy beaches and mud flats also share some species. The moisture that remains in
some intertidal substrates during low tide allows the existence of some subtidal
species.
The species and the most important characteristics of groups of echinoderms of
different habitats are presented below.
Rocky Cliffs
Tectonic processes along the Pacific coast of Colombia have resulted in abundant
steep cliffs and to rocky shores with sloping platforms composed of boulders,
pebbles, and gravel formed by cliff erosion. In the northern basaltic cliffs the rocks
are igneous, very hard, and consolidated. In the southern part most rocks are
sedimentary (mudstones and shale), subject to rapid erosion by living organisms
(bioerosion). These cliffs have nearly vertical walls that fall directly into the sea or
may be separated from it by sandy beaches. Most of these are located in areas
exposed to strong waves and high hydrodynamic conditions. The erosion forms
caverns and abrasion terraces, which are inhabited by sea urchins of the family
Cidariidae (Eucidaris thouarsii and Hesperocidaris asteriscus), Diadematidae
(Centrostephanus coronatus), and Echinometridae (Echinometra vanbrunti). Some
ophiuroids can be found occasionally (Ophioderma teres, O. panamensis, and
Ophiocoma alexandri).
Rocky Shores
This habitat is located along the whole Pacific coast. It is characterized by igneous
and sedimentary rocks at different stages of fragmentation and by variability of
environmental conditions. Rocky platforms are covered with hard blocks of var-
ious sizes (20 cm to 1 m in diameter), pebbles (5–10 cm), or gravel (2–5 cm) or
are partially covered by sand or mud. In some cases these platforms only emerge
during the minimal astronomic tides. This ecosystem is one of the richest in
species, with 23 species (two asteroids, seven echinoids, nine ophiuroids, five
holothurians). The most common species of rocky shores, living under rocks or in
cavities or holes, are: Phataria unifascialis, Pharia pyramidata, E. thouarsii,
H. asteriscus, D. mexicanum, C. coronatus, Echinometra vanbrunti, O teres,
O. panamensis, Ophiothrix (Ophiothrix) spiculata (associated with clusters of
polychaetes and sponges), Ophiocoma aethiops, Ophionereis annulata, Ophiactis
savignyi, and Holothuria (Cystipus) inhabilis. O. savignyi and Ophiothela mira-
bilis are found associated with the gorgonian Leptogorgia alba (Neira and Cantera
2005).
Coral Reefs
The coral areas are found mainly in some localities of the Chocó (Cabo Marzo,
Cupica Gulf, Utría Bight) and the islands of Gorgona and Malpelo (Zapata and
Vargas-Ángel 2003). The major coral reefs are found on the island of Gorgona,
where there are true reefs 2 or 3 km long, and some coral patches with species of
scleractinian corals of the genera Pocillopora, Pavona, Porites, Psammocora.
These reefs have four areas: the backreef, comprising sandy areas and a platform
with dead coral and few living colonies (depth between 1 and 3.5 m); a platform-
crest region, with the highest coralline coverage (but largely dead) and dominated
by Pocillopora coral (depth between 0.5 and 3 m); a reef front, with a decreased
coverage of Pocillopora and an increased number of massive coral colonies (depth
between 1 and 4 m at low tide, reaching 8 m at high tide); and finally, a reef slope,
depth to 12 m, characterized by low coral coverage, and some dead colonies
scattered on the sandy bottom. Patches also occur, with massive and branching
corals (Cantera et al. 2001; Zapata 2001). Malpelo Island has coral colonies
growing on steep rocky platforms, with Pocillopora species having a different

external morphology in response to the slope of the substrate and the strength of
the currents (Brando et al. 1992).
This ecosystem has 32 species (four asteroids, eight echinoids, 11 ophiuroids,
and nine holothurians). In the Gorgona reef, the area with the greatest richness of
echinoderms is the reef front with species of both intertidal and subtidal habitats.
In this ecosystem occur, from the shallow zones to the outer edge, the sea urchins
A. pulvinata, C. coronatus, D. mexicanum, E. vanbrunti and E. thouarsii, between
the coral colonies and in the cavities eroded by them at the base of coral colonies.
Toxopneustes roseus is found in the sandy patches between corals at different
depths. Under the bases of corals and rocks within the reefs are Ophioderma teres,
O. panamensis, Ophiolepis plateia, O. variegata, O. aethiops, O. alexandri,
Ophiocomella schmitti, Ophionereis annulata, and Ophiostigma tenue. Ophiactis
savigny is associated with the bivalves Pinna rugosa and Pinctada mazatlanica),
and O. spiculata is associated with the bivalve Pinna rugosa.
The Asteroidea of the reef areas are Pentaceraster cumingii and Mithrodia
bradleyi, found in the deeper parts of the reef. Cohen-Rengifo (2007) and Narváez
and Zapata (2010) have reported Acanthaster plancii from the Malpelo reefs, and
some unpublished data record it from Gorgona Island. Finally, in the reefs there
are holothurians: Holothuria (Thymiosycia) arenicola, H. (Platyperona) difficilis,
H. (Stauropora) fuscocinerea, H. (Mertensiothuria) leucospilota, H. (Thymiosycia)
impatiens, H. (Mertensiothuria) hilla, H. (Lessonothuria) pardalis, Labidodemas
maccullochi, H. (Cystipus) inhabilis and Pentamera chierchia. All these species
are found at the base of the colonies or out (at night) in the sandy patches.
Sandy Beaches
Sandy beaches of the Pacific coast of Colombia are intertidal zones formed mainly
by sediments from the discharge of rivers. These beaches have an upper area,
emerged most of the time and covered only at high tides and a lower area that
remains submerged much of the time. These beaches may consist of siliceous
materials of medium size (between 1 and 2 mm) mixed with silt and clay brought
from land or mangrove areas through runoff (Von Prahl et al. 1990). This habitat is
associated with external areas of bays which can be subject to strong wave action.
The nine species found on these beaches are irregular urchins of the family
Mellitidae. Their presence at these beaches is seasonal. Very large numbers of
individuals (between 9 and 20 m-2) are found between September and November
and are almost absent at other times (Cantera 1991). The species involved are
Encope insularis and Mellita longifissa, which migrate from the sublittoral zone
(submerged up to a depth of 16 m) to the upper intertidal zone, in places where
tidal changes maintain wet conditions, such as hydraulic dunes characterized by
presence of ripple marks.
Mangrove Areas
Echinoderms are virtually absent from the mangroves on the Colombian Pacific
coast, except for a few records of the ophiuroids Amphiodia oerstedii, Microp-
hipholis geminata, and A. squamata, which were found under hard objects on mud
flats, near the roots of red mangrove (Rhizophora sp.), outside of the Bay of
Buenaventura.
Submerged Rocky and Sandy Substrates
These bottoms are located within bays or on the continental shelf of the Colombian
Pacific area. Most sandy and muddy bottoms are enriched with abundant organic
matter from coastal mangroves and rivers, deposited between depths of 5 and 20 m
along the coast. The most abundant species found in these submerged muddy and
organic substrates are Astropecten armatus, A. verrilli, A. regalis and A. brasili-
ensis peruvianus, Luidia columbia,, L. superba, and Luidia latiradiata. Sandy
bottoms are inhabited by the echinoids Encope laevis and E. micropora
ecuadoriensis.
On submerged rocky bottoms located near Gorgona and Malpelo Islands and
the northern part of the coast (Chocó) the most abundant species are the asteroids
Nidorellia armata, P. unifascalis, P. pyramidatus, the ophiuroid Ophiothrix
spiculata (on sponges), the echinoid Echinometra vanbrunti, and the holothurians
Holothuria portovallartensis and Isostichopus fuscus.
5.4 Aquaculture, Fisheries and Environmental Threats
In the Colombian Caribbean Sea, echinoderms are not included in the diet of native
people. However, some marine resources such as species of sea cucumber are caught
for export. The over-exploitation of sea cucumbers in areas traditionally fished
(Asian countries) has resulted in drastic population declines and even local
extinctions. This has led to the expansion of this fishery in the world, including the
Caribbean, as well as an increase in the number of exploited species. In Colombia,
the first documented record of this fishery is in 2006 (Borrero-Pérez et al. 2010),
when there was a massive collection, mainly of Isostichopus badionotus and
Holothuria mexicana, in the area of Portete (Guajira). However, there are records of
the export of dried sea cucumber (0.55 t) from Colombia to the Asian market in the
year 2001 (Bruckner et al. 2003). In 2005, resolution 2467 of INCODER authorized
two exploratory fishing permits in the Guajira. In none of these cases have the
harvested species or biomass been reported. Recently, in 2008 and 2009, there were
several catches of sea cucumbers in the area of Santa Marta, including the species
Holothuria glaberrima, Holothuria grisea, and Ocnus suspectus (Benavides-Serrato
2009). Despite the ecological and economic importance of these invertebrates, there
is a lack of information in Colombia and in most Caribbean countries about their

biology, population dynamics, and fishery, laws (legislation) for their management
are lacking. In Colombia, interest in this resource is growing, and studies on the
reproductive biology of some species (e.g. H. glaberrima) and on the potential of
aquaculture are beginning. However, the current status of natural populations
remains unknown, while exploitation continues. At the last workshop organized by
the IUCN in Cartagena de Indias, in June 2010, I. badionotus and H. mexicana were
mentioned as vulnerable species. However, due to lack of information, these species
were placed in the insufficient data category.
Sea cucumbers were not considered in the Red Book of marine invertebrates of
Colombia published in 2002. At this time the only three species evaluated and
analyzed were Diadema antillarum, Ophiothrix synoecina categorized as Deficient
Data (DD), and Oreaster reticulatus categorized as Least Concerning (LC). Dia-
dema antillarum suffered massive mortality in 1983 due to an unknown pathogen,
which spread throughout the Caribbean Sea, causing a mortality of over 97 % of
mature individuals. This trend in the Caribbean Sea was also seen in Colombia but
was not documented. Even today, there are no detailed data on the population
dynamics of this species. However, González-Gaviria (2010) found that abun-
dances appeared to be lower compared to pre-mortality levels, indicating that as in
the rest of the Caribbean, recovery in this region has been slow. Oreaster retic-
ulatus is especially vulnerable to human exploitation due to its size, its slow
locomotion and easy access to its habitat. Individuals of this species, dried and
painted in bright colors, have been sold as decorative objects on beaches for many
years, including Colombian beaches. Because of overexploitation, O. reticulatus is
now considered a rare species in some localities of the Caribbean Sea where
formerly it was very common. In Colombia this species is still common in some
areas. However, there is insufficient information to assess the dynamics of their
populations and there are no regulations for their protection. Ophiothrix synoecina
was included in the Red Book because it has a very limited geographical distri-
bution, being endemic to rocky shores in the area of Santa Marta and Tayrona
National Park (Schoppe and Werding 1996). There has been no recent information
on its populations. It is now known that the species is only present on the rocky
shores of Santa Marta and the Guajira and the density of its populations has
declined significantly since 1996. Thus a change in conservation status to vul-
nerable (VU) has been suggested (Monroy and Solano 2008).
Echinoderms are not used as food by the Pacific coast inhabitants, and for this
reason they are not fished or cultivated. Similarly, there are no reports (even oral)
of commercial fishing for echinoderms. Some species are threatened by incidental
capture (Astropecten, Luidia) by small trawling nets. Most populations are not
impacted by anthropic factors. Some species (Encope and Mellita in sandy beaches
and D. mexicanum and probably C. coronatus in coral reefs) undergo great vari-
ations in number of individuals, as was observed in other echinoderm populations
of the world (Uthicke 2009), displaying cyclic population dynamics, probably in
relation to specific seasons, or climatic conditions. In some places, populations
may decrease because of human activities in coastal areas. These activities can
increase sedimentation rates, changing rocky substrates to sand or mud and con-
sequentially, change the relative abundance of different species affect assemblages
composition. A conservation status assessment is difficult because of the lack of
periodical monitoring and insufficient data from deeper waters.
5.5 Recommendations
The advance in the study of echinoderms has been notable in recent years, mostly
at species level in deep habitats, specifically soft bottoms. Interesting results,
including the discovery of new habitats, show the need to continue with these
investigations, particularly at depths greater than 1,000 m. This requires improved
sampling techniques than are currently being used, which are less destructive and
more efficient.
Although in 1998 the largest amount of information about echinoderms was on
the shallow-water species, during recent years, research has been focused on
deeper ecosystems, which has increased our knowledge of these zones, mainly in
the Caribbean, while there has been little recent research in shallow ecosystems.
It is now clear that more taxonomic and ecological studies are needed of these
invertebrates in shallow areas in the Caribbean and of deeper zones in the Pacific.
Although there has been a significant increase in ecological studies of echi-
noderms in Colombia, there still is little information about this important group of
invertebrates. Further studies are necessary especially considering the commercial
interest that is being focused on some species such as sea cucumbers; the current
status of their natural populations and their dynamics is not known.
Building on all the information that has been collected during recent years and
specimens that have been deposited in the echinoderm collections of MHNMC and
the Universidad del Valle, future research could utilize new tools such as
molecular techniques to investigate cryptic speciation, population connectivity,
species adaptation and conservation biology, considering the privileged location of
Colombia to answer questions about the evolution of species in the Americas and it
high biodiversity and resources to conserve and use sustainably.
Although studies on Colombian Pacific echinoderms have been carried out for
many years since the initial international expeditions that found the first species
and in recent years have increased, knowledge of this group in this region is still
incomplete. Taxonomy and descriptive ecology of the species of coastal ecosys-
tems of the mainland and islands are relatively well known, and there are abundant
but not complete references for many species of some deep areas. For these
reasons it is necessary to continue to expand the inventory of species, visiting
poorly known locations on the coast (i.e. northern Chocó), or increasing the
coverage of oceanographic cruises that explore the offshore bottoms of the Pacific
Ocean in search of benthic fauna.
Similarly, although there is some biological data for some groups (mainly
asteroids and echinoids), it is necessary to increase research on species richness
and diversity, ecological studies that include structure of assemblages of species,

distribution, population densities, and breeding seasons. These biological charac-
teristics must be related to the environmental and oceanographic conditions of the
study areas (e.g. coastal and marine National Natural Parks, such as Ensenada de
Utría, Bahia Malaga, and Gorgona and Malpelo Islands).
Bioerosion activity caused by some sea urchins has been studied around
Gorgona Island, but there is no information on bioerosion rates and the causes of
greater or lesser bioerosion action. Similarly, further studies are necessary to
explain the causes of important and sudden variations in the populations that occur
in some beach echinoderms (Encope and Mellita) and coral reefs (Diadema,
Centrostephanus and Holothuria) at certain times of the year. It is also important
to study these variations in relation to oceanographic features that affect the
Eastern Tropical Pacific, particularly El Niño phenomenon that apparently
increased populations of some echinoderms in Gorgona Island in 1982 and 1997.
Finally, although there is no commercial exploitation of echinoderm species in
the Colombian Pacific, it is necessary to initiate studies to establish the potential of
some species that are exploited in other regions. It may also be important to initiate
preliminary assays of culture of the potentially important species.
Acknowledgments The authors wish to express thanks to J.J. Alvarado for the invitation to
participate in this synthesis on the Latinoamerican echinoderms. The results presented in this
chapter have been produced by different kinds of research projects on both Colombian coasts. We
are grateful for the support of the Ministry of Environment, Housing and Territorial Development
(MAVDT), the Administrative Department of Science, Technology and Innovation
(COLCIENCIAS), the Autonomous Regional Corporation of La Guajira (CORPOGUAJIRA),
and the National Hydrocarbons Agency (ANH), among others, all of them with the participation
of INVEMAR. For the Pacific coast, the projects ‘‘The relationship between bioerosion and
growth rates in Colombian Pacific coral reefs of the island of Gorgona (project UNIVALLE
1106-09-069–96), Biomalaga (project INVEMAR-UNIVALLE INCIVA 210509-16821), and
Biodiversity of vulnerable stages of benthic fauna of Bahía Malaga (project UNIVALLE-IN-
VEMAR 1106-405-20155) were logistically and financially supported by the Universidad del
Valle and COLCIENCIAS with the participation of INVEMAR. We thank G. Hendler, D.Paw-
son, C. Ahearn, and C. Mah, C. Messing, R. Mooi, J. Lawrenc, and F. Solís-Marín, who have
contributed permanently to the echinoderm collection of the Universidad del Valle (CRBMUV),
and the Colombian Museum of Marine Natural History MHNMC-INVEMAR through confir-
mation of species identification, advice, and literature. We also would like thank Colombian
researchers dedicated to taxonomy, especially G. Navas and O.D. Solano. Finally, we would like
to thank J. Espriella and R.Pinzón Corredor, members of the System Information Laboratory
LABSIS-INVEMAR for preparing the maps elaboration in this chapter and P. Silverstone-Sopkin
for correcting the English grammar.
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Chapter 6
Echinoderms of Ecuador
Jorge Sonnenholzner, Margarita Brandt, Vanessa Francisco,

Alex Hearn, María Luzuriaga, Paulina Guarderas
and Juan Carlos Navarro
J. Sonnenholzner (&)
Universidad Laica Eloy Alfaro de Manabí, Departamento Central de Investigación,
Manta, Manabí, Ecuador
e-mail: jsonnenholzner@hotmail.com
M. Brandt
Department of Ecology and Evolutionary Biology, Brown University,
Providence, RI, USA
e-mail: margarita.brandt@gmail.com
V. Francisco
Laboratorio de Ecología de Ecosistemas de Arrecifes Coralinos,
Centro de Investigación y de Estudios Avanzados del Instituto Politécnico Nacional,
Mérida, Yucatán, Mexico
e-mail: vanewagen@gmail.com
A. Hearn
Department of Wildlife, Fish and Conservation Biology,
University of California, Davis, CA, USA
e-mail: arhearn@ucdavis.edu
M. Luzuriaga
Laboratorio de Plancton e Ictioplancton, Instituto Nacional de Pesca del Ecuador,
Guayaquil, Ecuador
e-mail: mluzuriagav@yahoo.com
P. Guarderas
GEOcentro, Universidad San Francisco de Quito, Cumbayá, Ecuador
e-mail: pagigv@yahoo.com
J. C. Navarro
LECROY S.A., Km 2.5, Avenida Carlos Julio Arosemena,
Bodega No.5, Guayaquil, Ecuador
e-mail: jcnavarrogo@gmail.com

184 J. Sonnenholzner et al.
6.1 Introduction
6.1.1 Ecuadorian Continental Shelf
Geography and Climate
The continental coastal territory of Ecuador extends 4,403 km from north to south,
including islands, islets, and estuarine borders. The width of the continental shelf
exceeds 100 km, particularly off the Gulf of Guayaquil (associated with the most
important watershed from the Guayas River basin). The influence of the Guayas
River extends over two-thirds of the surface along the continental coast. This
causes coastal sedimentation, and a complex estuarine-deltaic system, that extends
close to the edge of the continental shelf (Benítez 1995 in Morante-Carballo 2004).
Approximately one-third of the coastline in Ecuador consists of stretches of
mangroves on mudflats, with major concentrations along the southern half of the
northern Ecuadorian coast and in the Gulf of Guayaquil. There are substantial
stretches of rocky shores scattered throughout the coast. The longest uninterrupted
areas occur in the central coast of Ecuador. Long stretches of sandy beaches are
mostly concentrated along the northern-central Ecuadorian shorelines. Coral reef
development in this region is prevented by unfavorable conditions that result from
freshwater input from river runoff, siltation, nutrient enrichment, and upwelling
influences. The majority of reef habitat in this region consists of rocky reefs.
Suitable conditions for coral development and persistence are found around islands
and rocky promontories located away from the central mainland coast, such as Isla
de la Plata, Isla Salango, and Bajo Montañita.
According to the classification of climate and life zones proposed by Cañadas
(1983) and Sierra (1999) there is a gradient in rainfall levels from north to south,
with a complex spatial and temporal variation. The northern part is a wet tropical
region (with [2,000 mm yr-1 of rainfall and without ecologically-dry months
throughout the year) and the central and southern regions, range from a dry
tropical desert to a sub-tropical desert (with \1.9 mm yr-1 of rainfall). The
monthly maximum temperatures are higher in the central and northern Ecuador
and decrease to the south, at the entrance to the Gulf of Guayaquil. Overall, the
average annual temperature is above 22 °C. The maximum temperatures fluctuate
between 32 and 38 °C and the minimum is around 15 °C. The relative humidity is
above 80 % and with considerable cloudiness. The rainy season runs from January
to April, with temperatures averaging 28 °C. The rest of year includes short rainy
summers and relatively dry seasons.
Oceanography
Ecuador is located at the confluence of two major marine ecosystems: the Pacific
Central American Coastal System and the Humboldt System. The Ecuadorian
6 Echinoderms of Ecuador 185
Fig. 6.1 The Ecuadorian continental shelf and current system. Source Google (Digital Satellite
Imagery, TerraMetrics, 2011) modified by L. Molina 2011
marine and coastal area constitutes a remarkable transition zone in the eastern
Pacific, characterized by a significant spatial and temporal variability of the
physical-oceanographic conditions.
The ocean circulation in the eastern equatorial Pacific area is dominated by the
action of the winds and the circulation patterns of several currents, mainly
the North and South Equatorial Currents, the Equatorial Counter Current, the
California Current, the Panamá Current (PC), the Perú or Humboldt Current, and
the Equatorial Undercurrent or Cromwell Current (Allauca 1990). All of these
currents carry water that contributes to the formation of the Equatorial Front, which
moves from north to south depending on the displacement of the Inter-tropical
Convergence Zone (ITCZ, Wyrtki 1965). Stevenson (1970) detected the presence
of a circulation system located between the mainland and the Galápagos Islands
(Fig. 6.1), which varies in size between the wet and dry seasons. A decrease in the
gradient of the prevailing trade winds and a reduction in the intensity of the Perú
Current, shifts the balance of forces along the frontal boundary. This results in a
southerly movement of a narrow stream of nutrient poor, warm water from the
basin of Panamá (Panamá Current) down to the coast of Ecuador and Perú between
December and April (Fig. 6.1). On the other hand, during the period of strong NE
trade winds, the Panamá Current can extend to 3°S (Enfield 1975).
The Peruvian Currents
Both the oceanic and coastal currents are initiated in at 40–458S and transport
highly saline, cold water (14–23 °C) to the north. The flow continues in a
northwesterly direction along the coast of Perú to the Ecuadorian coast where it
joins the South Equatorial Current system (Fig. 6.1). This current system prevails
from May to November and it represents a ‘‘change of season’’ in Ecuador, as it
significantly decreases the sea surface and air temperatures. This system provides
favorable conditions for fisheries, because its waters are rich in nutrients. The
greatest intensity of this current is between July and September. It gradually
weakens and disappears completely in December, when the Panamá Current,
warm and low in nutrients, starts to dominate Ecuadorian waters (Fig. 6.1).
The Equatorial Undercurrent
This current runs eastwards along the equatorial Pacific right in front of Ecuador,
reaching speeds of approximately 1.5 m s-1. When it reaches the Galápagos
Islands it weakens and splits into two branches, one north and one south of the
archipelago. The southern branch moves to southeast and generates outcrops that
occur along the northern coast of Perú (Fig. 6.1).
The Ecuadorian Coastal Current (ECC)
The Ecuadorian Coastal Current (ECC) moves northwards, paralleling the coast of
Ecuador. The northern boundary of this current is linked to the position of the
ITCZ, and begins to descend toward the south during December (Moreano 1983).
An important aspect to consider is the influence of the Gulf of Guayaquil in the
area around its mouth, because it has a well defined circulation regime. Yet it
somehow alters the normal course of the circulation of the ECC, mainly through
the influence of tidal currents that are stronger into the Gulf (Allauca 1990;
Fig. 6.1). Two semidiurnal tides occur both along mainland Ecuador and at the
Galápagos. The ECC influences the local currents along the continental coast of
Ecuador, and the marine ecosystems and coastal geomorphology.
6.1.2 The Galápagos Islands
Geography and Climate
The Galápagos Islands are situated astride the equator, between 01°400 N and
01°250 S, and between 89°150 W and 92°000 W, about 1,000 km from the Ecuadorian
mainland. These oceanic islands have been in existence for at least 3.3 million
years. They are made up of 13 major islands, the smallest having an area of
14.1 km2 (Genovesa Island), and the largest of 4,588 km2 (130 9 40 km, Isabela
Island). There are also over 115 islets and rocks, most of which have vegetation.
The most recent islands, Fernandina and Isabela, are moving slowly from west to
east, while the oldest islands are San Cristóbal and Española (Cox 1983; Geist
1996). The total land area of the archipelago is about 8,000 km2. It is surrounded
by a Marine Reserve that covers an area of 137,975 km2, with a total of 1,800 km
of coastline (Snell et al. 1995; Toropova et al. 2010) making it the ninth largest
marine protected area in the world. Outside the Marine Reserve, national waters
extend 200 nautical milles from the island baseline. Climatologically, these islands
are not a uniform group, as they extend both north and south of the equator and are
influenced by different oceanographic conditions (Abbott 1966). More than 90 %
of their surface area is located south of the equator. In spite of their equatorial
position, a rather temperate climate (with an annual mean temperature of *24 °C)
prevails because the islands are located in a climatic transition zone between the
west coast of South America and the dry zone of the Central Pacific Ocean. These
islands are influenced by both warm and cold currents (Figs. 6.1, 6.2) which meet
in the Galápagos region. Most climate data available come from the arid coastal
zone where the year is divided into a dry season (June–October) and a rainy season
(December–April). During the dry season, upwelling cold oceanic water (19 °C)
flows along the southern part of the archipelago. The extreme temperatures in the
‘‘arid zone’’ are rarely below 13 °C or above 23 °C, which are cool for this latitude.
There is occasional or intermittent drizzle in the coastal zone but almost continuous
drizzle in the higher zones of the islands. This kind of rainfall, locally called
‘‘garua’’, is also typical of all South American coasts influenced by the Humboldt
Current (Chile, Perú, and southern Ecuador). In the arid zone, the monthly rainfall
from May to October never exceeds 25 mm. During the rainy season, the dominant
wind changes from south-southeast to east, with light winds in the lower regions.
Upwelling of cold oceanic water stops during this period and genuine tropical
conditions prevail. The heaviest rainfall is on the southern slopes of the large
islands above 250 m altitude. There is almost no rainfall on the leeward side of the
coastal region (northern side).
Oceanography
Several marine currents converge at the Galápagos Islands (Fig. 6.2). This
determines the great ecological uniqueness of these marine ecosystems. Seven
ocean currents have been identified. Three of these are the most decisive currents
that determine the seasonal periods of the islands, in terms of intensity, direction
and temperatures: (1) the South Equatorial Current (SEC); (2) the warm Panamá
Current; and (3) the deep cold Cromwell Current (Chávez and Brusca 1991;
Banks 1999).
South Equatorial Current (SEC)
The SEC is the main surface current that flows westward. It is a product of the
confluence of two currents with different direction and intensity throughout the
Fig. 6.2 The current system and biogeographic zones of the Galápagos Islands according to
Edgar et al. (2002b). Source Galápagos National Park Service and Charles Darwin Foundation
2003
year (Fig. 6.2), the tropical Panamá Current (*30–34 %, depending on rainfall),
and the sub-tropical Perú Current (*35 % close to Ecuador). The dominant flow
occurs during the rainy season (from May to November) resulting from the winds
blowing from the southeast (Muromtsev 1963). Advection of the cold surface
waters from the west (product of the equatorial upwelling, induced by wind at low
latitudes) that maintains cold local temperatures in Galápagos (Wyrtki 1966,
1974). The differences in temperature and salinity between the two currents form
an oceanic front at the crossroads between the two bodies of waters producing
gradients of up to 5 °C at 50 km (Banks 1999). The front descends from north of
the Galápagos during the southern summer hemisphere (December to May)
reducing the temperature gradient from northwest to southeast, while the sea
surface temperature is homogenized in the archipelago (Banks 2002). Both surface
currents are relatively impoverished in nutrients after extensive circulation in the
open ocean gyres (Thurman 1996). The Perú Current is an extension of cold waters
originating in equatorial upwelling zones between Galápagos and the coast of
mainland Ecuador and Perú.
The Panamá Current (PC)
Every year a body of surface water (warm and low in salinity and nutrients) that
flows from Panamá (Fig. 6.2) reaches Galápagos in January. At this time of year,
the SE winds decrease in intensity and the currents flowing westward also weaken.
This allows the ITCZ warm water masses to invade from the north. This mass of
warm water flows southward along the NW coast of South America. When the PC
penetrates to the south and lasts long enough, the event known as El Niño occurs.
This has devastating impacts on the marine ecosystems of the region (Jiménez
1987).
The Subsurface Equatorial Current (or Cromwell Current)
The Cromwell Current directly or indirectly generates a series of unusual effects.

These include cool surface water temperatures on the equator (17 °C on average)
and a large amount of plankton that is responsible for the richness of fauna in the
archipelago. It has a thickness of 300 m and a width of 400 km. It is normally
positioned at a depth of about 100 m, well below the euphotic zone. Upwelling
occurs when it hits the Galápagos platform (Fig. 6.2) bringing nutrient rich, cold
waters to the surface (Wyrtki 1985). This current produces persistent upwelling
zones enriched with nutrients in the western coasts of most islands and consistently
colder water masses. This allows the occurrence and persistence of temperate
species in Galápagos, which is an equatorial zone (Feldman 1985). The branches
of this water mass come together again in one undercurrent about 100 km east of
the archipelago (Banks 1999). Based on analysis of satellite remote sensing data
(SeaWiFS), Banks (2002) identified regions where annual primary production is
consistently high, in particular to the west of Isabela, known for its high pro-
ductivity and fishing intensity, but also at some easterly points of Isabela and SE
Santa Cruz, which may be due to local patterns of currents and seamounts. It has
been suggested that an elevated productivity (10–30 mg Chl a m-3) occurs in
sporadic but intense events of short duration, while occasional, spatially large but
moderately intense (0.5–1.0 mg Chl a m-3) productivity occurs throughout the
archipelago. This is possibly due to a mixing of points of freshwater runoff and
nutrient enrichment from the coast, surface advection from other islands, and
retention effects of nutrients from areas with high sediment content and mangrove
bays (Wellington 1984; James 1991).
Explorations of echinoderms in Ecuador have been intensive, particularly around

the Galápagos Islands, where several expeditions made collections over the past
100 years. These include the Albatross in 1894, 1898, 1899, 1905; the Eagle and
Table 6.1 Summary of the total number of echinoderms species

Continental Galápagos C Depth distribution (m)
shelf Islands
0– 20– 100– [1000 ND Total
Habitat 20 100 1000
Shallow Open
Crinoidea ND ND 3 0 ND ND ND 2 1 3
Asteroidea 19 5 67 15 11 5 12 16 27 71
Ophiuroidea 21 8 82 11 3 3 18 28 42 94
Echinoidea 24 2 43 16 7 4 16 10 14 51
Holothuroidea 19 13 47 11 11 2 2 24 24 63
Total 83 28 242 53 32 14 48 80 108 282
Information is referred for habitats and depth distribution along the continental shelf and the
Galápagos Islands, Ecuador. C common number of species for both the continental and the
Galápagos Islands. ND no data
Ara in 1921–1928; the Presidential Cruise in 1938; the Velero III and IV between
1932–1954; and the New York Zoological Society in the late 1930s. More
recently, an expedition was undertaken using the research submersible Johnson-
Sea-Link II in 1998 (Pawson and Ahearn 2001). These expeditions produced the
majority of articles dealing with taxonomy, distribution and diversity of echino-
derms from subtropical and tropical waters of the Western Americas. Publications
relevant to Ecuador, include Agassiz (1892, 1898, 1904)), Ludwig (1894, 1905),
Lütken and Mortensen (1899), Clark (1902, 1917, 1920a, 1920b, 1939, 1940,
1948), Fisher (1911), Boone (1933), Ziesenhenne (1935), Deichmann (1938, 1941,
1958), Caso (1948, 1958, 1961, 1962, 1965, 1976, 1978, 1979), Barham et al.
(1973), Brusca (1980), and Maluf (1988a, b).
This chapter includes information about the echinoderms of the continental
coast and the Galápagos Islands, based on all known records of the literature and
on published data from the catalogues of the Allan Hancock Foundation Collec-
tions, Los Angeles County Museum and from holdings at the Scripps Institute of
Oceanography and at the National Museum of Natural History of the Smithsonian
Institute. Furthermore, species observed in the field, but not necessarily reported in
the literature or present in collections (Chiriboga et al. 2011) are also reported
here. All species are listed in the Appendix.
6.2.1 Diversity
Ecuador possesses 282 species of echinoderms (Table 6.1). Eighty-three and 242
species have been on the continental shelf of Ecuador, and the Galápagos Islands,
respectively (Table 6.1).
6.2.2 Ecuadorian Continental Shelf
In general, the location with the greatest diversity of marine invertebrate species
(including echinoderms) seems to be in the Gulf of Guayaquil (which is the main
estuary of Ecuador). Maluf (1988a, b) identified this Gulf as a transition area
because there are a large number of southern echinoderm species whose ranges are
between 1° and 4°S. Transition zones may have a greater number of species
because co-occurrence in the ecotone between two similar habitat types is possi-
ble. However, this zone may have a higher biodiversity because it has been better
studied compared to the other areas along the continental coast (Cruz et al. 2003).
Even so, information is still insufficient for echinoderms (Villamar and Cruz 1983;
Avilés 1984; Mair et al. 2002; Sonnenholzner and Lawrence 2002). So far, sci-
entific studies have been limited to basic research and the taxonomy of the biota.
To date, there is no proper analysis of the distribution of the fauna of echinoderms
based on the limited available records (Appendix).
The echinoderm fauna presently known is composed of 26 species: nine species
of Asteroidea, two species of Ophiuroidea, ten species of Echinoidea, and four
species of Holothuroidea. Of these, 17 species have been reported from the
intertidal area and shallow water environments, less than 20 m depth (Mair et al.
2002). Sea urchins, such as Echinometra vanbrunti and Eucidaris thouarsii; sea
cucumbers such as Holothuria theeli, and Holothuria arenicola; and sea stars such
as Pharia pyramidata and Phataria unifascialis are extremely common along the
central and southern coast of mainland Ecuador.
6.2.3 The Galápagos Islands
The Galápagos Islands are the southern distributional limit for many species.
Although these islands are separated from the nearest American mainland by
1,000 km of open water, there are several factors influencing the dispersal process
that favor affinity between the Panamá Bight (Panamá/Colombia/Ecuador), and the
Galápagos, while a smaller affinity is expected between the Galápagos and Perú
(Glynn et al. 1983). Only one-third of Perú’s echinoderms are shared with the
Galápagos Islands (Maluf 1991). The pronounced offshore movement of waters in
northern Perú would favor selection for non-planktotrophic species along the
mainland coast with reduced larval dispersal to the Galápagos. The islands lie to the
south of the normal path of the North Equatorial Countercurrent, the best potential
dispersal route for shallow-water tropical forms between the Central and the
Eastern Pacific. The greatest influence of this current is between 4° and 10°N
latitude, along the coasts of Costa Rica, Panamá, Colombia and the northern part of
Ecuador. Trans-Pacific species may have secondarily dispersed to the Galápagos
Islands from established mainland populations. The Equatorial Undercurrent could
transport (at great depth) central Pacific tropical larvae to the Galápagos.
Two hundred and forty-two species of echinoderms that are known to occur in
Galápagos and the Galápagos Ridge, including three crinoids, 67 asteroids, 82
ophiuroids, 43 echinoids, and 47 holothurians. These islands have a high per-
centage of endemic species relative to other offshore islands of the Central Eastern
Pacific (between 34°300 N and 18°000 S), because of the high environmental and
spatial heterogeneity of the archipelago, due to its geographic location (Maluf
1988a, 1991). Seventeen percent of echinoderm species in Galápagos (a total of 34
species) are endemic, particularly the asteroids and ophiuroids.
Of these species, 27 species inhabit deep waters between 105 and 925 m (See
Appendix), and 107 inhabit depths between 20 and 100 m (30 species of asteroids,
46 species of ophiuroids, and 31 species of echinoids). Deep sea urchins, such as
Plagiobrissus pacificus, Brissopsis pacifica and Idiobryssus coelus have been
recorded. For intertidal and shallow water environments (\20 m depth), 14 species
of asteroids, 28 species of ophiuroids, 25 species of echinoids, and 17 species of
holothuroids have been reported. Hickman (1998) and Chiriboga et al. (2011)
prepared a list with the most common species of echinoderms along the intertidal
and shallow waters around the Galápagos. Among the holothuroids are Holothuria
arenicola, H. impatiens, H. imitans and H. theeli, and Isostichopus fuscus. The
latter species is common in the cold western region of Galápagos (Fig. 6.3a).
Among the echinoids are Eucidaris galapagensis (Fig. 6.3b), Lytechinus semitu-
berculatus (Fig. 6.3c), Tripneustes depressus (Fig. 6.3d), Diadema mexicanum
(Fig. 6.3e), Toxopneustes roseus (Fig. 6.3f), Astropyga pulvinata (Fig. 6.3g),
Agassizia scrobiculata, and Clypeaster rotundus. Eucidaris galapagensis was
formerly identified as E. thouarsii, but new genetic evidence recognized it as a
species endemic to the Galápagos Islands, Cocos Islands and Clipperton Atoll
(Lessios et al. 1999). Among the asteroids are Pharia pyramidata (Fig. 6.3h),
Phataria unifascialis (Fig. 6.3i–j), Nidorellia armata (Fig. 6.3k), Pentaceraster
cumingi (Fig. 6.3l), and Mithrodia bradleyi (Fig. 6.4a) are the most common,
while Astropecten armatus (Fig. 6.4b), Leiaster teres (Fig. 6.4c), Heliaster cum-
ingi (Fig. 6.4d), and Heliaster solaris are rare. Among the ophiuroids are Ophi-
actis savignyi, Ophiocoma aethiops, O. alexandri, and Ophioderma teres.
The distribution and diversity of echinoderms in the Galápagos Islands differs
greatly among biogeographic regions (Maluf 1991). Edgar et al. (2004) proposed a
bio-regionalization scheme of five regions within the Galápagos, based on the
scheme proposed by Harris (1969) using a data set from communities and species
richness of reef fish and macro-invertebrates. These bioregions are the ‘Far-
northern’, ‘Northern’, ‘Southeastern’, ‘Western’ and ‘Elizabeth’ (Fig. 6.2). These
authors emphasized that both faunal abundance and species richness indicate that
Galápagos inshore reef ecosystems are divisible into three major regional com-
ponents: (1) the far-northern islands of Darwin and Wolf; (2) the central, southern
and eastern islands, including the east coast of Isabela; and (3) Fernandina and
western Isabela. Concerning the echinoderm fauna, the following patterns were
identified:
Fig. 6.3 Common echinoderms off the mainland and at the Galápagos Islands, Ecuador.
a Isostichopus fuscus (Ludwig, 1875), b Eucidaris galapagensis Döderlein, 1887, c Lytechinus
semituberculatus (Agassiz and Desor, 1846), d Tripneustes depressus A. Agassiz, 1863,
e Diadema mexicanum A. Agassiz, 1863, f Toxopneustes roseus (A. Agassiz, 1863), g Astropyga
pulvinata (Lamarck, 1816), h Pharia pyramidata (Gray, 1840), i–j Phataria unifascialis (Gray,
1840), k Nidorellia armata (Gray, 1840), l Pentaceraster cumingi (Gray, 1840). Photos by: J.
Sonnenholzner (a–b); Charles Darwin Foundation (c); and J.C. Navarro Gómez (d–l). Note that
photos are not to scale
1. The islands of the Far Northern and the Northern islands include a large
component of species from the Indo-Pacific and Panamá, with no endemic
species. Species associated with warmer waters and coral environments are
Fig. 6.4 Common echinoderms off the mainland, and at the Galápagos Islands, Ecuador.
a Mithrodia bradleyi Verril, 1870, b Astropecten armatus Gray, 1840, c Leiaster teres (Verill,
1871), d Heliaster cumingi (Gray, 1840), e Asteropsis carinifera (Lamarck, 1816), f Luidia
bellonae Lütken, 1865, g Linckia columbiae Gray, 1840, h Caenocentrotus gibbosus (L. Agassiz
and Desor, 1846), i Stichopus horrens Selenka, 1867, j Paulia horrida (Gray, 1840), k Acanthaster
planci (Linnaeus, 1758), l Coronaster marchenus Ziesenhenne, 1942. Photos by: J.C. Navarro
Gómez (a); X. Romero (b); A. Chiriboga (c–l). Note that photos are not to scale
Mithrodia bradleyi and Asteropsis carinifera (Hawaiian species, see Fig. 6.4e)
are found on the northern edge of the Galápagos (Darwin and Wolf Islands).
2. Western islands (west of Isabela and Fernandina) and the Elizabeth bioregion,
contain numerous endemic species and those whose distribution includes the
cool-temperate Peruvian waters (Heliaster solaris and Luidia superba). Their
distributions extend to the south of Ecuador. The colder water species (surface
temperature can drop to 14 °C in August and December) include Astrometis
sertulifera, Luidia bellonae (Fig. 6.4f), and Linckia columbiae (Fig. 6.4g).
Echinometra vanbrunti is a rare species in this region, despite being common in
almost all the islands. It is presumed that the Peruvian species occur here due to
the environmental conditions that include colder temperatures and more pro-
ductive waters, similar to the conditions prevailing along the coast of South
America where these species initially evolved. This is also the case of the sea
urchin Caenocentrotus gibbosus (Fig. 6.4h), an immigrant from the temperate
region of the Peruvian-Chilean banks, which is dominant in the lower areas of
the western coast of Isabela and Fernandina Islands.
3. The Central/Eastern/Southern bioregions contain a combination of species from
various regions. Widely distributed in shallow waters are the sea urchins Eu-
cidaris galapagensis (Fig. 6.3b), Tripneustes depressus (Fig. 6.3d), Diadema
mexicanum (Fig. 6.3e), Toxopneustes roseus (Fig. 6.3f) and Centrostephanus
coronatus. Lytechinus semituberculatus is native to the Galápagos, and replaces
the ubiquitous L. pictus of the Eastern Pacific. On sandy and rocky bottoms, the
most frequent sand dollars species are Clypeaster elongatus, C. europacificus
and C. rotundus.
Edgar et al. (2004) emphasized that the variation in the abundance of echino-
derms around individual islands was high, often greater than the variation between
islands, thereby significantly lowering the density of certain species of sea stars.
For example, Heliaster cumingi (Gray, 1840) was not found during baseline sur-
veys despite being previously abundant, occurring in densities greater than 2–
3 ind m-2 in the intertidal (Wellington 1975). Nevertheless, it has been recently
found in Pinzón Island (see Fig. 6.4d). On the other hand, densities of some other
species seem to be increasing, for example the sea urchin Tripneustes depressus. It
appears that its abundance has increased at least an order of magnitude since the
1970s. This species was recorded in just one of ten sites investigated by Strickland
(1968), Wellington (1975), and Glynn (Edgar et al. 2002b). However, it is also
known that its populations greatly fluctuate through time, absent during some
periods, but very abundant during others (Brandt and Guarderas 2002).
Lawrence and Sonnenholzner (2004) documented the occurrence of the most
conspicuous echinoderms at seven sites with distinct habitats in central Galápagos
(i.e. sites were considered upon spatial gradients, from north to south, on hydro-
dynamics, wave energy, and sloping). Eighteen species (eight species of echinoids,
five species of holothurians, and five species of asteroids) were recorded. Of the
echinoids, Eucidaris galapagensis occurred in most of the sites ([90 % of the
quadrats at every site except one), and reached the highest densities (between
10.1 ± 4.8 and 15.6 ± 6.1 ind m-2). Lytechinus semituberculatus was found in
[90 % of the quadrats at two sites. Other echinoderms present in more than 35 %
of the intervals along transects at a site), were the echinoids Tripneustes depressus,
Diadema mexicanum, Toxopnesutes roseus and Caenocentrotus gibbosus, the
asteroid Pentaceraster cumingi and the holothuroid Holothuria atra. Diversity was
greater at the northern sites than at the southern sites, but there were low densities
of asteroids and holothurians. This may be related to a lack of appropriate substrata
or food availability at the sites. This is in agreement with Roff et al. (2003) and
Edgar et al. (2004) who indicated that, in most cases, abiotic factors such as
salinity, temperature, depth, rock type or sediment particle size are important
factors for biota that act in synchrony with biological process to affect macro and
micro distribution.
Pawson and Ahearn (2001) collected echinoderms from bathyal and abyssal
waters and reported four new records for the Galápagos Islands and eight new
species. The new species included four holothurians (one representing a new
genus), one echinoid, and three asteroids. The cidaroid echinoid Centrocidaris
doederleini that was formerly considered rare occurs in numerous populations,
reaching average densities of 185 ind m-2 (Pawson and Ahearn 2001). The
ophiuroids Ophiothix spp. and Ophiacantha spp. often occur together in large
aggregations of more than 100 individuals in black sandy areas with negligible
current activity (Pawson and Ahearn 2001). The asteroids Henricia spp. and
Coronoaster sp are fairly common on a variety of habitats, but were found mostly
on rocky substrates; the stalked crinoids Thalassometra agassizii and Calamocrinus
diomedeae are very rare (Pawson and Ahearn 2001).
6.3 Research
6.3.1 Ecology and Population Biology
Sea Cucumbers
Isostichopus fuscus
Distribution and Density

Isostichopus fuscus is a deposit-feeding aspidochirote species distributed from the
Gulf of California to Ecuador, including the Galápagos Islands (Maluf 1991).
The cylindrical, tough and flexible body is brownish with darker warts (Fig. 6.3a).
The color may change to dark brown depending on the color of the substrate. This
species is found mainly on rocky reef areas from 2 to 40 m depth (Hickman 1998)
although it has been seen at 60 m. In Galápagos, its distribution is not continuous.
Therefore there is a high level of uncertainty regarding its population dynamics.
The largest population was concentrated in the western region of the archipelago
along the Bolívar Channel (Murillo et al. 2002a). The average size of the indi-
viduals was 20 cm in length and the range in wet weight was 100–410 g (average
271 g) (Sonnenholzner 1997). At this location densities fluctuated between 0.27
and 0.40 ind m-2 from 1994 to 1997, but decreased to 0.13 ind m-2 due to an
illegal fishery (Piu 1998, 2000; Martínez 1999; Murillo et al. 2002a, b). In 2003,
the lowest adult density was recorded in this area since 1999 (with a reduction of
62 %). Juveniles (\16 cm) were almost completely absent from the survey area
from December 1994 until July 1999, after which there was a sudden increase,
with juvenile densities increasing by a factor of 20, reaching a peak value of
0.47 ind m-2 in April 2001. This maximum density of juveniles implies settlement
sometime in early 1998, at the height of El Niño event (Hearn et al. 2005).
Following this peak, there was a steady decline, with densities as low as
0.20 ind m-2 in the pre-fishery survey of 2003 (Hearn et al. 2005). The Bolívar
Channel has been recognized as a sea cucumber nursery ground and has remained
closed to fishing activities for the fishing seasons of 2003, 2004 and 2005 (Toral-
Granda 2005). Lawrence and Sonnenholzner (2004) reported densities of I. fuscus
in seven localities at the central region of Galápagos of between 0.02 ± 0.03 and
0.05 ± 0.04 ind m-2.
Growth and Reproduction

There is no knowledge of the recruitment dynamics or growth patterns of the
population of I. fuscus in Galápagos. Herrero-Pérezrul et al. (1999) and Reyes-
Bonilla and Herrero-Pérezrul (2003) obtained preliminary population parameters
of I. fuscus from México, including estimations of asymptotic length
(L? = 32.613 cm) and growth coefficient (K = 0.2115 yr-1). However, Hearn
et al. (2005) questioned if those growth parameters were applicable to Galápagos,
where individuals might reach 16 cm in length approximately 3.3 yr after settle-
ment and the minimum landing size of 20 cm in length after 4.5–5 yr. Isostichopus
fuscus reproduces sexually and can spawn throughout the year. Complete infor-
mation on its reproductive biology is limited for the whole archipelago. Toral-
Granda (1996) studied the reproduction of I. fuscus in a population at Caamaño
Island, Galápagos in 1995. Fifty percent of individuals were sexually mature at
23.6 cm and between 161.0 and 170.0 g dry weight. The smallest individual (less
than 16 cm) found with mature gonads had a dry weight between 121.0 and
130.9 g. Four reproductive peaks were found, two reproductive events in each cold
and warm water seasons, suggesting a continuous reproduction. The highest gonad
index (GI) value was found in December whilst the lowest was recorded in April
and September. Mean fecundity (eggs per gonad) for this species was
567,884 ± 95,528. Many animals were regenerating on June 2002 at the western
side of Galápagos at the Bolívar Channel (Sonnenholzner, personal observation).
These specimens were between 8 and 10 cm in length, and were without a mouth
or an anus, indicating an asexual reproduction episode by fission in this population.
Diet and Feeding Habits

Sonnenholzner and Cajas (1993) determined the most dominant food components
of I. fuscus collected early in the morning at Bolívar Channel at a depth of 12 m.
The gut contents contained a high percentage (50.3 %) of polyps of stony corals,
followed by benthic diatoms of the genus Melosira sp. (12.3 %), Grammatophora
sp. (11 %) and Navicula sp. (9 %), and low percentages of other species of
microalgae. A variety of microorganisms, such as foraminifera and tintinnids
(0.7 %), as well as abundant detritus was found. Isostichopus fuscus is mostly
active at night (Hickman 1998; Hearn and Pinillos 2006).
Stichopus horrens

Stichopus horrens (Fig. 6.4i) is a large holothurian that can grow to a length of
50 cm (Schoppe 2000). The largest specimen from Galápagos was collected in the
western region with a length of 42 cm (Hearn and Pinillos 2006). This species is
found in shallow waters at depths between 5 and 20 m, mostly occurs on rocky
reef substrates in the central and southern region of Galápagos (Edgar et al.
2002b). Stichopus horrens tends to contract to less than half its length during its
cryptic behavior during the day. It emerges around dusk to form aggregations with
densities exceeding 1–2 ind m-2 (Hearn and Pinillos 2006).

Throughout the Galápagos Islands, the population structure of Stichopus horrens
has a uni-modal length frequency distribution ranging from 14 to 42 cm. The mean
length of individuals was around 28 cm at most islands, although smaller indi-
viduals occur at Rábida (22 cm) and Wolf (24 cm). No individuals smaller than
12 cm in length were recorded. Nothing is known about the larval biology and
recruitment of this species. Hearn and Pinillos (2006) made preliminary estimates
of the Von Bertalanffy population parameters using size frequency distributions
(L? = 38.8 cm, K = 0.17), which indicated that an individual of 20 cm in length
would correspond to an age of 4 years. Stichopus horrens reproduces asexually by
fission (Kohtsuka et al. 2005), as well as sexually. Hearn and Pinillos (2006) found
individuals greater than 16 cm in length were mature. Fecundity was estimated to
vary greatly between 1 9 103 and 1 9 106 eggs per individual.
Holothuria theeli

Sonnenholzner and Lawrence (2002) and Sonnenholzner (2003) evaluated the
distribution and density by microhabitat preference of Holothuria theeli along the
continental coast of Ecuador. The maximum density was at the lowest tidal level
and there was a progressive decrease in abundance with depth. It was observed on
exposed surfaces at low tide, suggesting a resistance to desiccation. The capacity
to withstand and exploit air-exposed areas during the tidal cycle has been observed
in other species such as Stichopus badionotus (Sloan and Von Bodungen1980),
Holothuria atra and Actinopyga echinites (Massin and Doumen 1986). Holothuria
theeli ranged in size from \5 to 20 cm in length. The relative absence of juveniles
(individuals less than 5 cm in length) and distinct cohorts upon the midshelf reef
suggest different habitat preferences between juveniles and adults (5–11 and
11–20 cm long, respectively). This probably also occurs with the population of
Isostichopus fuscus in the Galápagos Islands (Martínez et al. 1998), and with
H. atra and S. chloronotus from the Great Barrier Reef (Uthicke 2001). Small and
medium juveniles inhabit areas with boulders, cobbles, pebbles and sand, whereas
large adults mainly inhabit areas with boulders and cobbles. In order of preference
the most used microhabitats were boulders/cobbles [ pebbles/sand [ thick
pavement/bedrock (Sonnenholzner and Lawrence 2002).

Sonnenholzner (2003) determined abundances of H. theeli on transects on a
shallow tidal reef flat at the Puntilla Santa Elena (Fig. 6.1) in 1994. Abundances
were variable in May but increased by the end of August as a result of asexual
reproduction by fission (during the sampling period many individuals were
observed without a mouth and none without an anus). The fluctuation in abun-
dance following the period of fission could be related to the survival rates for
posterior and anterior body parts (Reichenbach and Holloway 1995). The impor-
tance of fission in reproduction is a well-known feature in the biology of this
species (Bonham and Held 1963; Uthicke 1997). Many individuals were observed
regenerating after asexual reproduction in May. The fission rate of individuals in
the intertidal zone to near shore reef was much higher (15–36 %) than at the high-
energy midshelf reef zone (3–7 %) (Sonnenholzner 2003).

Sonnenholzner (2003) determined that the distribution pattern of H. theeli on a
shallow tidal reef flat was apparently not limited by food availability (Yamanouchi
1956). Roberts (1979) and Pomory et al. (1998) suggested differences in feeding
time as a means of niche partitioning, where the habitat was partitioned into
microhabitats to avoid competition with other holothurians species found in the
area. Holothuria theeli generally begins burrowing around sunrise and emerges
close to sunset, showing a semi-cryptic behavior at night. Sonnenholzner (2003)
determined that the feeding activity of H. theeli may be low in May compared to
March and August, since the intestine length was shorter in May, the capacity of
the intestine may fluctuate as a physiological response to food availability (Tanaka
1958). For example, Stichopus japonicus ceases feeding after spawning and its
intestines degenerate and become smaller in size (Tanaka 1958). The dormant
condition in H. theeli may result from the change in water temperature from high
to low, which may stimulate physiological processes such as feeding inactivity
after spawning (Yamanouchi 1942). Under these conditions, H. theeli maintains
normal feeding activity in March and August but not in May. The dormant con-
dition is evident in juvenile H. scabra, where individuals of 4–14 cm in length
responded to changes in temperature. Specifically, the maintenance of a constant
warm temperature prevents H. scabra from burrowing (Mercier et al. 1999).
Holothuria theeli is undoubtedly a non-selective, generalist feeder throughout the
rainy and dry seasons, and appears to be a forager. It feeds at an almost constant
daily rate and probably continues to feed through stormy periods. The lowest
percentage of prey organisms (such as micro invertebrates and microphytoben-
thos) occurred in March and the highest in August. The major food sources of
H. theeli were crustaceans, foraminiferans, diatoms, and detritus. High levels of
meiofauna and microphytobenthos in the gut contents of H. theeli indicate that
both play a significant part in the nutrition of the holothurians without any rela-
tionship to the size frequency distribution of the sediment around holothurians
(Sonnenholzner 2003).
Sea Urchins
Eucidaris galapagensis

Although there were different expeditions since the 1800s to qualitatively study the
Galápagos marine diversity, including sea urchins, it was not until the 1950s that
information on abundances of sea urchins became available. In 1954, the slate-
pencil urchin Eucidaris galapagensis Döderlein, 1887 (Fig. 6.3b) occurred at a
mean density of 19 ind m-2 in the western islands (Malmquist 1991). During the
1970s, some studies indicated that its densities around the archipelago fluctuated
between 2–8 and 34–50 ind m-2 (Wellington 1975; Glynn et al. 1979; Glynn
1990).
Some years later, Glynn (1988) compared the densities of E. galapagensis
before and after El Niño 1982–1983 and found that they had increased from 5 to
30 ind m-2. He also found that E. galapagensis fed on hermatypic coral polyps
and dead corals, which were highly impacted by the warming event. This
explained the dramatic increase of the urchin abundances after El Niño (Glynn
et al. 1979; Glynn 1988). Similarly, Edgar et al. (2010) examined historical data to
document the spatial and temporal scale of recent changes in species distributions
and important benthic habitat types. Part of the study included the analysis of
underwater images (taken in 1982, 1983 and 1984) to assess changes in the
densities of E. galapagensis before and after El Niño 1982/1983. They found that
densities of this species significantly increased by a factor of about two from 1982
(0.45 ind m-2) to 1984 (1.0 ind m-2).
Brandt and Guarderas (2002) reported E. galapagensis was the most common
and abundant sea urchin species in the whole archipelago, with an average density
of 3.2 ind m-2. They also pointed out that it was abundant in the islands of Rábida
and Santiago with average densities of 5.0 ind m-2. The lowest densities occurred
in the northern islands of Darwin and Wolf with average densities of 2.1 ind m-2.
Irving and Witman (2009) carried out experiments and density surveys of E.
galapagensis in different habitats in Academy Bay, Santa Cruz Island. They found
maximum densities of 28 ind m-2 in algal turfs, and only 6 ind m-2 in barren
habitats. Sonnenholzner et al. (2009) studied the density of E. galapagensis in 20
shallow rocky reefs in the southeastern biogeographic region of the Galápagos
Marine Reserve (off Santiago, Santa Cruz, Baltra and Seymour Islands), specifi-
cally in sites open and closed to fishing. Eucidaris galapagensis densities (±SE)
were higher in open than in closed sites to fishing (4.5 ± 1.55 vs.
2.2 ± 1.55 ind m-2, respectively). More recently, Paredes (2010) conducted
population surveys of E. galapagensis at four sites in San Cristóbal Island between
August 2009 and March 2010. The highest densities were in Punta Carola and
Playa Mann (1.3 ind m-2) followed by La Lobería (1.1 ind m-2) and Tijeretas
(0.9 ind m-2).
Size Structure and Gonad Production

Brandt and Guarderas (2002) indicated that the mean test diameter of E. gala-
pagensis was 5.12 ± 0.01 cm around the archipelago. The test diameter ranged
from 1 to 9 cm and varied between regions/islands (H = 1462.162, p \ 0.0001,
df = 6), with the largest test diameters occurring in Floreana and the southeastern
islands.
During the Base Line survey collections between September 2000 and April
2001 Francisco and Smith (unpublished data) estimated morphometric relation-
ships and gonad index of the most common sea urchins. Twenty individuals of
each species (E. galapagensis, Lytechinus semituberculatus and Tripneustes de-
pressus) were collected haphazardly at a depth of approximately 6 m at 12 sites on
seven islands. Individuals were dissected and gonad index (GI) was calculated by
dividing the drained gonad weight by the total body weight, multiplied by 100.
The mean GI (±SE) for E. galapagensis was 2.98 ± 0.11 (Francisco and Smith,
unpublished data).

Eucidaris galapagensis feeds on encrusting algae (Wellington 1975; Hickman
1998), live hermatypic polyps, dead corals and barnacle plates (Glynn et al. 1979).
It contributes to the production of sediment in the subtidal community of the
Galápagos (Wellington 1975). It also has the ability to bore deeply into the sub-
strate to create crevices, which reduces its risk of dislodgement from hydrody-
namic forces (Witman and Dayton 2001).
Lytechinus semituberculatus

The green sea urchin Lytechinus semituberculatus (Fig. 6.3c) has been reported as
the second most abundant sea urchin species (1.2 ind m-2) of the Galápagos’
rocky reefs (Brandt and Guarderas 2002). L. semituberculatus was widely dis-
tributed in islands with colder and higher productivity water regimes, particularly
on Isabela Island (2.7 ind m-2). The lowest densities were in the northern islands
(0.1 ind m-2), and on Santa Cruz island (0.2 ind m-2). L. semituberculatus shows
a significant density pattern associated with depth, suggesting it prefers shallow
habitats (\10 m) (Brandt and Guarderas 2002). Edgar et al. (2010) documented
changes in its density before and after El Niño 1982/1983. They found a decrease
from 0.52 ind m-2 in 1982 to 0 individuals in 1984. In another study, L. semi-
tuberculatus had higher densities in barren grounds (51 ind m-2) than in turf
habitats (1.6 ind m-2) (Irving and Witman 2009). Finally, Paredes (2010) con-
ducted population surveys at four sites in San Cristóbal Island between August
2009 and March 2010. Density data showed that Bahía Tijeretas had higher
densities of L. semituberculatus than the other surveyed sites. The highest density
was 4.8 ind m-2, followed by La Lobería (1.6 ind m-2), Playa Mann
(0.9 ind m-2) and Punta Carola (0.6 ind m-2).
Size Structure, Allometric Scaling and Gonad Production

Brandt and Guarderas (2002) reported a mean test diameter (±SE) for L. semi-
tuberculatus in the whole archipelago of 4.3 ± 0.02 cm, with a range of 1–7 cm.
There were significant differences in test diameters among islands (H = 691.363,
p \ 0.0001, df = 6). Populations at Floreana and Santa Cruz islands had the
highest mean test diameters (Brandt and Guarderas 2002). In relation to gonad
production, Francisco and Smith (unpublished data) found a mean gonad index
(±SE) was 7.36 ± 0.43.
In general, body size affects most aspects of an organism’s biology (Calder
1984), and scaling of structure and function to body size can be considered in
terms of ontogeny and phylogeny. Lawrence et al. (2010) studied the ontogenetic
and phylogenetic scaling of lantern components (weight of demipyramid, rotula
and epiphysis of the lantern and test diameter) of L. semituberculatus from the
Galápagos Islands versus three other species of Lytechinus (L. variegatus from
west coast of Florida, U.S.A. and San Blas Islands, Panamá; L. williamsi from San
Blas Islands, Panamá; and L. pictus, northern Baja California, México) and one
species of Tripneustes (T. gratilla from Okinawa, Japan). Results showed that all
components were largest in the smallest species (L. williamsi), and smallest in the
largest species (T. gratilla), suggesting a genetic basis and interrelated scaling. All
components are larger in L. variegatus from Panamá than those from Florida,
which could have a genetic and/or environmental basis. These results are very
important, since the arguments proposed by Ebert (1980) suggested that large
demipyramids of a lantern of sea urchins increase their capacity to feed.

Nonetheless, Lawrence et al. (2010) determined that the Aristotle’s lantern of sea
urchins can show an allometric relation with body size and that it varies intra- and
inter-specifically. The reasons for this have not been well established, but these
results should be considered for well-focused ecological studies along the
Galápagos Islands, concerning growth and reproduction of this abundant species.
Because ingestion is the largest term in the balanced growth equation, the feeding
apparatus of animals is of particular interest in scaling (Peters 1983).

Hickman (1998) reported L. semituberculatus feeds on algae and debris in the
Galápagos Islands. It is commonly covered by pieces of seaweed or shells, which
would probably be a protection from ultraviolet radiation (UVR). In this sense,
Sharp and Gray (1962) provided some of the first evidence that artificial UVR can
induce covering in L. variegatus.
Tripneustes depressus

Idrovo and Sonnenholzner (1994) studied the distribution and abundance of
T. depressus (Fig. 6.3d) at Punta Los Frailes, Machalilla National Park (Fig. 6.1)
along the coast of mainland Ecuador. This species had an aggregated distribution
(Morisita index 2.9–1.6), with a density between 1.5 and 1.8 ind m-2.
Luna (2000) conducted density surveys of T. depressus at 35 sites around the
Galápagos Islands from 1996 to 1997. Density fluctuated from 0 to 1.5 ind m-2
with an average value of 0.3 ± 0.4 ind m-2. Higher densities were found in the
central region of Galápagos: Rocas Gordon (1.5 ind m-2), Caleta Bucanero
(1.2 ind m-2) and Cabo Marshall (0.8 ind m-2), whereas consistent lower den-
sities (mostly no individuals) were found in the western islands (Cabo Berkeley,
Bahía Urbina, Bahía Elizabeth, Punta Moreno, Cabo Rosa, Punta Espinoza and
Cabo Douglas), with the exception of Caleta Iguana which had densities of
0.2 ind m-2. Luna (2000) also conducted monthly surveys (from October 1996 to
April 1997) at one site, Playa Ratonera, Santa Cruz Island. The density of
T. depressus decreased from 0.5 to 0.1 ind m-2 over this period.
To understand spatial and temporal patterns of abundance of T. depressus,
Guarderas (2000) conducted monthly population surveys (from September 1998 to
September 1999) in intertidal and subtidal habitats in Santa Cruz Island. In this
study, this species had an aggregated distribution (the ratio variance to average
was higher than 1) with high temporal and spatial variation in abundance within
sites. Populations in subtidal sites had higher densities and were more stable and
more spatially homogenous than in intertidal sites, although there were no sta-
tistical differences among habitats. On average, El Barranco had significantly
higher average densities (3.0 ind m-2) than Barco Hundido site (0.6 ind m-2), the
Charles Darwin Research Station (0.5 ind m-2) and Punta Núñez (0.3 ind m-2).
Within all sites, density decreased over time. This result was explained by the
movement of sea urchins. It was concluded that T. depressus in intertidal sites
moved from shallow to deeper areas, probably to find more stable environmental
conditions. On the other hand, there was no clear direction or pattern of movement
in subtidal sites. Brandt and Guarderas (2002) described abundance of T. depressus
around the Galápagos Archipelago. The highest densities were found in Floreana
(0.9 ind m-2) and in the southeastern islands (Española and San Cristóbal with
0.6 ind m-2) while the lowest values were found on Fernandina (with
0.05 ind m-2). T. depressus showed significant density patterns associated with
depth, suggesting that it prefers shallow habitats less than 10 m depth (Brandt and
Guarderas 2002).
Size Structure, Growth and Gonad Production

Tripneustes depressus is the largest sea urchin species in the Galápagos Islands
(Hickman 1998). Size structure data for T. depressus is available for more sampling
periods than for any other sea urchin species in the archipelago. Population surveys
done in 1996 showed that the size frequency distribution had two peaks: (1) 7.0 cm,
and (2) 12.0 cm. In 1997, the size-frequency distribution had a single mode
(12.5 cm) and a mean size of 11.5 cm. Consequently the former surveys may have
presented a bimodal shape due to a smaller sample size (Luna 2000). In 1998,
population surveys conducted in the western islands of the archipelago, showed a
mass recruitment event (Brandt and Guarderas 2002). In 1999, intensive sampling
was conducted around three islands of the archipelago (Española, Floreana and San
Cristóbal). The results showed a bimodal size frequency distribution for each island
(Brandt and Guarderas 2002). On the other hand, the information for the 2000–2001
periods indicates that the mean test diameter around the archipelago was
10.0 ± 0.04 cm, with a range of 1–14 cm. The size frequency distribution had two
peaks (5.0 and 9.5 cm). Test diameter was significantly different (H = 499.9,
p \ 0.0001, df = 6) between regions/islands (Brandt and Guarderas 2002)
Guarderas (2000) determined growth parameters for T. depressus using modal
progression of size-frequency distributions of monthly data (September 1998 to
September 1999) from four sites in Santa Cruz Island (El Barranco, Barco Hun-
dido, Muelle de la Estación Charles Darwin and Punta Núñez). Data from all sites
were pooled to estimate population parameters. Analyses and processing of length
data were conducted using the Length Frequency Distribution Analysis (LFDA
Version 4.01), from the FISAT (FAO/ICLARM) package. Growth rate K (1 - L
(t)/L?) declines with time as the diameter of the animal reaches the asymptotic
length. Estimation of growth parameters (K, L?) of the Von Bertalanffy growth
curve was done followed by definition of grid boundaries using the methods of
Powell-Wetheral and ELEFAN (Electronic Length Frequency Analysis). The
asymptotic diameter (L?) ranged from 144.7 mm in the Punta Núñez site to
178.8 mm Barco Hundido. In the same way, the body growth rate coefficient (K)
varied from 0.50 in the Barco Hundido site to 0.90 in the Punta Núñez site. The
parameters estimated from all sites were L? = 182.3 mm and K = 1.38. The
growth function of Von-Bertalanffy used in this study has often been used to
calculate growth parameters of fish and many marine invertebrates, including sea
urchins. However, other models such as Richards, Jolicour, Tanaka and Gompertz
described better growth curves of sea urchins. All these models require informa-
tion obtained from capture–recapture methods of marked individuals. Therefore it
is necessary to carry out direct studies of growth for T. depressus and to compare
with the growth parameters obtained in this study (Guarderas 2000). Growth rates
for T. depressus were also estimated in laboratory by monthly measurements of
test diameters of different cohorts. Growth rates ranged from 4 to 10 mm per
month with an average of 5 mm per month (Guarderas 2000).
In relation to gonad production, Luna (2000) estimated gonad index from
bimonthly collections of 20 urchins from Caamaño islet (Santa Cruz Island)
between January 1997 and January 1999. This study showed that there was no
clear evidence for a distinct reproductive annual cycle for T. depressus. It seems
that there is spawning activity throughout the year. However, two peaks were
observed during the sampling period; one in November 1997 and the other in June
1998 (Luna 2000). On the other hand, Francisco and Smith (unpublished data)
determined that the mean gonad index for T. depressus was (±SE) 6.09 ± 0.36.
This information was obtained of twenty individuals collected haphazardly at
approximately 6 m depth at 12 sites on seven islands. Field sampling was con-
ducted between September 2000 and April 2001.

Guarderas (2000) studied the diet composition of T. depressus in Caamaño islet,
Santa Cruz Island from January 1998 to March 1999. Stomach contents were
sampled monthly. The diet consisted mainly of algae (98 %), although remains of
sponges, and unidentified invertebrates (2 %) were also found. For the algae, red
filamentous algae were the most abundant (49 %) and most frequent functional
group (99 %) in its diet (Guarderas 2000). There was temporal variation of dif-
ferent algal functional groups in the diet, which suggests that food availability
could determine the food of T. depressus. In addition, laboratory observations
showed that T. depressus in high density and starvation conditions (with low food
availability) are cannibalistic. These findings suggest that T. depressus tends to be
more a generalist than a specialist (Guarderas 2000).
Encope micropora
Size Structures and Allometric Scaling

Differences between paired structures sometimes occur normally, but random
variation can give insight into developmental stability and environmental stress.
In clypeasteroid sand dollars, the modified ambulacra, or petals, are among the
most conspicuous bilaterally paired structures. Lawrence et al. (1998) studied the
bilateral symmetry of the petals in the Ecuadorian sand dollar Encope micropora
L. Agassiz, 1841 versus others two species of sand dollars (considering large and
small specimens): (1) Mellita tenuis collected at Mullet Key, Florida, USA
(Scutellina, same sub-order of E. micropora); and (2) Arachnoides placenta col-
lected at South Mission Beach, Queensland, Australia (Clypeasterina, different
sub-order of E. micropora). Analysis of the anterior and posterior petals of
E. micropora suggested they were asymmetrical. This fluctuating degree of
asymmetry for some petal pairs might be a means to evaluate environmental stress.
Knowing the type of asymmetry is necessary to understand its origin, because
certain environmental conditions and forces can affect test form in sea urchins
(Ellers and Telford 1992; Ellers 1993).
Other Sea Urchins

Pinela and Calvache (1997) studied the distribution (Morisita index), and density
of Echinometra vanbrunti in four different localities along the southwest coast of
Ecuador. They found that this sea urchin had an aggregated distribution at all sites,
and that its densities ranged from 0.87 to 8.80 ind m-2. The abundance of this
species along mainland Ecuador is remarkably higher than in the Galápagos.
Brandt and Guarderas (2002) reported that E. vanbrunti presented densities of
0.3 ind m-2 in the Galápagos Archipelago. In addition, Wellington (1975)
reported that E. vanbrunti was abundant in the southern, central (including the
eastern side of Isabela Island) and northern regions (with 4.38 ind m-2), but rare
in Fernandina and on the western side of Isabela Island where Caenocentrotus
gibbosus (Fig. 6.4h) was the dominant species. Three other sea urchins (D. mex-
icanum, see Fig. 6.3e; Toxopneustes roseus, see Fig. 6.3f; and Centrostephanus
coronatus) were also reported by Brandt and Guarderas (2002). However their
abundances were significantly much lower (\0.01 ind m-2) than the other sea
urchin species found there. General findings of the Galápagos Marine Biodiversity
Baseline, including density patterns of sea urchins, were also published by Edgar
et al. (2004). Lawrence and Sonnenholzner (2004) documented the occurrence of
eight species of echinoids at seven sites with distinct habitats in central Galápagos.
The most diverse assemblage was at Itabaca Channel with E. galapagensis,
L. semituberculatus, T. depressus, T. roseus, C. coronatus, E. vanbrunti, C. gib-
bosus, and D. mexicanum. All had densities \2.3 ind m-2. The assemblages of
species are similar to those reported for the Florida Keys and the Hawaiian Islands
(Kier and Grant 1965; Ebert 1971).
Sea Stars
Luidia (Platasterias) latiradiata
Distribution
The living genus Platasterias is a surviving somasteroid and Luidia is a surviving
basal asteroid (Fell 1963). Luidia (Platasterias) latiradiata, based upon its skeletal
morphology, is considered a ‘‘living fossil’’ (Fell 1963; Caso 1970; Blake et al.
2000). The biogeographical distribution of this species has been extended to
Ecuador. It had been found close to the coast of México and Central America
(Caso 1970, 1976; Solís-Marín 1998). Villamar and Cruz (1983) found large
specimens on a sandy bottom in the intertidal zone at Playas, General Villamil, in
the mainland of Ecuador (Fig. 6.1). In México, this species have been reported by
incidental mortality due to shrimp fisheries, where this assumes an impact that can
cause a decrease in their populations. For more information, review the México
chapter.

Villamar and Cruz (1983) reported L. latiradiata fed on crustaceans (i.e. cuma-
ceans, miscidaceans), gastropods (Olivella semistriata), and bivalves (Donax
mancorensis).
Phataria unifascialis and Pharia pyramidata

Distribution of sea star species in shallow waters along the mainland of Ecuador is
essentially unknown. Sonnenholzner and Lawrence (2002) conducted a brief
survey focused on two sea stars, P. pyramidata (Fig. 6.3h), and P. unifascialis
(Fig. 6.4 g–h) at Los Frailes, Machalilla National Park, Manabí. They made sixty-
one transects (2 m wide 9 25 m long ‘‘double’’ quadrants, a total area of
3,100 m2). The density of P. unifascialis and P. pyramidata was 0.017 ind m-2
and 0.0006 ind m-2, respectively. These species were found mainly on open rocky
flat bottoms among the rocks, on or under boulders and dead coral.
Arm Size
Sonnenholzner and Lawrence (2002) measured P. unifascialis and P. pyramidata

in the southern and northern embayment at Los Frailes, Manabí. The arm radius of
P. unifascialis ranged from 40 to 80 mm, with most being 60–70 mm. The arm
radius of a small distinct cohort was 90–95 mm. The arm length of P. pyramidata
was between 93 and 121 mm.
6.3.2 Immune Response
Sea Urchins
It is well known that high temperatures have caused a widespread mortality of key
species, coral bleaching in many regions of the world, a decrease in number,
density and biomass of species with high immigration of predators and a decrease
in number of grazers (Epstein et al. 1998). Considering the ecological role of sea
urchins as key species in shallow coastal and marine rocky environments, and the
paucity of knowledge of their immune response during strong natural disturbances,
Sonnenholzner (2001) documented in detail the immune response of the sea urchin
E. vanbrunti during El Niño episode of 1997–1998 and the cold phase La Niña of
1999–2000. He made observations under laboratory (in vitro) and natural condi-
tions (in vivo, where specimens were collected along a gradient of intertidal rocky
pools at Anconcito, Santa Elena, see Fig. 6.1). Four coelomocyte types were
identified in the perivisceral coelom of E. vanbrunti: (1) the phagocytic
amoebocytes, which were the only phagocytic cells studied in this experiments
(Fig. 6.5a, b); (2) spherule cells, divided into colorless (with intense amoeboid
movements, and colorless cytoplasmic granules, see Fig. 6.5c), and red cells
(similar to colorless spherules, but with red granules, see Fig. 6.5c); and vibratile
cells (flagellated and small rounded cells, see Fig. 6.5d). The effect of three dif-
ferent temperatures: 19 ± 1 °C (low); 24 ± 1 °C (medium); and 29 ± 1 °C
(high) on the immune response of E. vanbrunti was evaluated using the perivis-
ceral coelom fluid and four immune assays: (1) antibacterial activity from the
plasma (clearance efficiency); (2) total hemocytes count (THC); (3) differential
hemocytes count (DHC); and (4) microbicidal activity (oxidative activity by
nitroblue tetrazolium reduction related to phagocytosis generated by stimulated
phagocytes in phorbol 12-myristate 13-acetate) (Sonnenholzner 2001). Both he-
mograms (THC and DHC) were useful assays related to the microbicidal activity.
At the high temperature in the laboratory conditions and during the mid phase of
the El Niño 1997–1998 warming event, four remarkable aspects of the phagocytes
were observed in natural conditions: (1) phagocytes were very motile cells and
spread rapidly, accompanied by a complete change in morphology (Fig. 6.5a, b);
(2) the aggregation-clotting process was accelerated (1.0 ± 0.5 min) at high
temperature conditions ([24 °C); (3) the oxidative activity increased (‘‘respiratory
burst’’); and (4) production of spheres-cellular masses with formation of microvilli
(Fig. 6.5b). A significant difference in clearance efficiency was observed with
different temperatures. This study showed that hemograms (DHC and THC),
clearance efficiency, and phagocytosis are reduced in E. vanbrunti at high tem-
perature conditions. These data are important because they provide baseline values
for tropical echinoid species in their natural environments that are affected by
strong environmental stress episodes. Thus, it is an excellent opportunity for study
and verifying through these immune indicators the health condition of this
potential bioindicator (as E. vanbrunti). Nonetheless, these findings indicate that
Fig. 6.5 Photomicrography of four types of coelomocytes identified in the perivisceral coelom of
the sea urchin Echinometra vanbrunti: a phagocytic amoebocytes in two different morphological
phases: a bladder or petaloid phase (into the square), and a filiform phase (black arrow);
b aggregate of filiform phacogytic amoebocytes; c red and colorless granulocytes; and
d flagellate cell, arrow indicate long flagellum. Photos taken by J. Sonnenholzner
more experimental studies are necessary to reach a definitive conclusion about the
possible effect of temperature in the immune response in the tropical sea urchin E.
vanbrunti.
6.3.3 Larval Ecology
In general, the larvae of echinoderms are an integral part of marine zooplankton

collections in the Ecuadorian waters, but their abundance and distribution have
been reported as rare (Luzuriaga 2011). Luzuriaga (2011) determined areas of
distribution and abundance of larvae of echinoderms (mainly ophiopluteus) based
on the analysis of time series data of zooplankton from fixed coastal stations (from
1996 to 2009) and oceanographic cruises (from 2004 to 2009). In some regions and
years an increase of the number of larvae of echinoderms along the continental
coast of Ecuador was observed, in areas possibly linked to large populations of

brittle stars, and sea urchins, and sand dollars (larva echinopluteus larvae)
(Luzuriaga 2011). Luzuriaga (1993) and Luzuriaga et al. (2007) reported high
densities of pluteus larvae ([200 pluteus larvae m-3) along the inner estuaries and
inland ecosystems of ‘‘Mataje-Cayapas’’ at Esmeraldas, at the mouth the rivers
Tambillo and Najurungo (Fig. 6.1), in August and December 1979 and November
2000 when there is a large influence of oceanic waters into estuaries. However,
few ophiopluteus larvae were reported in the open sea in October 2003, at the
south and north of the Ecuadorian coast, near the archipelago of Jambelí, El Oro,
and Punta Galera, Esmeraldas (Luzuriaga 2011). The higher concentrations of
ophiopluteus larva were documented in surface and vertical collections (using nets
with a mesh size of 200 and 300 lm) in October 2004, November 2005 and
February 2009 cruises near the southern coast of Ecuador, from Puntilla de Santa
Elena to the estuary of Gulf of Guayaquil and Jambelí Archipelago. This area is
greatly influenced by the Humboldt Current off the coast of Ecuador (Fig. 6.6 a–j).
Therefore the waters are rich in nutrients and plankton biomass that favor larval
echinoderm development (Coello and Prado 1999; Luzuriaga and Elías 1999).
6.3.4 Predation, Parasitism and Disease
Sea Cucumbers
Sonnenholzner and Molina (2005) provided information on the host–parasite

relationship that occurs along the southeastern and western bioregions of the
Galápagos Islands by examining two families of aspidochirote holothurians: (1)
Holothuridae: Holothuria portovallartensis (n = 1557) and H. theeli (n = 983),
and (2) Stichopodidae: Isostichopus fuscus (n = 850) and Stichopus horrens
(n = 150). Holothuria portovallartensis and H. theeli were infected by the same
gastropod parasite Balcis panamensis. The prevalence of infection was high, with
95 and 79 %, respectively. The maximum number of parasites (adult and juvenile)
in the same host was eight and six, respectively. Both I. fuscus (n = 850) and S.
horrens (n = 150) had a low frequency of parasitism by Melanella falcata.
Sand Dollars
Echinoderms are subject to predation by crabs, gastropods, lobsters, fish, and other
species, and their populations are affected by biotic diseases caused by a variety of
parasites and microorganisms (Jangoux 1990). In particular clypeasteroids are
preyed upon by gastropods that drill holes into the coelomic cavity, although the
attack is not always successful (Moore 1956; Gerace and Lyndsay 1992;
McClintock and Marion 1993). Disease symptoms of populations undergoing mass
mortalities include lesions and necrotic tissue of the body wall caused by vibrios
Fig. 6.6 Location of stations 1996–2011 a Abundance of larva ophiopluteus larvae 100 m-3
during research cruises, in surface waters: b October 2004, c November 2005, d November 2006,
and e February 2009; taken from Luzuriaga (2011)
Fig. 6.7 Encope micropora. a Aboral view showing irregular portions of the test. Note also the
indication of disease (dark spots), magnification 1.2X; b Oral view showing diseased portion of
the test at the posterior lunule, magnification 2.1X; c Aboral view showing bore hole in
ambulacrum 1, magnification 1.2X; and d Record of bore holes from 500 specimens, where
interambulacrum regions (I–V) were most affected. Photos taken by J. Sonnenholzner, and figure
by M. Burgos
and aeromonads (Gilles and Pearse 1986). Sonnenholzner and Lawrence (1998)
studied the occurrence of predation and disease in a population of the sand dollar
Encope micropora on the coast of Ecuador (Playas de Villamil). Thirty-eight
percent of the specimens had pronounced irregularities on the test margin
(Fig. 6.7a). Irregularities were more common in large individuals (81 % of the
specimens between 52.0 and 98.5 mm in length). Seventy-four percent of the
irregularities were on the posterior half of the test. Small rounded holes on the
aboral test occurred in 48.4 % of the specimens, usually in ambulacra I, IV and V
and the interambulacra 3 and 4 (Fig. 6.7c). Irregularities in the test of this sand
dollar species may be caused for crabs Callinectes arcuatus and C. toxotes,
because they are potential predators of E. micropora, as Callinectes sapidus preys
on Mellita quinquiesperforata (Weihe and Gray 1968). Pagurid hermit crabs are
other potential predators. The bore holes found in the tests indicate gastropod
attacks (Fig. 6.7c, d). The gastropod Northia pristis and Polinices sp. are common
in the area.
Eighty-seven percent of specimens were diseased (Fig. 6.7b). Symptoms were

most frequent on the oral side (65 %) and in the posterior lunules (89 %). Lesions
extended to the adjacent posterior interambulacrum (Fig. 6.7b). These lesions were
found in the larger specimens (46.4 to 106.0 mm in length), with a mode (38 %) at
62.7 mm. Some specimens had small black spots with surrounding lesions on the
oral surface (Fig. 6.7a, b). The authors indicated that oils from fishing boats and
commercial crafts are common in this area, and these conditions probably lead to
lesions of the epithelium of the test of E. micropora that subsequently became
infected.
Sea Urchins
Sonnenholzner et al. (2009) studied the effects of predation by fish (the Mexican
hogfish Bodianus diplotaenia), and lobsters (red spiny lobster Panulirus peni-
cillatus, and rocky lobster Scyllarides astori) on the herbivore E. galapagensis.
The authors sampled 10 highly fished and 10 (putatively) lightly fished shallow
rocky reefs in the southeastern area of the Galápagos Islands. Eucidaris gala-
pagensis showed a negative association with non-coralline algae. In addition,
pencil urchins were less abundant where there were many predators indicating an
indirect positive association between predators and non-coralline algae occurred.
Fishing appeared to affect this trophic cascade. The spiny lobster P. penicillatus,
the slipper lobster S. astori, and the hogfish B. diplotaenia were significantly less
abundant at highly fished sites. Sea urchin density was higher at highly fished sites.
Non-coralline algae were nearly absent from highly fished sites, where a contin-
uous carpet of the anemone Aiptasia sp. occurred. The algal assemblage was
mainly composed of encrusting coralline and articulated calcareous algae.
Sonnenholzner et al. (2011) conducted an experiment for measuring lobster
predation on Eucidaris galapagensis in Galápagos. Six slipper lobsters Scyllarides
astori (19.0–26.2 cm total length, 0.50–0.91 kg) and six red spiny lobsters Pan-
ulirus penicillatus (25.0–28.0 cm total length, 0.60–1.13 kg) were used as pre-
dators. The sea urchin E. galapagensis (36.7–51.6 mm test diameter, average
44.9 mm) was offered as prey to each lobster. Two treatments were used for each
species of lobster and a control treatment of sea urchins with no lobsters. Sea
urchin survivorship was measured for 3 days. Spiny lobsters ate three sea urchins
and attacked an additional four urchins. Slipper lobsters did not interact with sea
urchins. This experiment identified spiny lobsters as potential predators of urchins.
There are many other important predator–prey interactions in the system (Witman
et al. 2010). Slipper lobsters might consume sea urchins in the wild, even though
they did not prey on them in the laboratory. Predation in aquaria does not nec-
essarily indicate predation rates or preferences in the field only that these predators
are capable of eating the prey offered to them. Still, observations in aquaria are
wholly consistent with the limited field observations of trophic interactions
(Sonnenholzner et al. 2011).
Although the taxonomy and distribution of echinoderms in the Galápagos

Islands are well known, their biological interactions with parasitism are not.
Echinoderms are intensively parasitized by eulimid gastropods (Pearse and
Cameron 1991), but there has been little research about the eastern Pacific eulimid
species and their ecology (Skoglung 2004). Sonnenholzner and Molina (2005)
provided information on the sea urchin–eulimids host–parasite relationship along
the southern and western regions of the Galápagos Islands. A total of 1,855
specimens of E. galapagensis were examined. Of them, 925 individuals were
mainly affected by Sabinella shaskyi. The maximum number of parasites in a host
was 63 (both adult and juvenile parasites), seven on the same spine. Between 25 %
and 42 % of the host spines were affected. Eucidaris galapagensis (n = 930) had
another parasite, Pelseneeria sp., which crawled freely on the host test and was
commonly found attached but not on or inside the spines as S. shaskyi was.
Pelseneeria sp. affected the gonads of the host in weight (a low gonad index) and
color (female gonads were pale). The infestation reached high percentages (95 %)
and the maximum number of parasites on the same host was five (adult parasites
only). At night, snails occasionally occurred on the substrate near sea urchins
(J. Sonnenholzner, pers. obs.), suggesting that snails may migrate between sea
urchins. It is unknown how S. shaskyi affects urchins at the population level, but
loss and reduction of spine number (Metz 1994; Sonnenholzner and Molina 2005)
would appear to increase susceptibility to predation and energetic costs for spine
repair (Heatfield 1971; Denny and Gaylord 1996).
Parasites dominate food web links (Lafferty et al. 2006). For this reason,
Sonnenholzner et al. (2011) took advantage of the variation in fishing pressure to
investigate complex indirect effects on two eulimid snails that parasitize the most
abundant sea urchin E. galapagensis in the Galápagos. Past work in the Galápagos
suggests that fishing reduces lobster and fish densities and this relaxation of pre-
dation pressure indirectly increases sea urchin densities, creating the potential for
complex indirect interactions between fishing and parasitic snails. To measure
indirect effects of fishing on these parasitic snails, the spatial relationships among
sea urchins, parasitic snails, commensal crabs, and large sea urchin predators
(hogfish and lobsters) was investigated. Parasitic snails had higher densities at sites
where sea urchins were abundant, probably due to increased resource availability.
Commensal crabs that shelter under sea urchin spines, particularly the endemic
Mithraculus nodosus, preyed on the parasitic snails in aquaria. Snails were less
abundant at field sites where these crabs were common. In aquaria, hogfish and
lobsters readily ate crabs, but crabs under sea urchin spines were protected from
predation. This would lead to a facultative mutualism between commensal crabs
and sea urchins. In the field, fishing appeared to indirectly increase the abundance
of sea urchins and their commensal crabs by reducing predation pressure from fish
and lobsters. Fished sites had fewer snails per sea urchin, probably due to
increased predation from commensal crabs. However, because fished sites also
tended to have more sea urchins, there was no significant net effect of fishing on
the number of snails per square meter. These results suggest fishing can have
complex indirect effects on parasites by altering food webs.
Sea Stars
Arm regeneration in starfish is related to sub-lethal predation. Sonnenholzner and

Lawrence (2002) collected few starfish with signs of sub-lethal predation at Los
Frailes, Manabí. They found that one of the 54 specimens collected in their study
had four arms and two others had regenerating arms. They observed that the
parrotfish Scarus perrico is the predator of the tan starfish P. unifascialis. None-
theless, the strength of the body wall, and the availability of food may affect the
susceptibility to predation (Lawrence 1987). The low incidence of wounded or
regenerating arms in the population of P. unifascialis suggests parrotfish are less
inclined to feed on it, probably due to its strong body wall.
6.3.5 Interacting Effects of Echinoderms on Benthic Communities
Sea Urchins
Of all benthic consumers, sea urchins are known to be one of the most influential
taxa affecting community structure (Lawrence 1975). Sea urchins have the
potential to influence the distribution, relative abundance and species composition
of algae and other sessile invertebrates (Kitching and Ebling 1961, Paine and
Vadas 1969), thereby determining the biomass, diversity and productivity of
marine communities. In Ecuador, studies on this topic have been done only in the
Galápagos Islands, with the pioneer work of Peter W. Glynn in the late 1970s. In
an assessment of coral growth versus coral attrition from grazing, Glynn et al.
(1979) showed that E. galapagensis reduced coral growth in the Galápagos. He
reported individuals of E. galapagensis grazing heavily on live hermatypic po-
cilloporid corals, including Pocillopora damicornis, P. elegans, P. capitata and
also Pavona clavus. Given that urchins were more abundant along the reef edge,
grazing was more intense there. The study suggested E. galapagensis might limit
reef growth, especially by interfering with the development of reef frame and its
lateral expansion. By comparing Panamá to the Galápagos, Glynn et al. (1979)
attributed the lower buildup of coral in the Galápagos to a much heavier sea urchin
grazing and suggested this difference was due to a lower predation of sea urchins
in the Galápagos.
Almost a decade later, Glynn (1988) showed that the distribution and popula-
tion densities of E. galapagensis changed dramatically following the 1982–83 El
Niño disturbance. Once abundant in off-reef and reef edges areas, sea urchins
moved to the center of the dead reef frameworks of Porites and Pavona, causing
bioerosion rates that ranged from 49 to 99 g CaCO3 m-2 day-1. Before 1983,
E. galapagensis at high densities grazed extensively on live pocilloporid corals
(Glynn et al. 1983). However after the El Niño event these corals were locally
extinct in Floreana and the remnant non-pocilloporid corals, with less than 0.1 %
of cover, were avoided by the urchins. Glynn (1988) suggested that the destructive
effect of sea urchins were the result of erosion of reef structures rather than direct
mortality of surviving corals. The impact of E. galapagensis on habitats other than
coral reefs was described by Brandt (2003). This study was done at a depth of 5 m
off Caamaño Islet (immediately to the south of Santa Cruz Island, outside Acad-
emy Bay). It consisted on removing all E. galapagensis from a 6 9 6 m treatment
plot. After 12 weeks macroalgal biomass was 20 times higher in the treatment than
in the control plot, which had densities of *40 ind m-2. In addition, benthic
diversity, including sessile invertebrates and algal species, was higher in the plot
devoid of E. galapagensis, while plots with high densities of the sea urchin were
characterized by crustose coralline algae and bare rock. Similar to Glynn (1988),
who reported that coral erosion rates by urchins were slower when the damself-
ishes Stegastes beebei and Stegastes arcifrons were present, Brandt (2003) also
noted that the abundance of the green foliose algae Ulva sp. was maintained in
plots where both urchins and damselfishes were abundant. The role of damselfishes
in mediating the impact of urchins thus seems important. Irving and Witman
(2009) explored the interactions between the yellow-tail damselfish Stegastes
arcifrons with E. galapagensis and with the ‘‘green’’ sea urchin L. semitubercul-
atus. They found that the ability of sea urchins to switch habitats by removing
algal turf to create coralline barrens was dependent on the absence of damselfish as
well as the species of urchin. In the presence of damselfish, neither species of
urchin caused any significant change to algal assemblages. However, in their
absence, only L. semituberculatus was able to switch the habitat towards greater
cover of encrusting algae by grazing filamentous turfs. In addition, one of their
results contrasts with previous findings about the impact of E. galapagensis on the
abundance of macroalgae described above. Irving and Witman (2009) found that
removal of E. galapagensis from algal communities dominated by encrusting
coralline algae had no effect on algal community structure and abundance, while
the removal of L. semituberculatus resulted in a significant decrease of encrusting
coralline algae and an increase in the abundance of green and other erect algae.
Since the densities of E. galapagensis and L. semituberculatus manipulated by
Irving and Witman (2009) were different, they concluded that relatively high
densities of L. semituberculatus impact algal habitats, but relatively low densities
of E. galapagensis do not. Nevertheless they noted that the densities manipulated
were the same as natural densities at the study site. The study of the impacts of sea
urchins on benthic communities has greatly improved our understanding about the
effects of their grazing and bioerosion in the Galápagos. However more research is
needed with other species of sea urchins, such as T. depressus and D. mexicanum
in the Galápagos, and also with sea urchin species on the coast of Ecuador.
6.4 Fisheries and Aquaculture
6.4.1 Sea Cucumbers
Isostichopus fuscus
Sea cucumbers received little or no attention from the scientific community in
Ecuador prior to the beginning of commercial harvesting of I. fuscus in the early
1990s (Aguilar et al. 1993; Richmond and Martínez 1993; Sonnenholzner 1997).
Since then, most studies focused on the biology of this species and other poten-
tially commercial species (such as S. horrens and H. theeli (Sonnenholzner 2003;
Mercier et al. 2004; Hearn 2005, Hearn and Pinillos 2006; Toral-Granda and
Martínez 2007), on the social and political implications of the fishery, especially in
the Galápagos Islands (Bremner and Pérez 2002), and on the potential for aqua-
culture (Mercier et al. 2004). In addition, the Charles Darwin Foundation in
Galápagos undertook a long term subtidal rocky reef monitoring program to obtain
information on the size and abundance of the sea cucumbers found in the Galá-
pagos Islands was recorded (Murillo et al. 2002a). No studies have been done in
Ecuador mainland.
About 40 species of commercial sea cucumbers, including I. fuscus are har-
vested throughout the world’s oceans (Conand 2004). The fishery of I. fuscus has
been relatively well documented since the start of its legal fishery in 1999. This
fishery began in the mainland Ecuador in 1988. Four years later it was centered
mainly on the western side of the Galápagos Archipelago because the sea
cucumber population rapidly collapsed on the mainland. This caused the transfer
of fishing activity to migrate to the Galápagos Islands (Aguilar et al. 1993; De Paco
et al. 1993; Camhi 1995; Powell and Gibbs 1995). At least 100 fishers, mainly
from continental Ecuador, began exploiting sea cucumbers in Galápagos in 1992
(Shepherd et al. 2004) although the first signs of sea cucumber harvesting were
detected in 1991 (Carranza-Barona and Andrade-Echeverría 1996). Nevertheless
the incipient fishery lacked any form of regulations or technical guidelines for its
sustainable management (Martínez 2001). After several years of unregulated
exploitation, an experimental fishery was opened in 1994, with a catch quota of
500,000 individuals over a 2-month period. However, the fishing season was
closed after 6 weeks, with an estimated catch of 8–12 million sea cucumbers (De
Miras et al. 1996). Ecuador’s General Fisheries Direction reported that ten com-
panies were exporting dried sea cucumbers. Data showed that sea cucumber
fishery was less than 0.04 t (fresh total weight) from 1983 to 1988 and 50.3 t from
1989 to 1995. The highest catch (30 t) was reported in 1995. These products were
all exported, 67 % to the U.S. and 33 % to Taiwan. The maximum value paid per
net kg in Ecuador was USD$30 (Sonnenholzner 1997). Although it is not the most
valued sea cucumber species (Conand 2004), this species has been recognized as
an important food item in the Chinese market. Sonnenholzner et al. (1997)
determined the proximate composition of the body wall of I. fuscus collected in
April 1993 at the eastern coast of the Fernandina Island along the Bolívar Channel
from Punta Mangle to Punta Espinoza. The concentrations (per cent wet weight) of
ash (1.0–1.5 %), protein (2.0–4.0 %), fat (3.1–3.6 %), and carbohydrates (\1.0 %)
were all low as the water content was high (93.5 %). Carbonate and chloride
concentrations were very low (\1.0 %). This indicates the body wall of I. fuscus is
not a high-quantity protein food source.
The I. fuscus fishery was banned from 1995 to 1998. In 1998, the Galápagos
Marine Reserve (GMR) was created, along with a novel management system based
on the participation of local stakeholders (Galápagos National Park Service,
Charles Darwin Research Station, tourism, fishing and local guide sectors). One of
the first issues addressed by the new participatory system was to agree upon
management measures to re-open the sea cucumber fishery in 1999. Since then,
this fishery has been characterized by annual negotiations, conflicts, inconsisten-
cies and a deterioration of the resource which has resulted in its collapse (Shepherd
et al. 2004). At first, local fishers remained focused on whitefish and lobster, but as
the sea cucumber fishery became established, this figure rose to a maximum of
1,229 active fishers in 2000. After this year, the number declined to just 436 active
fishers in 2007 (Murillo and Reyes 2007). In the first official fishing season (1999)
4.4 million sea cucumbers were harvested, 4.9 million in 2000, 2.7 million in
2001, 8.3 million in 2002 and 5.0 million in 2003 (Toral-Granda et al. 2006).
From 2002 onwards quotas were set in place which generally outstripped the
capacity of the resource, and catches declined steadily until the fishery was closed
in 2006. After pressure from fishers and local politicians, it was re-opened in 2007.
Overall, more than 25.3 million sea cucumbers (over 6800 t live weight) have been
extracted legally from the GMR since 1999 (Murillo et al. 2002a, b). Since its
re-opening in 1999, the fishery for I. fuscus in the Galápagos Islands has been
regulated by a minimum size limit for both fresh and processed sea cucumbers
(among other regulatory measures). The size limits are not regarded as a useful
tool by the fishers because they believe that sea cucumbers change length too
easily to be accurately measured. Nonetheless, local fishers mostly agree, in
principle, to have minimum legal size limits rather than none at all. Both man-
agement authorities and scientific bodies concur that this regulation is not as
effective as others.
Sea cucumber density displayed an irregular pattern with high values of
[0.4 ind m-2 between 1995 and 1997, but with very low values in 1999 (only
0.13 ind m-2). No surveys were made in 1998. However, after 1999, there was a
rapid recovery to prior levels, peaking in 2002, after which densities dropped
rapidly and showed no further signs of recovery. At the same time, catch per unit
effort (CPUE) measured by onboard observers, increased from 90 to 136 ind di-
ver h-1 (between 1999 and 2002, respectively), and then dropped to 55 ind diver
h-1 by 2005. The decline of I. fuscus populations has led to the illegal exploitation
of three other, less valuable, sea cucumber species in the GMR: Stichopus horrens,
Holothuria kefersteinii and H. atra (Hearn and Pinillos 2006; Toral-Granda et al.
2006).
Several methods, such as molecular phylogeny, morphology, visual, and skeletal
features, have been used to try to identify different species of sea cucumbers while
live or fresh (Uthicke and Benzie 2003; Uthicke et al. 2004). Concerned with the
illegal fishery of I. fuscus in the GMR, Toral-Granda (2005) studied the shapes and
combination of microscopic spicules from the skeletal components in the body wall
and tube feet of I. fuscus. Spicules are shaped like tables and C-shaped rods in the
body wall and perforate supporting tables in the tube feet plates and perforate
supporting tables. This provides a means to diminish the illegal trade in this species,
and support conservation efforts. Processed I. fuscus looks similar to some other
species and can be almost impossible to distinguish from them. This includes S.
horrens, which is harvested illegally together with I. fuscus (Arellano 2004). This
methodology can be used in a quick, precise and easy identification. It is cost-
effective and can be done in a timely manner by custom officers to distinguish this
species from others.
Isostichopus fuscus has been intensively fished along the mainland and around
the Galápagos Islands. Therefore its populations have thus been severely depleted
over the past decade. Mercier et al. (2004) first attempted to breed I. fuscus in land-
based installations on the coast of Ecuador. They studied spawning, fertilization,
larval rearing, disease control and juvenile growth. Data pooled from monthly
trials conducted over 3 years indicate that, under optimal conditions, juveniles can
be grown to a length of ca. 8 cm in length in 3.5 months. The survival rate is
typically between 30 and 50 %. Furthermore, preliminary experiments showed
that growth of young I. fuscus in abandoned shrimp ponds is a promising option.
Overall, this study demonstrates that I. fuscus can be reared in captivity. Thus, this
provides an alternative to fisheries, or a way to maintain sustainable harvests and
eventually contribute to the restoration of the natural populations.
6.5 Threats
6.5.1 Anthropogenic and Environmental Effects
The fuel tanker Jessica grounded on Shiavioni reef at the entrance of Wreck Bay,
San Cristóbal Island (Galápagos Islands) on 16 January 2001. The vessel carried
approximately 240,000 gallons of fuel oil consisting of 160,000 gallons of Diesel
Oil #2 (DO#2) and 80,000 gallons of Intermediate Fuel Oil 120 (IFO 120, or
bunker fuel). By 29th January, most of the remaining 180,000 gallons of fuel oil
(75,000 gallons of IFO 120 and 105,000 gallons of DO#2) had escaped from the
hull and dispersed to waters within the archipelago. The results of the Galápagos
Coastal Oil Survey indicate that environmental contamination from the Jessica
spill was broadly and thinly dispersed throughout the southwestern archipelago.
Survey work confirmed that bunker fuel went ashore at sites as remote as the
coastlines of Fernandina and Isabela and in sensitive habitats such as mangroves
(Edgar et al. 2002a; Marshall and Edgar 2003).
Intertidal and shallow subtidal sites on shores affected by oil in San Cristóbal
and Santa Fé Islands had been sampled prior to the oil spill. These data were
compared with data collected at the same sites 1 month later and at reference
locations that had not received oil. No clear effects of oil were detected, indicating
that impacts on shore communities of seaweeds, marine invertebrates and fishes
were probably slight. Increases in the density of several algal, sea urchin, hydroid
and fish taxa were detected adjacent to the wreck, and were probably caused by oil
pollution related to the Jessica spill (Edgar et al. 2002a).
The greatest oil pollution of shoreline contamination occurred on Floreana and
Isabela Islands. Lower levels of oil pollution were found at Santa Fé, Santa Cruz,
and San Cristóbal Islands and secondary contamination (oiled debris and foam)
was found on Española and Fernandina Islands. Only one survey site contained
sufficient coverage of oil to be termed ‘‘moderate’’. All other sites had ‘‘light’’ or
‘‘very light’’ surface oil (Lougheed et al. 2002).
Along the coast of mainland Ecuador, the absence of historical baseline data for
the invertebrate community is a real problem for determining impacts of this kind
of event on invertebrates.
Fishing has indirect effects on ecosystems, particularly through trophic cas-
cades (Sala et al. 1998; Pinnegar et al. 2000; Jackson et al. 2001; Bascompte et al.
2005), in which sea urchins are important grazers. For instance, because sea
urchins can structure reef communities (Schiel and Foster 1986; Hughes et al.
1987; Pearse and Hines 1987), the removal of sea urchin predators (large lobsters
and fish) by artisanal fishing can increase the abundance of sea urchins. They can
then overgraze algae (Tegner and Dayton 1981; Tegner and Levin 1983; Siversten
2006). Overgrazing probably magnified impacts of the El Niño 1982–1983 through
a cascade of indirect effects involving population expansion of populations of
grazing sea urchins.
Edgar et al. (2010) indicated that shallow reef habitats across the central
Galápagos Archipelago underwent major transformation at the time of the severe
El Niño 1982–1983 warming event with consequences that persist today (Bensted-
Smith 2002). Heavily grazed reefs with crustose coralline algae (urchin barrens)
replaced former macroalgal and coral habitats, resulting in large local and regional
declines in biodiversity. Also, the range of corals decreased. Their ranges and their
populations are now largely confined to localized areas in the west (Glynn 1994)
and the north of the archipelago (Banks et al. 2009). One echinoderm, the 24-rayed
sunstar Heliaster solaris, has not been sighted since 1983 (Cairns 1991; Hickman
1998; Edgar et al. 2010). It is considered probably extinct in coastal environments
in Galápagos as probably extinct, because it has not been seen for [28 years (Red
List IUCN 2008). Marine protected areas with adequate enforcement are predicted
to ameliorate but not eliminate ecosystem impacts caused by increasing thermal
anomalies associated with El Niño and global climate change (Robinson 1985;
Edgar et al. 2010).
6.6.1 Biodiversity
1. A diverse and complex set of marine ecosystems exist along the coast of
mainland Ecuador and the Galápagos Islands. These include rocky shores,
sandy beaches, belts of sandy banks, estuaries, mangrove forests, deep sea
mounts, and cliffs associated with pocket beaches. All these systems provide a
high variety of subtidal and intertidal habitats for a wide range of organisms.
Nevertheless information about echinoderms is limited and dispersed. A con-
tinuous, long-term monitoring program, which integrates all habitat types and
latitudinal variation, is needed in order to understand how environmental
conditions influence echinoderm distribution and diversity. In addition, almost
nothing is known about echinoderms of bathyal and abyssal zones, especially in
the vicinity of the Galápagos Archipelago.
2. Most studies in the inner and outer Gulf of Guayaquil have focused on estuarine
production and mangrove ecology. No information on the echinoderm fauna
inhabiting this complex and large ecosystem is available, especially on species
associated with rocky, sandy and muddy bottoms, such as sand dollars, sea
biscuits, sea stars, brittle stars, and sea cucumbers. Viable population sizes
should be determined for key groups of taxa and species. They should be
monitored throughout their distribution range to quantify changes in population
size.
3. No studies have focused on the relationship between climate change and bio-
diversity, neither for Ecuador mainland nor for the Galápagos Islands. The
taxonomic information available is largely at the most basic descriptive level.
There is little information about the phylogeny of species and their geo-
graphical distributions.
4. Scientists should identify priority areas and taxonomic groups of interest for
completing biological inventories. They should encourage the development of a
geographic information system for detecting areas of high diversity and design
species/area curves for predicting extinction patterns to use to formulate poli-
cies for conservation and economic development (Cruz et al. 2003).
6.6.2 Ecology
1. Population dynamics of key sea urchin species should be continuously moni-

tored, because many of the ecological changes in Galápagos might have been
exacerbated indirectly through trophic cascades, via loss of predation and
disease on E. galapagensis (Sonnenholzner et al. 2011).
2. In general, very little information about plankton ecology is available over in

Ecuador. There is a lack of information on echinoderm larval ecology, dis-
persion, and its development along the mainland Ecuador and the Galápagos
Islands. This field of research must be developed, because it is important for the
future. Key life history traits of stenotopic species need to be known in order to
determine their capacity to respond to extreme changes in temperature and
salinity in their environments. Additionally, samples of plankton and larvae of
echinoderms in the Reference Collections are not being well maintained. There
is a need to create a GIS-Database not only for biological but also for ocean-
ographic data.
3. Studies on meiobenthos on the mainland of Ecuador and Galápagos should be
continued. However it is necessary to carefully choose study sites for long-term
research to monitor changes in biodiversity.
4. As suggested by Okey et al. (2004) for the Galápagos Islands, mass-balanced
trophic models (Ecopath/Ecosim/Ecospace) for primary and secondary pro-
duction for rocky reefs, sandy and muddy beaches, and estuaries environments
along the mainland of Ecuador should be constructed. This is necessary to
detect those habitats that may act as sinks for oceanic plankton, because reef
systems and estuarine environments may be strongly coupled with physical
oceanographic conditions. This information should be available for the early
life stages of different species of echinoderms in Ecuador.
6.6.3 Fisheries
1. Although several factors contribute to variability in communities of algae,

fishes, lobsters, sea urchins, sea urchins parasites, and crabs, in Galápagos,
evidence suggests that artisanal fishing has both direct and cascading effects
throughout the community (Sonnenholzner et al. 2009, 2011; Edgar et al.
2010). Nonetheless, the magnitude, rate and severity of these changes are still
unknown for echinoderms (Sonnenholzner et al. 2011), and community struc-
ture (Ruttenberg 2001) because most studies have been observational. More
experimental work is needed in order to understand top-down control in the
Galápagos.
2. Capture-recapture studies for the sea urchin T. depressus (tagged specimens)
should be conducted to estimate basic population parameters, such as, age,
growth rate, natural mortality, and movement patterns (Sonnenholzner 2011).
These studies, combined with other population surveys, should provide baseline
data to develop an adequate strategy of coastal management for a potential
planned fishery of T. depressus in the mainland and the Galápagos Islands.
3. Techniques need to be developed for production of commercial juvenile sea

cucumbers (i.e. I. fuscus), and sea urchins (i.e. T. depressus) through laboratory
cultures for aquaculture and/or repopulation.
4. A better understanding of the social and political aspect of the sea cucumber
fishery is required. A key external stakeholder in the overexploitation of sea
cucumbers in Galápagos is the Asian market. Imaginative solutions are required
in order to escape the current situation, whereby social pressure forces the
fishery to open after the slightest hint of recovery. This jeopardizes both the
long term recovery of the resource as well as the economic stability of the
fishers.
6.6.4 Threats
1. Impacts can be grouped into six main categories: (a) fishery exploitation; (b)
global climate change; (c) invasions of exotic species; (d) pollution; and (e) loss
of habitat due to increased human population of the coastal zones both in
Ecuador mainland and in the Galápagos Islands.
2. The collateral effects of the sea cucumber fishery in the Galápagos Islands are
not fully understood. What is the damage to biodiversity of illegal fishing
camps on pristine islands? To what extent does the presence of a licensed sea
cucumber merchant in the islands open a door for the exploitation of other
protected species such as sea horses, sharks and sea lions? Socially and eco-
nomically, who has really benefited from this fishery?
3. Future research should address the following main questions considering the
immune response of two sea urchins species: E. vanbrunti (at the mainland) and
E. galapagensis (at the Galápagos Islands) to establish the exact chain of
causation that could kill different sea animals after low-level oil contamination,
or from global warming change (as was suggested by Romero and Wikelsky
2002). Is the immune response playing a direct role like bio-predictors under a
stress response?
Acknowledgments We would like to express our thanks to Manuel Cruz (formerly at the
Instituto Oceanográfico de la Armada del Ecuador), Soledad Luna, Kate Clark and Priscilla
Martínez (Instituto Ecuatoriano NAZCA de Investigaciones Marinas), Luis Molina (Charles
Darwin Foundation, CDF), and Manuel Burgos (Universidad de Especialidades Espíritu Santo,
UEES) for their assistance. We also thank Angel Chiriboga, Xavier Romero, and CDF for
photographical material included in this manuscript.
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Chapter 7
Echinoderms from Venezuela: Scientific
Recount, Diversity and Distribution
César Lodeiros, Alberto Martín, Vanessa Francisco, Nicida Noriega,

Yusbelly Díaz, Jeny Reyes, Orangel Aguilera and José Alió
C. Lodeiros (&)
Instituto Oceanográfico de Venezuela, Universidad de Oriente y Fundación para la
Investigación y Desarrollo de la Acuicultura del Edo. Sucre, Cumaná, 6101, Venezuela
e-mail: cesarlodeirosseijo@yahoo.es
A. Martín Y. Díaz
Laboratorio de Crustáceos Peracáridos. Centro de Biodiversidad Marina. Departamento de
Estudios Ambientales - INTECMAR, Universidad Simón Bolívar, Caracas, 1080, Venezuela
e-mail: amartinz@usb.ve
Y. Díaz
e-mail: yusdiaz@usb.ve
V. Francisco
Laboratorio de Ecología de Ecosistemas de Arrecifes Coralinos, CINVESTAV-IPN Unidad
Mérida, 97310, Yucatán, Mexico
e-mail: vanewagen@gmail.com
N. Noriega
Laboratorio de Biología Marina y Laboratorio de Bioquímica, Universidad Simón Bolívar,
Caracas, 1080, Venezuela
e-mail: nicidanoriega@gmail.com
J. Reyes
Laboratorio de Zoología de Invertebrados, Universidad del Zulia, Maracaibo 4001 y
Fundación para la Investigación y Desarrollo de la Acuicultura del Edo. Sucre,
Cumaná, 6101, Venezuela
e-mail: jenyreyesl@yahoo.es
O. Aguilera
Museu Paraense Emilio Goeldi, Coordenação de Ciências da Terra e Ecologia,
Departamento de Geociências Av. Perimetral, 1901 Terra Firme, Belém, PA,
CEP: 66077-830, Brazil
e-mail: orangel.aguilera@gmail.com
J. Alió
Centro de Investigaciones Agrícolas de Sucre y Nueva Esparta, Instituto Nacional de
Investigaciones Agrícolas, Cumaná, Estado Sucre, 6101, Venezuela
e-mail: jalio@inia.gob.ve

236 C. Lodeiros et al.
7.1 Introduction
Venezuela is located in northern South America between 00°450 and 15°400 N, and
between 59°470 and 73°220 W (Fig. 7.1). Its coastal waters include the Caribbean
Sea and the west central Atlantic Ocean, which gives it a special biogeographical
position. Ecosystems in the Caribbean Sea include sandy beaches, rocky shores,
seagrass meadows, coral reefs, coastal lagoons, soft bottom communities and
mangroves, which results in a high environmental variability as a consequence of
periodic coastal upwellings due to the trade winds. The Atlantic zone is mainly
influenced by the Orinoco River which flows north towards the Caribbean Sea and
promotes a brackish water environment in the coastal areas of its delta and the
Gulf of Paria, a situation that increases the environmental variability. On the other
hand, the 3,964 km coastline (68 % in the Caribbean Sea, 21 % in the Atlantic
Ocean and 11 % in the islands) is very diverse both in geology and topography,
and a great diversity of geomorphologies coexist including brackish lakes,
lagoons, bays, peninsulas and other geologic formations. Such environments,
together with 14 archipelagos, encompass multiple ecosystems resulting in one of
the most diverse areas in the world (Aguilera et al. 2003; Miloslavich et al. 2005;
Miloslavich and Klein 2008).
Sea surface temperature (SST) along the Venezuelan coastline is mainly
affected by upwelling, and presents yearly ranges between 20 and 29°C with daily
maximum variations of 2.5°C (Penchaszadeh et al. 2000). There are 12 upwelling
zones that could be identified along the Venezuelan coast using images from the
‘‘Advanced Very High Resolution Radiometer (AVHRR)’’ to determine the SST.
In western Venezuela, the upwelling zones are located to the east of the Guajira
Peninsula; in the Gulf of Venezuela and in Punto Fijo coast, western Paraguaná
Peninsula; in Cabo San Román and in Puerto Cumarebo, eastern Falcón state.
Upwelling is not frequent along the central coast of Venezuela. There are several
upwelling systems in the eastern coast between Puerto La Cruz and Cumaná, to the
north of the Araya Peninsula and Margarita Island, and the central coast of Sucre
state, near Carúpano. Coastal upwelling is frequent in all these locations, associ-
ated with the relatively high speed and predominance of the trade winds during the
first 6 months of the year. The intensity and propagation of the upwelling is
considerable and extends beyond 2,00,000 km2, contributing to establish the most
productive zone of the Caribbean Sea. Offshore from the Paria Peninsula and to the
north of the island of Trinidad, upwelling occurs during limited periods of variable
intensity. The cold water masses occasionally form eddies that migrate northeast,
probably due to the movement of less dense warm water masses from the Gulf of
Paria (Müller-Karger and Varela 1989; Penchaszadeh et al. 2000).
All these characteristics that result from environmental variability in the
Venezuelan coasts have allowed the establishment of 13 ecoregions according to
their ecological, physical and oceanographic characteristics (Fig. 7.1; Miloslavich
and Klein 2008).
7 Echinoderms from Venezuela 237
Fig. 7.1 Marine ecoregions of Venezuela (Miloslavich and Klein 2008)
Ecoregion 1: Delta of the Orinoco River and Atlantic Front. This ecoregion is
characterized by estuarine and marine ecosystems with a large sedimentary load,
high turbidity, sandy and muddy bottoms, with a well developed mangrove
community.
Ecoregion 2: Eastern upwelling region, which encompasses the area from the
eastern Paria Peninsula to Mochima Bay. This ecoregion is characterized by a
shallow continental shelf with coarse sandy sediments, a large rocky shore and few
coralline reefs. The region is highly productive due to the upwelling fronts. It
includes the islands of Margarita, Coche, Cubagua, Los Frailes, Los Hermanos and
Los Testigos Archipelago.
Ecoregion 3: Axis Unare-Píritu-Tacarigua, from Barcelona to Cabo Codera. This
ecoregion is characterized by the presence of large coastal lagoons with mangrove
community and long sandy beaches, with a large sediment load from the Unare
and Tuy rivers. There is a large secondary production in the region, mainly of
crustaceans and bivalves.
Ecoregion 4: Central coast, from Cabo Codera to Patanemo Bay. This ecoregion is
characterized by a narrow coastal fringe with rocky shores and abundant sandy
beaches of coarse sediments. The coral communities are associated with rocky
substrates, as well as with some mangrove communities and seagrass.
Ecoregion 5: Golfo Triste, from Patanemo Bay to the mouth of the Tocuyo River.
This ecoregion is characterized by a relatively large and shallow continental plat-
form, rocky shores and well developed coralline barriers, particularly in the National
Parks Morrocoy and San Esteban. The region has a moderate secondary production.
Ecoregion 6: Tocuyo region, from the Tocuyo River to the isthmus of the Paraguaná
Peninsula. This ecoregion is characterized by sandy beaches and rocky cliffs with a
high sediment load from the Rivers Tocuyo, Hueque and Ricoa, and low water
transparency.
Ecoregion 7: Paraguaná region, from the isthmus of the Paraguaná Peninsula to

Punta Cardón. This ecoregion is characterized by emergent zones of rocky shores
and great banks of macroalgae with carbonated sandy beaches and high primary
productivity due to seasonal upwelling fronts.
Ecoregion 8: Golfete de Coro, from Punta Cardón to the mouth of the Maracaibo
estuary. This ecoregion is characterized by shallow waters and high sedimentary
load and suspended materials from the Golfo de Coro. There is a high secondary
production mainly associated with penaeid shrimps and sciaenid fishes (corvinas).
Ecoregion 9: Estuarine region of Lake Maracaibo. This ecoregion is characterized
by a continuous perturbation caused by the dredging of the navigation channel
along the strait of Maracaibo. The region has high primary and secondary
productions, the latter associated with crustaceans (shrimp and crabs), bivalve
mollusks and fishes. It is threatened by pollution due to the intense activity of the
oil industry and sewage discharges.
Ecoregion 10: Golfo de Venezuela. This ecoregion is characterized by a wide
coastal platform with shallow bottoms (\50 m). It has a high primary and
secondary production of shrimp and fishes due to the upwelling fronts. Bottom
communities are dominated by detritivorous organisms.
Ecoregion 11: Oceanic Islands which includes all emerged land with the exception
of the Eastern Upwelling Region. This ecoregion is characterized by islands and
archipelagos, with reefs, seagrass and well developed mangrove areas. It has a
high biodiversity. It is a zone shared by marine turtles and birds.
Ecoregion 12: Oceanic area including the Exclusive Economic Zone of Venezuela,
an area with depths beyond 200 m. The biodiversity of this ecoregion is little
known. It has zones with moderately high primary productivity due to their
proximity to upwelling coastal zones.
Ecoregion 13: Cariaco Trench, the depression of the continental shelf below
150 m deep in front of Barcelona and Mochima Bay in eastern Venezuela. This
ecoregion represents a special ecosystem with maximum depths close to 1,400 m
and anoxic water masses below 250 m. The productive processes are based mainly
on chemosynthesis.
In Venezuela, echinoderms have been poorly studied in contrast to other groups
of marine organisms like fish, crustaceans (decapods and peracarids) and mollusks,
which have been grouped, classified and analyzed taxonomically (Rodríguez 1980;
Valdez and Aguilera 1987; Cervigón 1994; Lodeiros et al. 1999; Díaz and Martín
2001; Martín et al. 2002; Martín and Díaz 2003, 2005, 2007; Díaz et al. 2005).
Taxonomic and ecological studies on local echinoderms are scarce and limited, in
spite of the fact that they represent an important component of the biota in the
marine coastal zones (Penchaszadeh 2003). A compilation of studies on echino-
derms of Venezuela is presented here, which represents an effort of researchers
from several national and international institutions, whose analyses are a current
description of the status of these marine invertebrates.
7.2 Research
Along the Venezuelan coast there are few studies pertaining to species of the
Phylum Echinodermata, and most have focused mainly on the northern Caribbean
Sea region. Studies made in the Atlantic region have been financed by the
Venezuelan oil industry (PDVSA) and include the Environmental Base Line of the
Platform in the Orinoco River delta (Martín et al. 2007) and the Venezuelan
Atlantic Front (Gómez et al. 2005).
7.2.1 Taxonomy
The first studies dealing with the Phylum Echinodermata (Families: Astropectin-
idae, Luidiidae and Echinasteridae) in Venezuela, came from the Belgian research
cruise ‘‘Mercator’’ between 1935 and 1936, which followed a trajectory between
Florida and Venezuela (Engel et al. 1960). Furthermore, Ummels (1963) and
Jangoux (1978) published studies on asteroids where they incorporated locations
from the country.
At a regional level, the work on echinoderm taxonomy was initiated in the
1960s with the publication of Zoppi de Roa (1967), who made an exhaustive
identification of the species belonging to the different classes along the Venezuelan
coast, from where she reported 65 species. On the other hand, Work (1969)
reported 27 species for Los Roques Archipelago National Park distributed as
follows: eight echinoids, 11 ophiuroids, three asteroids and five holothuroids.
Later, Boada (1970) conducted a general study on the taxonomy of echinoderms,
but focused in the zone near Cumaná. Belmonte (1985) studied the asteroids,
echinoids and ophiuroids present in Margarita Island, while Amaro (2009) made a
revision of the echinoderms of the Gulf of Cariaco, and reported 50 species, nine
of which belonged to the Class Asteroidea, 17 Ophiuroidea, 12 Echinoidea and 12
Holothuroidea.
Perhaps the holothuroid has been the most highly studied echinoderm group of
Venezuela (Mago 1971; Martínez 1971, 1973, 1975, 1982, 1986, 1987a, b, 1988,
1989, 1991a, b; Martínez and Mago 1975), particularly due to their commercial
and ecologic interest. In what concerns to the remaining classes, Martínez (1969)
evaluated organisms belonging to the Family Brissidae (Class Echinoidea); while
Flores and Martínez (1970) described the status of the Family Goniasteridae (Class
Asteroidea) and Francisco and Pauls (2008) made a bibliographic compilation of
the Order Clypeasteroida (Class Echinoidea) reporting ten species, six of which
belong to the Family Clypeasteridae and four to the Family Mellitidae.
7.2.2 Ecology
Studies on populations and ecology of echinoderms have been made primarily in the
central-western and eastern coasts of the country. Most refer to abundance and
distribution of the species in the Class Echinoidea, Holothuroidea and Ophiuroidea.
With respect to the Class Echinoidea, six of the seven species known for the
Caribbean Sea have been reported locally (Hendler et al. 1995): Lytechinus
variegatus (Lamarck, 1816), Tripneustes ventricosus Lamarck, 1816, Diadema
antillarum Philippi, 1845, Arbacia punctulata (Lamarck, 1816), Echinometra
lucunter (Linnaeus, 1758) and Echinometra viridis Agassiz, 1863. These species are
distributed along the coast from Falcón to Sucre and Nueva Esparta states
(Cruz-Motta 2007). The sand dollar, Mellita quinquiesperforata (Leske, 1778) has
been reported from the coasts of Golfo Triste (Penchaszadeh and Molinet 1994).
Most records have been made between 0 and 50 m depth, especially on rocky
substrates and seagrass (Cruz-Motta 2007). For the Atlantic region, most species are
irregular urchins, among which are Agassizia excentrica Agassiz, 1869, Brissopsis
elongata Mortensen, 1907, Clypeaster euclastus Clark, 1941 and Schizaster orbi-
gnyanus A. Agassiz 1880 (Martínez 1969; Francisco and Pauls 2008).
Diadema antillarum is distributed along the entire continental coast and oceanic
islands of Venezuela, associated primarily with coral communities. The abundance
of this species in some coastal locations, before the massive mortality registered in
1984, was similar to those found in the rest of the Caribbean Sea. However, only in
the eastern zone of the country is there information at the population level before
and after the demise that this species underwent in the entire Caribbean Sea, which
also affected the populations present in Venezuela (Pauls 1982; Noriega et al.
2006). In Mochima Bay, eastern Sucre state, the average density before the
massive mortality varied between 0.28 and 4 ind m-2 (Pauls 1982). After the mass
mortality, the average density decreased to 0.15 ind m-2 in 2000 and 0.47 ind m2
in 2002 (Noriega et al. 2006). On the other hand, there have been records of
densities from 0.28 ind m-2 in 2002 to 1.05 ind m-2 in 2003 in Oricao Bay, and
from 0.84 ind m-2 (2002) to 0.74 ind m-2 (2003) in Chichiriviche de la Costa,
both locations in the central coast of Venezuela (Noriega et al. 2006). The pop-
ulations of D. antillarum in Morrocoy National Park and Cuare Wildlife Refuge
showed the highest average densities at Playuela (0.43 ind m-2) and Cayo Sur
(0.95 ind m-2), while the remaining areas showed densities \0.1 ind m-2. The
density found in Playuela in 2003 is lower than the one reported before the mass
mortality event (0.58–3.64 ind m-2) (Noriega et al. 2006). In Morrocoy National
Park, Weil et al. (1984) reported average densities between 2.6 and 3.7 ind m-2 on
three reefs (Pescadores, Playuela and Caiman). The highest densities (4.0–7.0 ind
m-2) were found in the shallow and frontal zones of the reefs, characterized by a
high structural complexity of the substrate. Low densities (1.5–4.0 ind m-2) were
found in deep zones of the reef platform with low to intermediate structural
complexity. Densities of D. antillarum in Los Roques Archipelago National Park,
specifically in Herradura Reef, varied between 0.22 and 0.23 ind m-2 (Noriega
et al. 2006). Currently, there are no data about the population densities of D. an-
tillarum in these or other areas of the country.
Other publications about the distribution and density of Venezuelan echinoids
have been mainly related with their association to seagrass, Thalassia testudinum.
Noriega et al. (2002) showed the relation between the density of L. variegatus and
several parameters of the seagrass (percent cover, density and length of the leaves,
density of the roots and total biomass) in Mochima National Park. Total biomass
was the most influential parameter on density (0.6–3.6 ind m-2) and distribution of
the species. At Margarita Island, Gómez (2002, 2003) also found a significant
correlation between the abundance of L. variegatus and the density of T. testudinum,
reporting maximum densities from 38 to 52 ind m-2.
The association of ophiuroids to specific habitats like sponges, corals,
octocorals and algae has been described by some authors (Boffi 1972; Hendler and
Miller 1984; Hendler and Littman 1986). Ophiuroids have been reported from the
central-western region of Venezuela to the Orinoco River delta, with 52 identified
species. The Family Amphiuridae is the most diversified one, being found both on
the benthos of shallow and deep water (Borges 2006), while Ophiodermatidae and
Ophiocomidae inhabit primarily shallow water. Villamizar and Laughlin (1991),
studied the animal communities associated to two species of dominant sponges
(Aplysina archeri and A. lacunosa), characterized by different degrees of structural
complexity, in a coral reef of Los Roques Archipelago National Park. Mollusks
had the largest number of species in both sponges (76.51 %); crustaceans had the
largest percent in number of individuals (67.27 %), while fish and ophiuroids had
the largest percent in dry weight (50.45 and 42.68 %, respectively).
In Cubagua Island, Nueva Esparta state, Hernández (2010) reported 24 species
of ophiuroids associated primarily with hard bottom, seagrass, aggregations of the
bivalve Arca zebra and the macroalga Halimeda opuntia. In the Gulf of Cariaco,
revision of the echinoderms resulted in 50 species, 34 genera, 22 families and four
classes: nine species of asteroids, 17 species of ophiuroids, 12 species of echinoids
and 12 species of holothuroids (Amaro 2009).
Studies have been made on the trophic ecology of species of Asteroidea,
Echinoidea and Holothuroidea. Studies on asteroids have focused mainly on the
composition of the diet and trophic interactions. In Golfo Triste, Falcón state,
Bitter (1984) found that Astropecten marginatus is a non-selective consumer,
although it shows a clear preference for mollusks, especially bivalves (33 %) and
gastropods (50 %). Similar results were found by Penchaszadeh (1973) with the
asteroid Astropecten brasiliensis, in which mollusks represented 80 % of the
stomach contents of the studied individuals. In relation to the trophic ecology of
Astropecten brasiliensis riensis and A. articulatus, Bitter and Penchaszadeh (1983)
found the trophic niches of both species overlap, although competition would be
reduced by specialization on the size of the preferred prey. In both species the diet
was mainly mollusks, with gastropods as the most important prey item. The
gastropod Finella dubia accounted for 50 % of the diet of A. brasiliensis riensis
and the gastropod Olivella myrmecoon was 26.5 % of the diet of A. articulatus.
Penchaszadeh and Lera (1983) found in their study of the diet of three species
of asteroids of the genus Luidia, that L. barimae is mainly a predator of ophiuroids
(85.8 % of its diet); while L. senegalensis feeds mainly on bivalves (49.1 %) and
L. clathrata on ophiuroids (35.3 %) and gastropods (24.4 %). Penchaszadeh and
Molinet (1983) evaluated the diet of the asteroid Tethyaster vestitus, and found
that it is a specialist predator of Astropecten spp. (which represented 55.8 % of the
total number of prey items). Other prey items of T. vestitus were L. clathrata and
ophiuroids (0.8 and 3.8 % of the diet, respectively). Bitter et al. (1980) studied the
trophic interactions between two asteroids (A. riensis and T. vestitus), and found
that echinoderms are the most important group in the diet of T. vestitus, with
asteroids representing 55.74 % of its diet and the remaining groups 44.20 %.
Among the latter, echinoids represented 12.30 %, crinoids 27.87 %, holothuroids
1.64 % and ophiuroids 2.4 % of its diet. The diet of A. brasiliensis riensis, was
composed of gastropods (67.8 %), pelecypods (20 %), crustaceans (4.71 %) and
scaphopods (2.23 %). A. riensis is an important element within the trophic
structure of the community studied, representing the second and third level of
consumers while T. vestitus represents the fourth level.
In Mochima Bay, Sucre state, the asteroid Oreaster reticulatus behaves mainly
as a predator of the sea urchin L. variegatus. However, some observations suggest
that it could be an opportunistic microphage, with densities between 0.0075 and
0.2 ind m-2 (Martín et al. 2001). The diet of A. marginatus at the mouth of the
Orinoco River and the Gulf of Paria was mainly bivalve mollusks and gastropods
(Ortega et al. 2011).
In the Caribbean Sea, many echinoids feed exclusively upon T. testudinum and
its epibionts (Lawrence et al. 2007). In La Tortuga Island, even though data on
density of T. ventricosus are not known, its stomach contents consisted primarily
of leaves of T. testudinum with epibionts although the macroalgae Acanthophora
spicifera, Hypnea cornuta and Cladophora spp. were found, showing their pos-
sible importance on the abundance and distribution of this species (Barrios and
Reyes 2009). These authors also indicate that the presence of calcareous algae like
Hydrolithon sp. and Jania sp. in the diet of T. ventricosus showed that they could
be important items in the calcium metabolism associated with formation of the
test. On the other hand, Montealegre and Gómez (1999) determined the stomach
contents of L. variegatus at Margarita Island, observing mainly remnants of
T. testudinum and smaller amounts of the macroalgae Ulva lactuca and diatoms.
With respect to holothuroids, Sambrano et al. (1990) characterized the ingested
items of Isostichopus badionotus and Holothuria mexicana from two types of
substrate (with calcareous and terrigenous origins) in T. testudinum meadows of
the western region of Venezuela. They found that the mean size of ingested species
differs significantly between these two holothurian species for both types of
substrate. The analyses of the stomach contents of Duasmodactyla seguroensis,
Euthyonacta solida, Ocnus pygmaeus, O. surinamensis, O. suspectus, Thyonella
sabanillaenis and Stolus cognatus from the northeastern region and some oceanic
islands, showed that the diet of holothuroids of the Order Dendrochirotida include:
planktonic and benthic diatoms, dinoflagellates and copepods as well as
filamentous cyanobacteria whereas species of the Order Aspidochirotida (I. bad-

ionotus, Holothuria grisea, H. occidentalis, H. pseudofossor and H. surinamensis)
consume detritus consisting of calcareous sand or quartz, coral fragments, bivalves
shells, test and spines fragments of echinoderms, sponge spicules, fish scales and
amorphous material. Holothuroids of the Order Apoda scrape the surrounding
ground with their feathered tentacles and consume mainly benthic diatoms and
remnants of green and brown algae (Martínez 1989, 1991a, b).
7.2.3 Reproduction
The biology and reproductive behavior of echinoderms in Venezuela is poorly

known. However, some studies suggest that temperature represents a key exoge-
nous factor in the onset of gametogenesis. In the northeastern region of Venezuela,
hydrographic and nutrient availability in coastal water masses are variable due to
the upwelling phenomena, which promote temperature fluctuations mostly during
the first half of the year (with variations between 20 and 28 °C) (Ferraz-Reyes
1987). Montealegre and Gómez (2005) indicate that L. variegatus exhibits the
highest spawning activity in June and July in coincidence with an increase of water
temperature. Espinoza et al. (2008), in spite of not finding a clear influence
of environmental factors on the reproduction processes of E. lucunter and
L. variegatus, indicated that low temperature (23–24 °C) seems to be an initiating
factor of gametogenesis and high temperature (26–28 °C) of maturation of
gametes of L. variegatus.
Muñoz (1980), Muñoz and Ellietes (1982, 1983) and Brey (1992), evaluated the
effect of different toxic substances (ferrous sulphate and sodium hypochlorite), as
well as temperature on embryonic development, fertilization and reproductive
processes in E. lucunter. Furthermore, Muñoz (1980) evaluated the fertilization of
eggs under variable thermal conditions (25, 30, 35 and 40 °C), finding a greater
motility of spermatozoa at 30–35 °C, a reduction in the percent of fertilization at
40 °C and a moderate reduction in the number of fertilized eggs at 25 °C. Brey
(1992) studied the effect of the ratio sperm/eggs and the time of post-dilution of
sperm in sea water on the fertilization success. Toxicity assays with Cr6+
(K2Cr2O7), Zn2+ (ZnCl2) and Cu2+ (CuSO4) found that zinc and chromium are less
toxic than copper (LD50 for copper between 36.1 and 119.3 ppb).
One aspect that is gaining importance on the eastern coast of Venezuela is
knowledge about larval and juvenile development of echinoid species with
potential for aquaculture. Some species in this category are L. variegatus (Butriago
and Lodeiros 2005; Butriago et al. 2005; Gómez and Gómez 2005; Domínguez
et al. 2007), T. ventricosus (Fernández 2005) and A. punctulata (García et al.
2005), in which the effect of salinity and food type were evaluated and are dis-
cussed in Sect. 7.5 of this chapter.
In holothurians, Palazón (2001, 2010) studied the reproductive biology of
I. badionotus from the islands of Margarita, Coche and Cubagua, in northeastern
Venezuela. He found that, in general, male:female sex ratio does not deviate
significantly from 1:1 and there were 2 % hermaphrodites. Sexual maturity was
reached at a horizontal diameter above 147 mm (55 g). Individuals with mature
gametes were found during the entire year, which suggests that reproduction is
continuous, although with a greater synchrony in spawning during the period from
September to November when water masses stratify and temperature is higher.
Fecundity is high, with an average number of eggs per individual of 1,417,330 (SD
151,186), but sometimes exceeded 5 million eggs.
7.2.4 Bioerosion and Herbivory
With respect to bioerosion by echinoderms, Weil (1980) evaluated the impact of

D. antillarum on coral reefs at Mochima National Park and Díaz et al. (1982) and
González (2005) evaluated it in Los Roques Archipelago. Losada et al. (1988)
manipulated densities of D. antillarum in the intake channel of Planta Centro
thermal electric plant, located at Punta Morón-Golfo Triste, and reported a sig-
nificant increment of bare substrate by grazing at densities that varied from
0.170 ind m-2 and bare space of 102.6 cm2 to more than 1,000 cm2 bare space
when the density reached 0.310 ind m-2.
Another aspect, though less known, is the effect of grazing by echinoids as
biocontrol agents. Lodeiros and García (2004) described for the first time in the
Gulf of Cariaco, Sucre state, L. variegatus and E. lucunter as control agents of
fouling in culture baskets and bivalve shells in suspended cultivation systems.
L. variegatus produced a 75 % decrease in the fouling of the culture baskets and
71 % that of the shells of Pinctada imbricata, suggesting the usefulness of this
method as a natural control of fouling in these systems.
7.2.5 Physiology and Biochemistry
There are few physiological studies of echinoderms in Venezuela. Studies with

holothurians have focused mainly on the ion-osmotic response to a great variety of
chemical compounds. The first studies were initiated in the 1970s by Zanders
(1970), Madrid (1971), Zanders and Herrera (1974) and Madrid et al. (1976), who
evaluated the effect of the salinity change of sea water upon the coelomic and
intracellular ionic composition of I. badionotus. Zanders (1970) studied the
histology, ionic composition and permeability to sodium of the intestine of
I. badionotus. The results showed that the organisms is isosmotic to sea water, but
levels of different ions in the tissues could differ from the latter, like potassium
which reaches intracellular concentrations 10–30 times higher than in the external
environment. Tissue water represents 80–90 % of the wet weight of the intestine
and 60–80 % of the tissue water corresponds to extracellular liquid. Zanders and
Herrera (1974) evaluated the distribution and ion flow in cells of the intestine and
stomach muscle of I. badionotus. They found that potassium concentration in
intestinal cells was 240 l-equiv g-1 cell and 252 l-equiv g-1 cell in those of the
stomach; sodium was 246 and 296 l-equiv g-1 cell, while chloride was 223 and
302 l-equiv g-1 cell, respectively. In muscle cells, potassium, sodium and chlo-
ride concentrations were 348, 119 and 127 l-equiv g-1 cell, respectively. Madrid
et al. (1976) found that intracellular ionic concentrations increase in direct relation
to the concentration of the external environment and the ionic concentrations of
the coelomic liquid of I. badionotus equilibrates with 80 and 120 % sea water
between 4 and 6 h.
Studies made by Herrera and Plaza (1981a, b), Jurisic (1982), Herrera and
López (1983), Jurisic et al. (1983), Sambrano et al. (1990), Conde et al. (1991),
Egea et al. (1991), Herrera (1991), Foglietta (1994), Foglietta and Herrera (1996),
Herrera and Foglietta (1998) and Herrera et al. (2000) focused their research on the
ion-osmotic response of the respiratory structures of I. badionotus and Holothuria
glaberrima. Herrera and Plaza (1981b), found that the respiratory tree and muscles
of H. glaberrima respire at a rate of 635.5 ± 10 l l O2/h 9 g dw and 137.4 ± 6.3
l l O2/h 9 g dw, respectively, in response to the concentration of sodium in
artificial sea water. They indicate that the substitution of sodium by chlorine,
lithium or Tris (pH 8.0) does not modify the oxygen consumption of the respi-
ratory trees structures. The low respiratory rate of muscles could be due to limi-
tations of oxygen diffusion in the tissue. Metabolic conditions could be involved in
the respiratory response of tissues to pH. Herrera and López (1983) evaluated the
effect of sodium concentration in the external medium and the intracellular
osmotic condition of the respiratory tree of H. glaberrima. They subjected the cells
of the respiratory trees to sodium concentrations reduced isosmotically (60, 30, 15,
6 and 0 % of normal sea water). The analyses showed an increasing loss of
sodium, potassium and chloride while maintaining the cytoplasm isosmotic with
the external medium. This indicates that the maintenance of isosmoticity with the
external medium assures the constancy of intracellular volume. The intracellular
concentration of potassium was regulated within relatively narrow limits in spite of
the changes in cell volume. In sea water with 6 and 30 % sodium, intracellular
sodium concentration changed only by 1.5 times which suggests a certain degree
of regulation. Herrera and Foglietta (1998) evaluated the effect of ouabain (which
blocks the Na–K pump) on the ion-osmotic properties of cells of the respiratory
trees of I. badionotus. Individuals were exposed to five concentrations of sea
water: 17.5, 21.0, 26.25, 35.0 and 42 % in the presence or absence of ouabain.
They found that control and ouabain-treated cells responded as perfect osmometers
to variations in sea water concentration implying that intracellular organic
osmolyte content also remained unchanged despite changes in sea water concen-
tration. In order to test if cells of marine stenohaline invertebrates can increase in
volume, Herrera et al. (2000) evaluated the effect of five different solutions of
artificial sea water with isosmotic sodium and KCl (100, 83, 71, 60 and 50 %),
with and without ouabain, in fragments of the respiratory tree of I. badionotus.
They found immobilization of extracellular sodium, depolarization of the
membrane, an osmometric increase of the cells and a decrease of potassium,

sodium and chloride when the cells were treated with the same dilutions of sea
water with distilled water. Jurisic et al. (1983) studied the role of chloride in the
intracellular isosmotic regulation in cells of the respiratory tree of H. glaberrima
exposed to different concentrations of chloride (100, 60 and 24 % of normal),
finding a small decrease in the intracellular potassium content. However, as a
consequence of the 50 % reduction in cell volume, the intracellular concentration
of potassium increased approximately by twofold, while that of sodium decreased
considerably. Respiratory rate of tissues increased with the reduction of external
chloride, which reveals an increase in the metabolic work needed for the sodium
and potassium pump against the ever increasing electrolytic gradient of these ions.
Foglietta (1994) studied the ion-osmotic response to changes in salinity of the
external medium in I. badionotus pre-incubated in 57, 100 and 137 % sea water
and the effect of ouabain in pre-incubated individuals at 100 % sea water. She
found an inverse linear relationship between the relative intracellular space to
incubation concentrations. These results indicate that cells show perfect osmo-
metric behavior. On the other hand, the intracellular concentration of K+ remained
unchanged, that of Na+ decreased with external salinity, and that of Cl- decreased
slightly when incubated above and below 83 % sea water and ouabaine.
Foglietta and Herrera (1996) performed studies similar to those of Foglietta
(1994), without the use of ouabain. They evaluated the ion-osmotic response of the
respiratory tree of I. badionotus pre-incubated in hyper-, iso- and hypo-osmotic sea
water. The cells and tissues of individuals pre-incubated in 20 and 48 % sea water
responded in a similar way to those pre-incubated in 35 % sea water. The intra-
cellular content of potassium was well regulated, remaining constant in spite of
changes in sea water concentration; that of sodium decreased with extracellular
concentration, while that of chloride increased in concentrations above and below
of 29 %. I. badionotus seemed to be more tolerant to hyposaline than to hyper-
saline sea water, which suggests that normal sea water represents the upper limit of
its salinity range. The osmotic condition of the cell and the response of the tissue
do not seem to be modified by the variation in the concentration of the means in
which it was pre-incubated.
In contrast to the studies just described, Conde et al. (1991) evaluated the
patterns of disintegration of fecal material of H. mexicana and I. badionotus. In
I. badionotus, the feces completely disintegrate around 8 h, suggesting a high
probability of dispersion in comparison with feces of H. mexicana which remain
on the substrate even after 25 h. The authors suggest that bacterial activity and
fungal growth are responsible for the disintegration of the fecal material of
I. badionotus, while the disintegration of the feces of H. mexicana is associated
with the activity of motile marine organisms.
There are few studies at the biochemical level. Urbaneja (1968) determined
steroidal and lipid contents in the sea star Echinaster sentus. Cedeño (1971)
identified the fatty acids of the holothurian, Ludwigothuria mexicana. Salazar et al.
(1994) made histochemical studies of cells of the echinoids, L. variegatus and
E. lucunter. Malavé (1995) characterized and quantified the lipid and fatty acid
content in the gonads of L. variegatus and E. lucunter. He found that the main
lipids present in both species were phospholipids and triglycerides, and in smaller
concentration free cholesterol, free fatty acids and ester of cholesterol. The higher
triglyceride and saturated fatty acid values present in the gonadic tissue of both
species coincided with their sexual maturity. The values of saturated fatty acids
were, in decreasing order of proportion, 16:0, 14:0, 17:0 and 18:0, while the
unsaturated fatty acids were represented by the mono-unsaturated fatty acids (16:1,
18:1 and 22:1) and the polyunsaturated fatty acids (20:5x3, 18:3 y 20:3). Díaz
et al. (1996) determined the total protein content and electrolytes in the hemal
lymph of L. variegatus and E. lucunter, finding that Na+ concentration varied from
466.42 to 470.30 meq l-1, Cl- from 372.78 to 739.2 meq l-1, and K+ from 5.73 to
3.25 meq l-1. Total protein concentration varied from 0.53 to 0.58 %.
The information in this section has been compiled from various bibliographic
sources which include studies made in Venezuela (Ummels 1963; Zoppi de Roa
1967; Martínez 1969, 1971, 1973, 1975, 1982, 1986, 1987a, b, 1988, 1989, 1991a,
b; Boada 1970; Flores and Martínez 1970; Mago 1971; Penchaszadeh 1973, 2003;
Martínez and Mago 1975; Bitter et al. 1980; Bitter and Penchaszadeh 1983;
Penchaszadeh and Lera 1983; Penchaszadeh and Molinet 1983, 1994; Bitter 1984;
Belmonte 1985; Noriega 1998; Francisco and Pauls 2008; Amaro 2009; Hernández
2010; Ortega et al. 2011) and in the Caribbean Sea (Jeannet 1928; Engel 1939;
Engel et al. 1960; Ummels 1963; Boffi 1972; Jangoux 1978; Hendler and Miller
1984; Hunte et al. 1986; Scheibling and Mladenov 1998; Donovan 2001;
Toral-Granda 2008) that provide information on echinoderm species found in
Venezuelan waters as well as the data base available in the web page of GBIF
(Biodiversity occurrence data provided by www.gbif.net, 2009-12-10). All echi-
noderm species from such studies have been placed in systematic order in the
appendix. The distribution of systematic groups as classes, families and species
was established considering the characterization of the Venezuelan ecoregions
(Miloslavich et al. 2005; Miloslavich and Klein 2008).
Two hundred and thirty four species of echinoderms have been reported for the
Venezuelan waters, with 97.87 % of them along the Caribbean Sea coast and only
2.13 % of them along the Atlantic coast (Table 7.1). For all coasts, the Asteroidea
is the most diverse with respect to the number of species (26.92 %), followed by
the Holothuroidea (22.65 %); while the Crinoidea is the least diverse (6.84 %). It
is important to emphasize, that in spite of a smaller diversity of species along the
Atlantic coast, Tethyaster vestitus (Say, 1825) (Asteroidea), Leptopentacta
deichmannae Domantay, 1953 (Holothuroidea), Paraster doederleini Chesher,
1972 (Echinoidea) and Amphiura otteri Ljungman, 1872 (Ophiuroidea) have only
been reported for this ecoregion.
Most records of echinoderms in Venezuelan waters have been made from

depths between 0 and 50 m (47.42 %), followed by the range 51–199 m
(26.12 %), 200–499 m (8.25 %) and depths [500 m (18.21 %). Now, considering
the distribution of echinoderms in the ecoregions described (Table 7.1, Fig. 7.2),
29.17 % of the known species are reported for the Eastern upwelling zone eco-
region; followed by the Oceanic islands ecoregión (23.33 %) and the Central coast
ecoregion (10.42 %). The smallest number was found in the Gulf of Venezuela
ecoregion (1.67 %). There are no reports of echinoderms within the Lake Mara-
caibo ecoregion, most probably due to the low salinity.
The Class Crinoidea includes 16 species, 15 genera, nine families and four
orders (Table 7.1), with members mainly in the Central Coast (23.91 %), Oceanic
Islands (21.74 %) and Eastern Upwelling ecoregions (19.57 %). They have not
been found in the Unare-Píritu, Golfete de Coro and Oceanic area ecoregions.
Comasteridae and Antedonidae showed a wider distribution, being present in
seven and five ecoregions, respectively. Families like Thalassometridae, Pentac-
rinidae and Bathycrinidae, have only been reported for a single ecoregion, Oceanic
Islands, Central Coast and Tocuyo, respectively. Forty percent of the crinoids
reported are found in the depth range 0–50 m, followed with 32 % in the range
51–199 m. The Charitometridae has the largest depth range, with representatives
from 51 m to more than 500 m deep. Tropiometridae, Colobometridae, Comas-
teridae and Antedonidae are only reported for shallow waters, in the depth range
0–50 m for the first, and the remaining in the range 0–199 m. Bathycrinidae and
Atelecrinidae show the deepest records ([500 m).
The Class Asteroidea is the most diverse, with 63 species, 40 genera, 15
families and six orders (Table 7.1), being mostly represented in the Eastern
upwelling (22.92 %), Oceanic Islands (22.92 %) and Paraguaná (11.81 %) eco-
regions; with the smallest representation in Unare-Píritu ecoregion (0.69 %, a
single record). The Astropectinidae and Luidiidae have the widest distribution
along the coast, with representatives in ten and nine ecoregions, respectively.
Families like Brisingidae, Asteriidae, Goniopectinidae and Benthopectinidae, are
only present in a single ecoregion, the first in Oceanic Islands ecoregion, the
second in Tocuyo ecoregion and the last two in the Eastern upwelling region. It has
been calculated that 30.53 % of the asteroid records has been made within
51–199 m, followed by depths greater than 500 m with 27.37 % of the records.
Members of Astropectinidae and Goniasteridae were present in all depth intervals.
Furthermore, the Oreasteridae, Asterinidae, Ophidiasteridae and Luidiidae, are
only reported for shallower waters, within 0–50 m for the first, and the remaining
from 0 to 199 m. Brisingidae, Asteriidae, Zorasteridae, Goniopectinidae and
Pterasteridae are only recorded at depths greater than 200 m.
The Class Ophiuroidea consisted of 52 species, 31 genera, 14 families and two
orders, being mostly represented in the Eastern Upwelling (34.07 %) and Oceanic
Islands ecoregions (21.98 %), and has not been found in the Cariaco Trench,
Tocuyo, Golfo de Venezuela and Oceanic area ecoregions. The Amphiuridae and
Ophiuridae have the widest distribution with representatives in seven ecoregions,
while the Asteroschematidae, Ophiochitonidae, Ophiolepididae, Ophiomixidae
Table 7.1 Number of echinoderm taxonomic groups in the marine ecoregions of Venezuela
Delta of the Orinoco River and Atlantic Front Orders 1 3 2 3 2 9
Families 1 4 4 4 2 15
Genera 1 7 6 6 2 22
Species 2 12 6 7 2 29
Eastern upwelling region Orders 3 5 2 10 3 23
Families 6 10 10 17 8 51
Genera 8 18 17 26 22 91
Species 9 33 31 33 35 141
7 Echinoderms from Venezuela
Axis Unare-Píritu-Tacarigua Orders 0 0 0 0 0 0

Families 0 0 0 0 0 0
Genera 0 0 0 0 0 0
Species 0 0 0 0 0 0
Central coast Orders 0 1 1 2 2 6
Families 0 1 4 2 3 10
Genera 0 1 6 2 4 13
Species 0 1 8 2 5 16
Golfo Triste Orders 3 2 3 8 0 16
Families 6 5 8 12 0 31
Genera 10 5 10 15 0 40
Species 11 8 14 17 0 50
Tocuyo region, Orders 2 3 1 6 1 13
Families 4 6 2 8 2 22
Genera 6 8 2 12 2 30
Species 6 16 2 15 2 41
Paraguaná region Orders 0 0 0 0 0 0
Genera 0 0 0 0 0 0
Species 0 0 0 0 0 0
249
(continued)
250

Golfete de Coro Orders 1 3 2 5 1 12
Families 1 5 5 6 1 18
Genera 2 10 8 10 1 31
Species 2 17 8 13 1 41
Estuarine region of lake Maracaibo Orders 0 1 1 1 0 3
Genera 0 2 2 2 0 6
Species 0 5 2 2 0 9
Golfo de Venezuela Orders 2 1 0 2 0 5
Genera 2 2 0 2 0 6
Species 2 4 0 2 0 8
Oceanic Islands Orders 0 0 0 0 0 0
Genera 0 0 0 0 0 0
Species 0 0 0 0 0 0
Oceanic area Orders 1 6 3 10 4 24
Families 6 11 9 18 8 52
Genera 9 30 13 25 15 92
Species 10 33 20 27 22 112
Cariaco Trench Orders 0 0 0 0 0 0
Genera 0 0 0 0 0 0
Species 0 0 0 0 0 0
Orders 4 6 2 10 4 26
Families 9 15 14 22 12 72
Genera 15 40 31 38 34 158
Total species 16 63 52 50 53 234
C. Lodeiros et al.
C Crinoidea, A Asteroidea, O Ophiuroidea, E Echinoidea, H Holothuroidea

Fig. 7.2 Sampling of sea urchins in the eastern upwelling ecoregion, in the Lobos island,
northern Araya peninsula Sucre state, Venezuela
and Ophionereididae are only reported for a single ecoregion. Records of ophiu-
roids show that 70.91 % of the species are found in shallow waters between 0 and
50 m deep, with the least number of reports at depths between 200 and 499 m
(3.64 %). Most of the reported families of ophiuroids in Venezuela are only found
at depths between 0 and 50 m (five families), while the Ophiactidae, Ophiole-
pididae and Ophiuridae are reported at depths greater than 500 m.
The Class Echinoidea is represented by 50 species, 38 genera, 22 families and
10 orders, with most members in the Eastern Upwelling (26.19 %), Oceanic
Islands (21.43 %) and Central Coast (13.49 %) ecoregions, and no records in the
Oceanic area region. The Clypeasteridae, Brissidae and Schizasteridae showed a
wider distribution along the coast, with representatives in eleven, eight and seven
ecoregions, respectively. Six families (Aeropsidae, Cassidulidae, Echinoneidae,
Echinidae, Hemiasteridae and Loveniidae) are reported for a single ecoregion, the
Oceanic Islands ecoregion for the first three and the remaining in the Eastern
Upwelling ecoregion. Most echinoids (45.76 %) in Venezuela occupy shallow
waters, between 0 and 50 m deep, followed by the range 51–199 m (42.37 %).
The echinoid families reported in this study are only present at most in two depth
ranges, most from 0 to 199 m (eight families). Only the Echinothuriidae was found
in two depth ranges, 200–500 m and beyond 500 m. The other families (13) were
only present in one depth range. The Echinometridae, Echinoneidae, Mellitidae
and Scutellidae, are reported between 0 and 50 m deep, while the Aeropsidae,
Aspidodiadematidae, Hemiasteridae, Loveniidae and Saleniidae were only found
in depths greater than 500 m.
The Class Holothuroidea consisted of 53 species, 34 genera, 12 families and
four orders, with most members in the Eastern upwelling (46.58 %) and Oceanic
Islands (30.14 %) ecoregions. Members of this class have not been found in the
Cariaco Trench, Central Coast, Tocuyo, Golfete de Coro and Golfo de Venezuela
ecoregions. The Family Holothuridae shows the widest distribution along the coast
(being found in five ecoregions), while the Chiridotidae, Deimatidae, Molpadiidae,
Psychropotidae and Psolidae were only present in a single one. Most records of
holothurians (69.64 %) are from the depth range 0–50 m. There are no records
from the range 200–499 m. Eight families of holothurians are only found in
shallow waters down to a maximum depth of 199 m. Representatives of four
families (Deimatidae, Molpadiidae, Psychropotidae and Synallactidae) have only
been found at depths deeper than 500 m.
7.4 Paleontology
Paleontological studies of echinoderms in Venezuela were initially developed in

the 1920s, as a supplementary component during the initiation of petroleum
exploration. The lack of knowledge at that time about the Venezuelan sedimentary
basins prevented the stratigraphic location of areas with fossil accumulations as
they are known today (Fig. 7.3). In addition, the limited attention to echinoderms,
being considered among the so called miscellaneous groups due to their apparent
lack of importance as paleo-stratigraphic markers, decreased the interest for
research on this group of fossils.
Early geological expeditions to Venezuela by P. Christ, C. Wiedenmayer,
Vonderschmitt and H.G. Kugler between 1921 and 1926 provided some specimens
for this study. Also, J. Nance in 1941 and D. Rusk in 1954, among several other
anonymous contributors, added specimens to the invertebrate survey from Venezuela.
Later taxonomic studies, based upon the first collections, led Jeannet (1928),
Cooke (1941, 1961), Anisgard (1954), Von der Osten (1957), Weisbord (1969),
Hoover (1981) and Mooi and Peterson (2000) to the formal classification of the
first fossil species of Venezuelan echinoderms. Mihaljević et al. (2010) placed the
study of fossil echinoderms in the paleoecological and paleobiogeographical
context, which emphasizes the importance of the study of fossil echinoderms for
the paleoenvironmental interpretation.
It should be emphasized, that the data provided in geological order in the
following section of paleoenvironmental descriptions are based on the Strati-
graphic Lexicon of Venezuela of the Venezuelan Ministry of Energy and Mines
(Ministerio de Energía y Minas 1997). This is also referred to in the web site
(http://www.pdv.com/lexico) as Venezuelan Geologic Code, and has been devel-
oped from Venezuelan fossil echinoderms references and records based on our
own field in situ collections (Fig. 7.4) and specimens from museum collections.
7.4.1 Permian
The earliest record of echinoderms in Venezuela is a species of undetermined

Cidaroida collected in the Palmarito Formation of Zulia state (Hoover 1981). This
corresponds to a paleoenvironment of transitional sedimentation between shallow
waters of variable salinity, and deeper waters with constant marine salinity.
Fig. 7.3 Geological map of Venezuela showing the fossil echinoid localities. 1 San Cristóbal, 2
Mérida, 3 Maracaibo, 4 Los Monjes, 5 Trujillo, 6 Barinas, 7 Guanare, 8 Barquisimeto, 9 Coro, 10
San Felipe, 11 San Carlos, 12 Valencia, 13 Maracay, 14 San Juan de Los Morros, 15 Los Teques,
16 Caracas, 17 Península de Paraguaná, 18 Las Aves, 19 Los Roques, 20 La Orchila, 21 La
Blanquilla, 22 Los Hermanos, 23 Los Testigos, 24 Porlamar, 25 Margarita, 26 Península de
Araya, 27 Cumaná, 28 La Tortuga, 29 Barcelona, 30 Maturín, 31 Tucupita, 32 Ciudad Bolívar, 33
San Fernando de Apure, 34 Puerto Ayacucho
7.4.2 Lower Cretaceous
The reports of Cooke (1961) show the presence of Tylocidaris sp. and Stereo-
cidaris sp. in the Barranquin Formation (Hauteriviense-Barremiense) of north-
eastern Venezuela (Sucre state), an area with a shallow marine paleoenvironment,
and Tetragramma sp. in rocks without formational information. For the Chimana
Formation (Albiense), also in eastern Venezuela (Anzoátegui state). Cooke (1961)
reported Pseudananchys sp. from a paleoenvironment characterized by external
Fig. 7.4 Echinoderm fossils of Venezuela from Falcón state: a Moira atropos from the San
Gregorio Formation, b Lytechinus cf. euerces, c Clypeaster subdepressus from La Vela
Formation and d Melitella falconensis from San Gregorio Formation
platform and continental slope. This author also described Phyllobrissus zulianus
and Heteraster (Washitaster) bravoensis from the Apon Formation (Aptiense-
Early Albiense) in western Venezuela (Zulia State), characterized by marine
environments of combined facies of littoral with sand bars, open platform and
restricted platform. Vivas (1987) reported Holectypus (Coenholectypus) planatus
aponensis. Samples of Enallaster cf. texanum from El Cantil Formation, in
Monagas state (Aptiense-Albiense) and from Capacho Formation (Albiense-
Turoniense) in Zulia state, come from an area with uniform sedimentary condi-
tions in a paleoenvironment of internal to medium platform, where water depths
should have been between 10 and 70 m. Cooke (1961) found Hemiaster sp. and
Epiaster whitei in a location without formational information in Río Caripe,
Monagas state.
7.4.3 Upper Eocene
In the Misoa Formation (a paleoenvironment with deltaic to coastal marine

characteristics) at Trujillo state, Eupatagus sp. was found, whereas Eupatagus
clevei was recorded by Cooke (1961) from rocks without formational information
from Lara state. Oligopygus rotundus, was collected from rocks without forma-
tional information from Guárico State (Cooke 1961), and Oligopygus christi and
Pauropygus sp. were found in rocks without formational information from Barinas
State (Mihaljević et al. 2010). The latter authors describe Oligopygus nancei, from
Caratas Formation (Tinajitas Member) from a reef environment with shallow
water in Monagas state.
7.4.4 Oligocene
Weisbord (1969) found Eucidaris tribuloides in rocks without formational infor-

mation from Nueva Esparta state. The stratigraphic record of this species extends
until the Recent.
7.4.5 Lower Miocene
Jeannet (1928) described Brissoma vonderschmitti, Opissaster kugleri and Opis-

saster sp., which were collected in the upper series of Agua Salada Formation, in
Falcón state. It is an hemipelagic deposit with paleoenvironmental variations
between a deltaic complex and deep waters.
7.4.6 Middle Miocene
All species found for this period come from formations and places in Falcón state
(Jeannet 1928; Cooke 1961). Jeannet (1928) reported Clypeaster sp. and
Pericosmus sp. in rocks of the Capadare region; Clypeaster kugleri, Echinolampas
sp. and Heteroclypeus wiedenmayeri from rocks without formational information
in the vicinity of Caño León, probably La Vela Formation. This should have been
a sedimentary environment in shallow waters, with salinity below normal sea
water and restricted communication with the open ocean. Mihaljević et al. (2010)
reported that Encope michelini, Encope wiedenmayeri, Encope vonderschmitti and
Encope spp. were found in rocks without formational information in Boca de
Hueque, probably of Punta Gavilán Formation from Pliocene, characterized by the
sedimentation in a marine environment, with depths higher than 100 m within an
external sublittoral zone. These authors also reported Eurhodia falconensis from
rocks without formational information of La Guarava-Santa Clara (probably of
Punta Gavilán Formation); Echinolampas lycopersicus in rocks without forma-
tional information from El Bejuco, in Valle de Araurima; and undetermined
species of Encope spp., from rocks without formational information from La
Caridad and Mirimire. Finally, Plagiobrissus lamberti was described in rocks
without formational information from Guaidima to Riecito.
7.4.7 Upper Miocene
Mihaljević et al. (2010) found Agassizia clevei, Antillaster lamberti, Brissopsis

antillarum, Clypeaster subdepressus, Lytechinus cf. euerces, and Prionocidaris sp.
in La Vela Formation of Falcón state. These authors also found Clypeaster
rosaceus in rocks of Guajira peninsula, probably associated to Castillete Formation
of Zulia state, from deposits formed in environments of brackish to shallow marine
waters, and in Margarita Island (probably from the Cubagua Formation, a member
of Las Tejitas, at Nueva Esparta state). Cooke (1961) and Jeannet (1928) reported
A. lamberti from Ojo de Agua Formation of Falcón state, a paleoenvironment of
deposits in brackish to shallow marine waters, not far from the coast.
7.4.8 Pliocene
Mihaljević et al. (2010) reported Lovenia sp. Rhynobrissus rostratus, Schizaster

eurynotus, Tripneustes tobleri, Brissopsis antillarum, and Cassidulus (Cassidulus)
falconensis from Punta Gavilán Formation of Falcón state. Cooke (1961) found
Encope emarginata, E. secoensis and Moira atropos, in San Gregorio Formation
of Falcón state, a place characterized for having been a shallow littoral marine
environment, with ostreid reefs and small waves. This author also reported
Agassizia excentrica, A. scrobiculata, arbacia punctulata, Encope kugleri, Lite-
chinus variegatus and Melitella falconensis from San Gregorio Formation in
Falcón state. Finally, this author indicated that Pericosmus stehlini, and Plagio-
brissus grandis, occur in Paraguaná Formation in Falcón state, a place which
similarly shows a littoral marine paleoenvironment with shallow waters.
7.4.9 Pleistocene
Weisbord (1969) recorded Brissopsis cf. atlantica, Echinolampas sp., Echinometra

lucunter, Encope emarginata Leske, 1778, E. secoensis Cooke, 1961, Leodia
divinata and M. atropos, from the Cabo Blanco Group of Vargas state, but the
author did not define the formational units. The Cabo Blanco Group with marine
fossils is made up by Playa Grande and Mare Formations. The paleoenvironment
of both corresponds to a littoral shallow water marine one, with coastal lagoons
and moderate surf zone. Clypeaster ravenelii, Meoma ventricosa and Encope
kugleri, were recorded for the Cumaná Formation by Macsotay et al. (2009), and
its paleoenvironment corresponds to a tropical marine one with quiet and shallow
waters with reefs.
7.4.10 Recent
Agassizia excentrica, A. punctulata, Clypeaster rosaceus, C. subdepressus, Encope

emarginata, E. secoensis, Lytechinus cf. euerces, Melitella falconensis, Moira
atropos and Prionocidaris sp., have been registered as fossils in Venezuela
(Mihaljević et al. 2010).
The geochronological descriptions of the echinoderm species, as expressed by
Mihaljević et al. (2010), show that the fossil echinoderms of Venezuela were most
diverse during Middle Miocene, in contrast to the rest of the Caribbean Sea where
they were most diverse during Eocene (Kier 1977; McKinney et al. 1992; Donovan
2001). However, it should be noted that the oceanographic and environmental
changes that took place in the Caribbean Sea, as a result of the elevation of the
Panama isthmus and the separation of the Pacific and Atlantic oceans during
Pliocene, resulted in the extinction and renovation of different invertebrate faunas

(Jackson et al. 1993; Cheetham and Jackson 1996; O’Dea et al. 2007; Johnson
et al. 2009; Aguilera et al. 2010).
Paleontological records are lacking for the upper Cretaceous, Paleocene and
lower and middle Eocene. This bias could probably be associated to sampling
problems or underdevelopment of taxonomic work in the units of these eras which
have fragmentary remnants of echinoderms without a posterior classification (e.g.
Navay Formation of upper Cretaceous: Sánchez and Lorente 1977; Morán For-
mation of Paleocene: Bushman 1965; Misoa Formation of lower and medium
Eocene: Miller and Collinson 1951).
There is a clear preferential bathymetric distribution of recent echinoderms in
the western tropical Atlantic and Caribbean Sea (Lessios 2005; Francisco and
Pauls 2008). Families like Echinometridae, Clypeasteridae, Mellitidae and
Toxopneustidae are found with greater frequency in littoral zones of shallow
waters down to a depth of 402 m. This is in extreme contrast to families like
Echinothuriidae, Aspidodiadematidae, Pedinidae, Hemiasteridae, Pourtalesiidae,
Urechinidae, Aeropsidae and Lovenidae whose preferential depth is between 740
and 5,200 m. Amongst these extremes, the families Schizasteridae, Cidaridae and
Diadematidae are distributed with some variation on the platform and continental
slope. This fact shows the usefulness of echinoderms as indicators of paleoenvi-
ronments, as suggested by Kroh and Nebelsick (2003) and Mihaljević et al. (2010).
This agrees with the environment of sedimentation of the formations which hold
the co-generic taxa shown in the results described before. In the case of extinct
species whose habitat is unknown, it is necessary to use the faunistic associations
and lithology of the sediment types to support any paleobathymetric hypotheses.
7.5 Extraction, Fishing and Aquaculture
Fisheries of echinoderms are centered on sea urchin and sea cucumbers. They
represent a product of high cost, since sea urchin gonads and the body wall of sea
cucumbers (bêche-de-mer) are considered delicacies and aphrodisiacs.
Consumption of sea urchin gonads is primarily centered in Europe and Asia.
Consumption of sea cucumber body walls is primarily found in Asia and the
southwest Pacific.
It is currently considered that most sea urchin and sea cucumber fisheries have
reached their maximum exploitation rates. In contrast, their aquaculture is in
expansion, particularly that of the sea cucumber Apostichopus japonicus. FAO
statistics (http://www.fao.org/corp/statistics/en/) show that echinoderm extraction
has decreased from 140,000 t at the end of 1990s to 90,000 t in 2007, mainly due to
a reduction of catches because of overexploitation of natural stocks. However,
their cultivation, practically nonexistent in the twentieth Century, started in 2003
with a production of 37,000 t and reached 85,000 t in 2007, for a value of US$ 300
Fig. 7.5 Sea urchin cone:

test with several gonads of
the sea urchin Lytechinus
variegatus prepared for
consumption in Margarita
island, Venezuela
million. Thus, the demand of the Asiatic countries is supplied both by aquaculture
and by importation of sea cucumbers fished in other regions.
7.5.1 Extraction and Fishing
The consumption of sea urchins in Venezuela is known since prehispanic times

(Pauls 1992). However, sea urchins in general are not highly appreciated in the
Venezuelan diet. There is only an artisanal exploitation of L. variegatus in Mar-
garita Island, Nueva Esparta state, without an official control for the exploitation of
this resource. Local people gather the gonads of several sea urchins and place them
into one of the tests, giving it a conic form (Fig. 7.5). The cones are roasted and
sold later informally for approximately one US$ each (Gómez 1999). In other
zones of Venezuela, like the surroundings of Mochima National Park, this species
is abundant and could be commercially exploited under the administration of the
National Institute of Fisheries and Aquaculture (INSOPESCA) and the Office of
National Parks. However, until now, the product is neither commercialized nor
exploited. Litechinus varieatus generally inhabits beds of the seagrass, T. testud-
inum. An uncontrolled exploitation could have negative effects on the marine
ecosystem.
There is no exploitation of the sea urchin E. lucunter, although it is occasionally
consumed by tourists and some local people. Since E. lucunter is associated with
rocky and coralline substrate of shallow waters where it can reach a relatively high
abundance, it could also be threatened by an uncontrolled exploitation. However,
there are not studies that suggest how its extraction and administration could be
regulated.
The sea urchin T. ventricosus has been fished in the Caribbean Sea, particularly
in Barbados, where its natural banks were overexploited decades ago (Scheibling
and Mladenov 1998). This species is also found in Venezuelan waters, more
associated with zones beyond the continental coastline. Its abundance is so low
that not even a subsistence fishery is possible.
The exploitation of sea cucumbers in Venezuela is centered on two species,
H. mexicana and, particularly, I. badionotus, and has been generally illegal.
Nevertheless, under pressure from dealers and Asiatic fishers for establishing a sea
cucumber fishery in Venezuela, by middle 1990s an experimental permit was
issued by Servicio Autónomo de los Recursos Pesqueros y Acuícolas (SARPA,
currently INSOPESCA) of Venezuela. The permit was intended to be for one-year.
It established a maximum weekly quota of 200 kg per vessel, as well as a
requirement for continuous supervision and evaluation of the resource (Conde
1997; Rodríguez and Pauls 1998). However, the conditions of this permit were not
properly fulfilled, since the information on the extracted amounts was considered
neither adequate nor trustable. This led to the suspension of the permit after 6
months, but the illegal exploitation of sea cucumbers has continued. Acccording to
Rodríguez and Pauls (1998), had the exploitation of holothurians been continued
in that form, the damage caused to these resources along the eastern coasts (where
fishing was permitted) could have produced major effects on the populations.
These authors, using a report from Marcano and Marcano (1993) about the average
dry weight ind-1, showed that landings could have surpassed 42,000 kg y-1,
equivalent to more than 3 million individuals.
Some results from this exploitation are provided by León and Cabrera (1994).
They found abundance indices of ca. 4 ind m-2 in the vicinity of Coche Island and
2 ind m-2 near Cubagua Island, which are considered high compared to other
zones in the Caribbean Sea (González et al. 2005). According to León and Cabrera
(1994), both species of sea cucumbers were distributed in depths from 2 to 40 m,
but their exploitation was concentrated between 10 and 15 m due to restrictions of
the breathing equipment used for the fishing process. They also found that
I. badionotus reaches sexual maturity at a length above 180 mm, although Palazón
(2001) reported sexual maturity at smaller size (body length of 147 mm).
The Ministry of Agriculture and Livestock established a general closed season
for sea cucumbers in Venezuelan waters until the needed technical studies for the
evaluation of the resources are made (Gaceta Oficial de la República de Venezuela
14-07-1997, MAC-DM 195, 10-07-1997). Nevertheless, illegal fishing is active
and there have been reports of Venezuelan sea cucumbers exported to Japan in
2005 (Toral-Granda 2008). In spite of the importance and uniqueness of the reports
by León and Cabrera (1994) and González et al. (2005) pertaining to the evalu-
ation of the sea cucumber resources of Venezuela, it is difficult to reach adequate
conclusions from them that would allow administration strategies for these
resources. Hence, there is need to generate new knowledge to establish an
adequate legal framework for the protection and exploitation of sea cucumbers,
Fig. 7.6 Major stages of larvae of Lytechinus variegatus: prisma larva, with 2 arms, with 4 arms,
with 8 arms and competent larva. The bar in each photo represents 10 lm at 100x magnification
(Butriago and Lodeiros 2005)
supported by technical evaluations in different regions of the country. The pres-

ence of a continuous illegal fishing of sea cucumbers in the Venezuelan coasts, as
revealed from sources in the coastal communities, impoundments made by
authorities and our own ‘‘in situ’’ observations, make it urgent to establish an
evaluation of these resources along the Venezuelan coast. In this regard, consid-
ering that previous studies were limited to a depth of 15 m, future studies should
cover explorations to deeper areas.
7.5.2 Aquaculture
The sea urchins (L. variegatus, E. lucunter, T. ventricosus and A. punctulata) have
been studied with the purpose of increasing production by aquaculture (Lawrence
and Balzhin 1998; García et al. 2005; Astudillo et al. 2005; Barrios and Reyes
2009). Furthermore, there is also interest in Venezuela for using the eggs and
larvae in sensitivity studies of toxic and contaminant compounds (Esclapés 1999).
This supports the importance of establishing techniques to induce spawning in
these species and production of larvae.
Most attention for aquaculture has been given to L. variegatus. Studies have
investigated mass production of larvae (Fig. 7.6), postlarvae and juveniles (Bu-
triago and Lodeiros 2005; Butriago et al. 2005) with the goal to establish initial
larval density and the microalgae diet appropriate for cultivation, as well as to
determine the conditions leading to metamorphosis and to estimate the postlarval
growth rate. These experiments showed that it was possible to mass produce
competent larvae with a length of 650 lm after 12–13 days with densities of
0.25–1 larva ml-1. The microalga Rhodomonas sp. was a convenient diet for
optimal larval growth and survival. Metamorphosis can be induced with biofilms
containing benthic diatoms (Navicula sp. and Amphora sp.) incrusted on plastic
layers obtained from sliced disposable beverage bottles (Fig. 7.7), reaching 100 %
96 h after the appearance of competent larvae. Nevertheless, a diet on benthic
microalgae is not sufficient for a good larval development, hence a supplementary
food source is necessary. For this purpose, the macroalga, U. lactuca, supports a
considerable growth. However, supplementary addition of feed pellets for marine
shrimp to the diet, increases survival (85–86 %). These results suggest studies
Fig. 7.7 Sliced plastic

bottles with biofilms of
benthic microalgae to induce
the larval metamorphosis of
Lytechinus variegatus
(Butriago and Lodeiros 2005)
should continue to optimize the initial stages of culture, as well as for the
following growth stages. Nutritional research should be emphasized to establish
the appropriate diet for growth and the production of gonads.
Studies in Venezuela concerning culture of E. lucunter, A. punctulata and
T. ventricosus have focused on conditions affecting larval development. García
et al. (2005) studied the effect of salinity (30 and 40 %) and type of microalga on
the larval development of A. punctulata, and suggested the use of the microalgae
Isochrysis galbana, Chaetoceros gracilis and Chaetoceros calcitrans for raising
larvae. They also found that using 40 % salinity reached the greatest survival,
while in 30 % salinity the growth rate was maximized. Astudillo et al. (2005)
concluded that E. lucunter is a species with excellent characteristics for cultivation
because it has short embryonic (16 h) and larval (12 days) periods and good
survival rates when fed a monoculture of C. gracilis. This microalga also supports
larval development of T. ventricosus, which completes larval development to
metamorphosis in 24 days after fertilization at temperatures from 23 to 29°C
(Fernández 2005). Survival is greater at lower temperatures.
Studies to increase production of sea cucumbers by culture have not been done
in Venezuela despite the high product market value and the detritivorous condition
that make their culture attractive.
7.6 Threats to Echinoderms in Venezuela
The echinoderms in Venezuela are not free from threats experienced by marine
organisms nowadays, like global warming (Byrne et al. 2009), increased UV
radiation associated with a reduction in the ozone layer (Lesser and Barry 2003)
and acidification of the oceans produced by an increase in the CO2 concentration
(Kurihara and Shirayama 2004).
At a local level, the different threats on echinoderm biodiversity can be divided

according to their origin, as natural threats and those generated by human inter-
vention. Considering the natural threats, during El Niño years most parts of
Venezuela showed a delay in the onset of the rainy season and a reduction in the
amount of precipitation (Lozada 2002). This can produce an increase in salinity of
coastal waters in some zones. However, the direct effect of El Niño-Southern
Oscillation phenomena upon echinoderm populations has not been described.
Other threats, like hurricanes, are relatively infrequent in Venezuela in comparison
with the rest of the Caribbean Sea due to the geographical position of the country
(Díaz-Martín 1995). There are no current records of their direct effects on popu-
lations of echinoderms. From a geological point of view, Venezuela is located
above a major fault zone, which produces relatively frequent earthquakes. In spite
of this, ‘‘tsunamis’’ have been rare (Grasses 1994). However, identification of a
series of submarine displacements in the Lesser Antilles reveal the vulnerability of
the Venezuelan coasts to this type of phenomenon (Theilen-Willige 2006).
It has been estimated that the most important potential natural threat to echi-
noderms in Venezuela is the large load of sediments carried by rivers during the
rainy season (May–November). This phenomenon can generate mass mortalities of
marine organisms (Goodbody 1961) and causes losses in the diversity of echi-
noderms, particularly in coral reefs (Vázquez-Domínguez 2003). In this sense, it is
important to emphasize the influence of the Orinoco River on the patterns of
salinity, currents, nutrient concentration and suspended material in the Atlantic
coast of the country and the eastern Caribbean Sea whose fluctuations are directly
related to the dry-rainy season regime (Penchaszadeh et al. 2000).
Regarding threats by diseases, there are reports that epidemics and the number
of emergent diseases are rising worldwide in corals and other marine organisms
(Richardson et al. 1998; Harvell et al. 1999; Green and Bruckner 2000). In
Venezuela, the echinoid D. antillarum did not escape from the mass mortality
registered in the general Caribbean Sea between 1983 and 1984 (Lessios et al.
1984; Hunte et al. 1986; Lessios 1988). The abundances of D. antillarum recorded
between 2002 and 2003 in different localities of the country show that they have
not reached yet the population density levels reported before the mortality period
of 1984 (Noriega et al. 2006), although our own observations in Mochima Bay and
the Gulf of Cariaco show a high density of individuals with signals of recovery of
the populations.
In addition to disease, an 80 % reduction in the coralline cover has been
reported in the Caribbean Sea (Gardner et al. 2003). In Venezuela coral cover has
decreased at Sombrero Key in Morrocoy National Park (Cróquer and Bone 2003).
This could also compromise the abundance of echinoderms like the echinoid
E. lucunter, generally associated with the terrace and the reef crest (Abbott et al.
1974) and D. antillarum which also occur in these types of environments.
Regarding anthropogenic threats, it is important to show that intense petroleum
exploitation and processing in Venezuela, with a heavy traffic of vessels and a
network of oil pipes, the chances for spills of oil and other derivatives represent a
latent and constant threat. This is particularly serious during the phase of
transportation towards storage places due to the eventual metal fatigue and rupture
of pipes, pumps and tanks (López et al. 1999). Populations of echnoids like
E. lucunter could be reduced by 80 % after an oil spill in the coasts of Panamá
(Jackson et al. 1989). Furthermore, oil spills (crude or refined) significantly affect
the early larval stages of echinoderms (Jangoux 1984). These effects vary
according to the composition of the hydrocarbon (Allen 1971).
More than 50 % of the human population in Venezuela is concentrated in or
near the coasts, increasing sedimentation considerably because of infrastructure
developments as well as the pollution by untreated sewage (Díaz-Martín 1995).
Other contaminants, sub-products of the industry and agriculture, can also become
a threat to the fauna in coastal marine environments of Venezuela (Díaz-Martín
1995; Penchaszadeh et al. 2000). Unfortunately, Venezuela does not have effective
and widespread sewage treatments and most contaminants reach the sea without
previous processing. This can produce high oxygen demand, eutrophication,
accumulation of heavy metals and other processes (Penchaszadeh et al. 2000). In
this sense, concentrations of some heavy metals, in particular mercury, have been
measured in asteroids like Luidia clathrata, L. bareimaes, L. senegalensis and
Tethyaster vestitus in the zone of Golfo Triste. Concentrations ranged between 3.8
and 8.87 lg g dry weight-1 in the gonads (Iglesias and Penchaszadeh 1983).
Concentrations of the metals Al, Cu, Mn, Ni, Pb, and Zn were measured in the
eviscerated digestive tissue of two species of holothurians (I. badionotus and
H. mexicana) in Morrocoy National Park. Concentrations of Cu, Ni, Pb and Zn
were higher than in the surrounding sediments (Laboy-Nieves and Conde 2001).
Industrial trawl fishing, recently prohibited by the Venezuelan government
(actually in March 2009), was considered a threat to echinoderms. However,
echinoderms represented a minor portion (\1%) of the incidental capture in the
trawls in eastern Venezuela (Marcano et al. 2001), although their representation in
the catches from trawl fisheries has been little studied in other parts of the world
(Alverson et al. 1994; Kelleher 2005). Unpublished observations from one of the
authors (JA) indicate echinoderm discards by industrial trawlers in eastern
Venezuela included echinoids, asteroids, ophiuroids and, in a lesser proportion,
holothuroids and crinoids. Most of them, with the exception of ophiuroids due to
their fragile structure, were returned alive to the ocean. Hill and Wasemberg
(1990) examined the fate of discards from shrimp trawlers in Torres Strait,
northern Australia, and recorded an 88 % survival rate in the asteroid, Astropecten
sp. However, the trawl gear can modify the habitat and generate a reduction in the
density of populations of echinoderms and other groups of benthic animals
(Bergman and Hup 1992). In this sense, the interdiction of industrial trawling in
Venezuelan waters could have reduced the threat to a considerable number of
organisms whose capture was intended or not for consumption. It could also have
prevented a possible suspension of contaminants present in the sediment, like
DDT, PCBs and PAHs, reducing their incorporation into the trophic chains
through the plankton (Schiff 2000). The interdiction only affected industrial
trawling, because artisanal trawling is still actively practiced in shallow areas of
eastern Venezuela. For example, trawl fishing of the turkey wing, A. zebra
Fig. 7.8 Samples obtained from a bivalve mollusk dredge with a large number of echinoderms,
Araya Peninsula, northeastern Venezuela
(Mollusca, Bivalvia, Acidae), which recently generated more than 50,000 t y-1,
has an important component of echinoderms in the incidental catch, which is
extracted and discarded maimed or dead, or their habitat destroyed by the low
selective fishing gear (Fig. 7.8).
Unfortunately, estimating the echinoderm discards in fisheries has been diffi-
cult. In this sense, Kelleher (2005) mentions that both fishers and observers tend to
focus on commercial species and recognized animals. There is a tendency to group
tunicates, sponges, echinoderms, hermit crabs, worms and corals with jellyfish and
perceive such biomass as debris, rather than as organic material. These non-
commercial animals are frequently ignored and not recorded as discards during
studies. This biomass tends to be omitted from estimates of discards. Even in
countries that prohibit discards, allowance is made for discarding of non-com-
mercial species (such as in Iceland). Many of these animals also pose practical
problems of measurement of the biomass but may constitute significant large
proportion of the total biomass harvested by trawls (Prena et al. 1999).
Considering the direct fishing of echinoderms, there exist evidence of
consumption of sea urchins in the Venezuelan territory since prehispanic times
(Pauls 1992), although this activity is currently restricted to artisanal fishers of
L. variegatus in Margarita Island (Gómez 1999, 2002) and has reduced local
populations (L. León, Universidad de Oriente, pers. comm.). This situation could
be worse in the future because the fishing of L. variegatus is still unregulated.
There is an interdiction of fisheries of sea cucumbers, which has apparently
reduced this activity. However, illegal fishing continues, mainly of I. badionotus,
and there are reports of large impoundments within Mochima National Park
(Rodríguez and Pauls 1998). Unfortunately, there are no current official figures
about the amount or species of sea cucumbers illegally fished.
Even though there is no strong tradition for consumption of sea urchins or sea
cucumbers in Venezuela, the threat of unregulated exploitation of these resources
is present due to the increased demand of Asian markets, the lack of evaluation of
their populations and absence of norms that regulate their local fisheries and
commerce.
A possible menace that requires more attention is the marketing of live orna-
mental species for sea water aquaria, about which there are no official data. Also,
some species are dried and sold as ‘‘souvenirs’’. These include the starfishes
O. reticulatus and Astropecten articulatus and, in a lesser degree, the sand dollar
(called ‘‘locha de mar’’ in Venezuela), Mellita quinquiesperforata. Sellers can be
seen in the more visited beaches of Margarita Island and also during the high
touristic seasons in Chichiriviche (Falcón state) in western Venezuela as well as in
Mochima (Sucre state) and Coche Island (Nueva Esparta state) in eastern Vene-
zuela. This activity is not regulated but is expected to increase with the touristic
development and will become an environmental problem worldwide (Lunn et al.
2008).
7.7 Recommendations
In spite of the heterogeneity of the ecoregions in Venezuela, the fact that some of
them have proposals for management and conservation, and the importance of
echinoderms in them, studies on echinoderms are still few. More intense research
about their biodiversity and ecology is necessary.
Although the present analysis has attempted to provide information on the
different classes, families and species of the extant echinoderms in some
ecoregions of Venezuela, the distribution of echinoderms requires incorporation of

more detailed spatial analyses that include the entire Venezuelan coastline. It is
expected that they will generate information about their density and other popu-
lation parameters, including growth, reproduction, feeding and their relation to the
biotic and abiotic characteristics of the environment. It is important to evaluate the
environmental factors that could influence their presence in a certain location,
including studies about their vulnerability to climatic change. This information
will be of great importance, not only for the scientific knowledge and a better
description the autoecologic (individual) and synecologic (community) relations,
but also for development of strategies leading towards the sustainable production
of echinoderms as a resource and their administration, as well as the search of
alternatives to increase their production through aquaculture.
Some studies have considered the echinoderms as model species for ecotoxi-
cologic analyses, as in the case of echinoids (Esclapés 1999). However, there has
been no continuity in these types of evaluations, which are very important because
the Venezuelan coastline is influenced by many anthropogenic influences that have
produced major changes to many habitats.
The studies on sea urchin aquaculture in Venezuela have been encouraging,
having completed the techniques for juvenile production. It is recommended that
such techniques be optimized and concentrate further studies on post-juvenile
growth. Because the gonad is the marketed product from sea urchins, it is nec-
essary to perform studies on the optimization of it production. In contrast with sea
urchins, there has been no research on sea cucumbers aquaculture in Venezuela,
despite their high market value and their detritivorous condition which makes
them attractive for cultivation. Considering that techniques for sea cucumber
aquaculture are being optimized in other countries, particularly in Asiatic regions,
we should take advantage of this technological expansion to set up culture
packages for Venezuelan species. This could also bring environmental benefits,
because it would decrease the current pressure on echinoderm resources from
illegal fisheries. In this sense, population studies on sea cucumber resources are
necessary for their administration. For this, the gathering of information on illegal
fishing is important, such as extracted species, the quantification of the landings
and the identification of places of extraction and commercialization.
The incidental fishing of echinoderms as bycatch also merits study, in particular
the effect of trawls for the ark shell (A. zebra) and other fishing gears on echi-
noderm populations in different zones of Venezuela.
It is possible that the list of echinoderms in Venezuela be larger than the one
presented here. In particular, the benthos of the deep ocean of Venezuela has been
little studied. Most expeditions covering this area have been made by institutions
of other countries where a great amount of information is deposited. Since access
to these sources is usually difficult, it is necessary to prepare catalogues of echi-
noderms with information about their biology, ecology, distribution and fisheries
to improve access to knowledge about local echinoderms and with it the promotion
of their studies in Venezuela. These catalogues should include paleontological
information of the species, produced from studies under a geologic and a
paleobiogeographical context. This would allow the ordination in time and space
of the ancestral forms with evolutionary continuity, as well as the origin, distri-
bution and ecology of this fauna in the Caribbean Province.
Acknowledgments The collaboration of L. León is appreciated for critical reading of the section
on fisheries, as well as that S. Paul for providing bibliographic material. J. Reyes, N. Noriega and
Y. J. Díaz, collaborated in the present study during their doctoral studies. The Fundación para la
Investigación y Desarrollo de la Acuicultura del Estado Sucre (FIDAES) partially contributed in
the research described in the section on aquaculture. This is contribution No 18 of the Fundación
para la Investigación y Desarrollo de la Acuicultura del estado Sucre (FIDAES).
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Chapter 8
Echinoderms of Peru
Yuri Hooker, Elba Prieto-Rios and Francisco A. Solís-Marín
8.1 Introduction
8.1.1 Geographic and Geologic Characteristic of the Peruvian

Coast
The Peruvian littoral extends 3,080 km. It goes from the border with Ecuador
(approx. 3°230 S) to the south (18°S) at the border with Chile. The Peruvian coast
faces the South East Pacific Ocean. Its coast is characterized by arid conditions,
Y. Hooker (&)
Laboratorio de Biología Marina, Departamento de Ciencias Biológicas y Fisiológicas,
Universidad Peruana Cayetano Heredia, Av. Honorio Delgado 430,
Urb. Ingeniería, S.M.P, Lima, Peru
e-mail: yuri.hooker@upch.pe
Y. Hooker
Unidad Marino Costera, Servicio Nacional de Áreas Naturales Protegidas (SERNANP),
Ministerio del Ambiente, Calle Diecisiete No 355, Urb. El Palomar - San Isidro, Lima, Peru
E. Prieto-Rios
Facultad de Ciencias Biológicas, Biología, Universidad Nacional Mayor de San Marcos,
Mesa de partes. Ciudad Universitaria de San Marcos, Av. Venezuela s/n, Lima 1, Peru
E. Prieto-Rios
Posgrado en Ciencias del Mar y Limnología, Instituto de Ciencias del Mar y Limnología
(ICML), UNAM, Apdo. Post. 70-305, 04510, Mexico, D.F., Mexico
F. A. Solís-Marín
Colección Nacional de Equinodermos ‘‘Ma. E. Caso Muñoz’’,
Laboratorio de Sistemática y Ecología de Equinodermos,
Instituto de Ciencias del Mar y Limnología (ICML),
Universidad Nacional Autónoma de México (UNAM),
Apdo. Post. 70-305, 04510, Mexico, D.F., Mexico

278 Y. Hooker et al.
especially south of 13°450 S where the Atacama Desert starts. The coastal deserts
are crossed by fertile valleys formed by rivers of the western slopes of the Andes,
which carry water mostly in summer months (December–April). These valleys are
vegetated oasis surrounded by extremely arid deserts that may lack any rainfall for
years. The desert plains of the Peruvian coast are limited to the east by the Andes,
which has heights over 6,700 m. North of 6°S, the coast receives minor seasonal
rains during the summer months, which allow the development of a peculiar dry
coastal forest. This extends to the border with Ecuador, where it becomes greener
and humid because it is located in a rainy area. At the northern tip of the Peruvian
coast is a mangrove forest which has a length of 20 km. This ecosystem extents
from Ecuador to the Tumbes River delta in Peru and is the southern reach of
continuous mangroves in the eastern Pacific. A small, isolated mangrove forest
occurs south of Tumbes, at the mouth of the Rio Piura (5°300 S), the most southern
reach of the ecosystem in Peru.
The general morphology of the coastline along the Peruvian coast lacks great
geographical variants (Fig. 8.1). The most noticeable changes of the coast are the
Sechura Bay-Illescas Peninsula (5°380 –6°540 S) and the Paracas Peninsula
(13°510 S). There are few large bays, such as San Juan de Marcona, San Nicolas,
Independencia, Paracas, Ancon, Tortugas, Samanco, Ferrol, Sechura, and Paita.
Along the Peruvian coast are 77 islands (94.36 km2). The biggest ones are Lobos
de Tierra off Lambayeque (16 km2); the island of San Lorenzo, off Callao (the
largest island with 16.48 km2) and the island of San Gallan (9.32 km2) and La
Vieja (11 km2) off Paracas. Geologically the littoral is formed primarily by rocky
coastlines of sedimentary origin, igneous rocks, and extensive sandy beaches
(Sandweiss et al. 1996). Petersen et al. (1972) described the geomorphology of the
Peruvian coast. North of Piura and south of Tumbes, plains are formed by an
elevated sedimentary platform, which dominates the coastline (western shelf) with
sediments of Upper Cretaceous and Tertiary origin. This area has creeks and water
run-off, formed during El Niño events which bring intensive rainfall. Between
Paita and Punta Illescas, the rocks are metamorphic, igneous, and slate. In Lam-
bayeque and part of La Libertad, desert plains, wide beaches and coastal dunes
dominate the coastline. Only offshore do some Paleozoic rocks reach the surface,
forming the Lobos de Tierra and Lobos de Afuera islands. The coast of Ancash is
dominated by igneous rocks from the Andes mountain range, which at this area
approach the coastline, forming one of the most variable geomorphologic sectors
of the coast, with numerous bays, islands, emerging rocky areas, and cliffs.
The coastline at Lima province is composed of a variety of geological features
that include sedimentary coastal terraces, multiple intrusions from igneous rocks
from the Andes that penetrate the sea, sandy plains, and wide valleys associated
with numerous rivers of seasonal flow. At Ica province, from Paracas to the south,
a geological complex of sedimentary rocks of diverse origin and age dominates the
coastline, penetrating towards the sea on the Paracas Peninsula. The eastern Andes
mountain range has wide desert plains up to 80 km in width, with spurs of low
height that penetrate into the sea. Erosion has formed some high islands such as
San Gayan and La Vieja (Independencia). In this coastline of exceptional
8 Echinoderms of Peru 279
Fig. 8.1 Bathymetry of the Peruvian territorial sea and localities where shallow water
echinoderms have been collected by Universidad Peruana Cayetano Heredia (UPCH). Isobath
1 (blue line) 50 m; isobath 2 (red area) 200 m; isobath 3 (yellow area) 1,000 m. Maximum depth
6,280 m
characteristics, emerging sedimentary deposits offer a rich register of fossils that

date to the Carboniferous. Further to the south, at the Arequipa region to near the
Chilean border, the Andes Mountains extend closer to the sea, forming edges
dominated by recent igneous rocks that form cliffs, deep coves, and islands,
interspersed by valleys with seasonal rivers. There are long sandy beaches and
coastal dunes, forming a coastline with high geomorphologic diversity. This in
turn generates great habitat diversity that favors the development of complex
biological communities. The marine sediments between 7° and 10°300 S are mainly
sand, which dominate the wide continental shelf. Sand also appears in patches
towards the south. Mud with high organic content covers the continental shelf and
slope south of 10°300 S and may appear in coastal patches to the north (Delgado
et al. 1987). The continental shelf of Peru has an average depth of 200 m. The
continental slope starts at that depth and descends rapidly towards the Peru ocean
trench that reaches 6,000 m in depth. This ocean trench is formed at the edge of
contact of the Nazca plate and the South American continental plate (Meschede
and Barckhausen 2000).
The continental shelf is very diverse. The area located off the Tumbes and Piura
provinces is narrow, with an average width of 40 km. It is wide from the Illescas
Peninsula (Piura) to the Paracas Peninsula, reaching up to 140 km in width at
Chimbote and 70 km off Lima. From the Paracas Peninsula to the Chilean border
the average width varies between 2 and 4 km (Fig. 8.1). Geotectonic blocks as
well as pyrogenic and orogenic submersions form the continental shelf. The group
of marine rocks of most of the Peruvian coastline was elevated due to tectonic
action by the end of the Pleistocene, emerging from the sea and penetrating in
many places at the coastline up to 100 km inside the continent. The rocks from the
continental shelf and the Peruvian littoral are made primarily of Paleozoic and
Mesozoic sedimentary rocks. Granite rocks, granodiorite rocks, and gneiss rocks
occur at a few localities (Petersen et al. 1972).
8.1.2 Oceanography and Marine Circulation
The oceanographic characteristics and the marine diversity of the Peruvian sea are
ruled by a complex system of currents, which produce one of the most important
upwelling systems in the world.
Most of the Peruvian sea is influence by the Peru Current (Humboldt), which
begins approximately at 40°S as a consequence of the winds produced by the
anticyclonic gyre of the South Pacific Sea. The flow of a branch of this current, the
coastal Peruvian Current, is pushed by the trade winds. It follows the topography
of the South American coast until 6°S (Illescas Peninsula) where it deviates to the
east. This current is characterized by temperate waters through the length of its
route, with temperatures between 13 °C and 18 °C according to year, season and
geographic position (Fig. 8.2). These temperatures are much colder than expected
because it is found in the tropics, close to the equator. This is caused by the intense
upwelling that carries cold waters rich in nutrients to the sea surface. These waters
have a stable salinity around 35 %. As suggested by its name, this current flows
along the coast with a width that varies between 96 and 160 km with a depth
normally below 200 m. The other branch of the Peru Current, The Peruvian
Oceanic Current, moves from south to north, to the east of the meridian 82°W and
reaches depths of 700 m. This oceanic current also has temperatures that are lower
than those of tropical oceanic waters (17–20 °C) due to its subantarctic origins
(Pizarro 2001; Tarazona et al. 2003).
In the Pacific Ocean, over the equatorial line, the Equatorial Counter Current
moves towards the east until it collides with the South American continent.
A branch of this stream deviates and moves to the south along the coast of Ecuador
and arrives to the north of Peru, where it is named El Niño Current. This stream of
warm waters flows up to Cabo Blanco (4°150 S–81°140 W) where it turns towards
the west to form part of the Equatorial Current. The area between Cabo Blanco and
Punta Illecas is recognized as a region of tropical water masses where waters
coming from the north mix with a branch of the Peruvian Current that maintains a
weak flow towards the north (Hooker 2010).
Beneath the mentioned surface currents, there are other subsurface current. The
Equatorial countercurrent (or Cromwell Current) flows from west to east towards
Ecuador. A branch deviates and flows to the southeast to near the Peruvian north
coast between 5° and 8°S. It flows under the Peruvian Current and in opposite
direction (from north to south). It is close to the coast between Paita and Punta
Falsa (5–6°S), flowing between 50 and 300 m in depth. It moves away from the
coast to be integrated into the Subsurface Peruvian Current and continues to the
south (Cromwell 1953). One of the most important characteristics of the current is
a high concentration of oxygen, higher than 1.0 ml-1 at 100–200 m (Morón 2000)
permitting processes that will be discussed later on Pequeño (2000) discusses the
Intermediate Antarctic Water masses in deeper waters. In southern Chile they are
rather shallow, going deeper to the north, reaching a depth of 1,500 m at
approximately 35°S and even deeper to the north. He also mentions that off Chile,
at depths of 3,500–4,200 m, the relatively salty, but very cold, Deep Antarctic
Currents flows towards the north over the Pacific sea floor, crossing probably the
Chilean-Peruvian trench. This facilitates the displacement of ichthyofauna of
Antarctic origin towards less than 20°S, off the Peruvian coast. Pequeño (2000)
also mentions that some authors (like Petersen et al. 1972) described the presence
of a mass of intermediate water of the southeast Pacific sea or Intermediate
Antarctic waters (between 500 and 1,500 m deep) located off Chile, Peru and
Ecuador
8.1.3 General Ecological Characteristic
Peru is situated in a peculiar geographical area where temperate waters coming

from the south and tropical waters from the north meet, generating a diverse and
dynamic ecological system. Furthermore, the recurrent phenomenon called ‘El
Niño’ makes the ecological processes even more complex, altering the abundance
and distribution of marine organisms. The Peruvian sea is considered one of the
more productive of the world because an intense upwelling system in most of
the littoral develops a productivity that has made Peruvian fishing industry one of
the most important in the world. The upwelling system is closely related to the
Peru Current that flows continuously from south to north, pushed by the trade
winds. Upwelling transports waters from depths of 50–150 m towards the surface
in areas close to the coast, carrying nutrients accumulated over the bottom
of the continental platform. Upwelling waters are characterized by low
concentrations of oxygen and high concentrations of nutrients (Zuta and

Guillén 1970; Strub et al. 1998).
The arrival of these nutrients to the surface generates intense primary
productivity of around 1 kg C m-2 y-1 (Tarazona et al. 2003). The high abun-
dance of plankton is the base of a trophic chain where the Peruvian anchoveta
(Engraulis ringens) is the dominant species and the primary food of enormous
populations of guano birds whose droppings (guano) are accumulated over the
islands and capes. Wind action spreads some of the guano, fertilizing the marine
waters even more (Palomares et al. 1987).
The influence of the upwelling reaches up to a depth of 100 km out into the
ocean (Tarazona et al. 2003). Its influence is also reflected in coastal waters.
The nutrients allow great beds of brown macroalgae as well as other algae to
develop. They are used by herbivorous organisms, such as several species of sea
urchins and gastropod mollusks.
Nevertheless, the primary productivity does not always favor the abundance of
organisms. In the area occupied by the Peruvian Current, oxygen decreases
rapidly, reaching hypoxic conditions (\1 ml l-1) at depths below 20 m (Rosen-
berg et al. 1983). In waters between 100 m and 500 m in depth, oxygen con-
centration usually is almost zero (\0.1 ml l-1) so diversity there is at a minimum.
The cause is the great abundance of plankton in surface waters that, after death,
generate a constant organic rain that decomposes as it sinks. The bacterial activity,
both in the water column as well as over the organic sediments accumulated on the
sea bottom, consumes the dissolved oxygen and generates a minimum oxygen
layer that extends the length of the Peruvian Current (Lama et al. 2009).
Outside the continental shelf oxygen reappears in the water column at depths
below 50 m carried by the Subsurface Peruvian Current. Over the bottom, at
600 m, biological diversity increases, reaching higher values at 1,000 m in depth,
where oxygen-rich Antarctic Intermediate Waters occur (Lama et al. 2009).
In northern Peru, the tropical oceanographic conditions result in totally different
ecological characteristics than those occurring south of 6°S. Tropical currents
coming from the north maintain surface temperatures at coastal areas usually
between 20 and 25 °C (Fig. 8.2). This allows the establishment of a purely tropical
ecosystem with a high diversity. Hooker (2009) indicates they include more than
70 % of the total richness of the Peruvian littoral at shallow depths. This region,
especially in the northern tip close to the border with Ecuador, is the only place
where the influence of low salinity waters is significant due to the great volume of
freshwater coming from Ecuadorian rainy areas, generating mangrove forests
between the delta of the Tumbes River and the Zarumilla River that mark the
border with Ecuador. Sediments carried out by these rivers also influence coastal
diversity because waters become murky close to the mouths of the rivers.
The effect of suspended sediment is dissipated to the South, where clearer waters
are found between 3°520 and 4°150 S. Species richness increases, especially on
rocky reefs. It should be pointed out that Peruvian marine waters do not have any
coral reefs and there is only one record of a hermatypic coral species.
Fig. 8.2 Sea surface temperature distribution in the Peruvian territorial sea during winter during
a year of ‘‘normal’’ oceanic conditions (11 August 2004) (source NAVO/NOAA)
In the entire sector between the border with Ecuador and 4°150 S the layer of
oxygen that exists south of 6°S cannot be detected. There, life is present in all
bathymetric gradients, maintaining high richness even below 1,000 m depth. The area
called ‘‘Banco de Mancora’’, located about 40 km off Zorritos, a submarine mountain
plateau between 90 and 500 m in depth, has high richness exists and some fish and
invertebrate species seem to maintain their population centers (Edgar et al. 2004).
Another important factor affecting richness in the Peruvian littoral is the El Niño
phenomenon. During these oceanographic events, trade winds push and weaken the
Peruvian Current, causing the current to lose strength. As a result, tropical waters
invade the Peruvian coast, reaching even to the Chilean north. The entire temperate
waters fauna is seriously affected by temperature increases up to 10 °C at some
points of the coast. Moreover, when upwellings decreases or disappear, nutrients
and primary productivity is notably reduced. This affects macroalgae and filter
feeders and the system collapses. In the tropical area of Peru, the water temperature
in the littoral reaches 30 °C (Fig. 8.2) and heavy rainfall over the desert brings
enormous quantities of sediments to the sea. This makes the sea completely turbid
and the submarine rocky formations become clogged due to the sediments, killing
the benthic fauna and transforming the coastal sea bottom into mud. The effect
of the sediment is even perceptible 2 years after the event, where compact mud
is found inside rock crevices and in the shells of dead barnacles.
During El Niño, the entire area is invaded by tropical waters and an aquatic
tropical fauna is established, significantly expanding its distribution to the south.
This expansion is only temporary. When the oceanographic conditions become
normal after 1–2 years, this fauna gradually disappears, leaving only some species
that are resistant inside localities that act as refugia, especially in enclosed bays
where temperature is normally higher than in open waters.
Few papers are dedicated to the taxonomy or systematic research on Peruvian

echinoderms. Alexander Aggasiz (1881) reported seven echinoid species, mostly
from shallow waters in Callao and Paita (at Paita Province). Later on, Clark (1910)
published the largest echinoderm list ever reported. He described 25 seastar spe-
cies collected in Peruvian territorial seas, 10 ophiuroids, 12 echinoids species, and
seven species of sea cucumbers. Deichmann (1941, 1958) reported 14 species of
holothurians collected between 1932 and 1938 during the Allan Hancock expe-
ditions to the American Pacific (Velero III and IV).
More recently, Bluhm and Gebruk (1999) made underwater images observations
at depths between 4,140 and 4,160 m for three Peruvian localities. They reported 11
species of deep sea holothurians and some non identified sea cucumbers. Hooker
et al. (2005) reported 39 echinoderm species at Islas Lobos de Afuera (Lambayeque):
seven Asteroidea, eight Ophiuroidea, 11 Echinoidea and 13 Holothuroidea.
Prieto-Rios (2010) reported 22 species of sea cucumbers for Peruvian waters,
including both shallow and deep areas. Finally, Prieto-Rios et al. (2011) reported
Florometra magellanica (Bell, 1882) in Piura between 360–814 m, the first crinoid
species ever recorded for Peruvian waters.
The Peruvian littoral is divided into two main areas included in two biogeo-
graphical provinces (Spalding et al. 2007): Warm Temperate South-Eastern Pacific
(Peruvian Province) and Tropical Eastern Pacific (Panamic Province). The former
is associated with the Peruvian Current with a high level of endemism. The
Temperate Eastern Pacific, on the other hand, is associated with warm waters
coming from the north, mostly by a branch of the South Equatorial Current. The
fauna of this province is widely distributed from the Gulf of California (Mexico) to
north of Piura region, Peru (4°150 S). The area between Cabo Blanco (4°150 S) and
Punta Aguja (6°S) is a transition zone between these two provinces (Hooker 2009).
Peruvian echinoderms are represented by 215 species: Crinoidea (1 species),
Asteroidea (64 species, distributed in seven orders and 22 families), Ophiuroidea
(42 species, distributed in two orders and 11 families), Echinoidea (35 species,
distributed in seven orders and 14 families), and Holothuroidea (73 species,

distributed in six orders and 16 families). Only 17 species are found in the
Temperate South Eastern Province, 64 in the Tropical Eastern Pacific, four are
found in both provinces. Seventy eight are deep water species. That means that
20 % of the 84 shallow water species are temperate, 75 % are tropical and 5 % are
both temperate and tropical.
When dealing with biogeography and the distribution of Peruvian echinoderms,
it is very important to use information gathered in normal oceanographic years.
This is because many tropical species widen their geographical distribution
southward in El Niño years (every 7–12 years). The species that extend their
distribution in El Niño years do so only temporarily, generally for less than
2 years, disappearing later from the colonized area. Our information has been
gathered during several years of observations and collections in about 20 localities
on the Peruvian coast.
8.3.1 Temperate South Eastern Pacific
South of Punta Aguja, around the Illescas Peninsula (6°S), shallow water echinoderm
richness is low, with most species belonging to the Warm Temperate South Eastern
Province and endemic to the Peru Current. In this area no differences in species
composition are found in the intertidal zone and abundance variations are not
significant. However, in the sublittoral zone differences do appear in the distribution
of some species, especially those that are mainly Chilean.
Species representative of the rocky intertidal and upper sublittoral are Stichaster
striatus (Fig. 8.3b), Heliaster helianthus (Fig. 8.3d), Tetrapygus niger and Pattalus
mollis. It is common to find juveniles of T. niger in rocky walls exposed to wave
action feeding on calcareous crustose algae. P. mollis occurs inside mussel
colonies. They use their tentacles to capture juvenile mussels, small crustaceans,
and snails detached from their colonies by wave action (Fig. 8.3h). The seastars
S. striatus and H. helianthus feed in the intertidal below the mussel belt where they
are not exposed to air during extended periods of low tide. The brittle star Ophiactis
kroeyeri lives inside mussel colonies by the thousands. A study carried out in Santa
Island, Ancash (Hooker et al. 2011) found an average of 27,000 ind m-2 inside the
mussel colonies.
The representative species in the sandy intertidal and upper sublittoral is
Athyonidium chilensis. It lives buried in sand. It has a U-shaped body. The ten-
tacles extend out on the sand and the cloaca shows just at the sediment surface.
This species feeds on suspended material from the water column.
In shallow subtidal waters the widely dominant species are the sea urchins
T. niger and Caenocentrus gibbosus (Fig. 8.4b) and Lytechinus semituberculatus
(Fig. 8.5c). Along the Peruvian central coast both species are equally abundant.
Tetrapigyus niger lives in shallower waters, usually over 5 m in depth, while
C. gibbosus is abundant from less than 1 m to about 10 m in depth. This species is
Fig. 8.3 Echinoderms from Temperate South Eastern Pacific (Peruvian Province): a Patiria
chilensis (Lütken, 1859); b Stichaster striatus Müller and Troschel, 1840; c Luidia magellanica
Leipoldt, 1895; d Heliaster helianthus (Lamarck, 1816). e Loxechinus albus (Molina, 1782);
f Arbacia spatuligera (Valenciennes, 1846); g Pseudocnus dubiosus (Semper, 1868); h Pattalus
mollis Selenka, 1868. (Pictures by Y. Hooker)
Fig. 8.4 Echinoderms from the Tropical Eastern Pacific (Panamic Province): a Astropecten
armatus Gray, 1840; b Phataria unifascialis (Gray, 1840); c Paulia horrida Gray, 1840;
d Ophiocoma aethiops Lütken, 1859; e Ophiothela mirabilis Verrill, 1867; f Ophiothrix
magnifica Lyman, 1860; g Eucidaris thouarsii (Agassiz & Désor, 1847); h Astropyga pulvinata
(Lamarck, 1816). (Pictures by Y. Hooker)
Fig. 8.5 Echinoderms from the Tropical Eastern Pacific (Panamic Province): a Centrostephanus
coronatus (Verrill, 1867). b Caenocentrotus gibbosus (L. Agassiz & Desor, 1846); c Lytechinus
semituberculatus (L. Agassiz & Desor, 1846); d Mellitella stokesii (L. Agassiz, 1841); e Dense
aggregation of Neothyone gibber (Selenka, 1867); f Cucumaria flamma Solís-Marín & Laguarda-
Figueras, 1999; g Holothuria cf. inhabilis; h Isostichopus fuscus (Ludwig, 1875). (Pictures by
Y. Hooker)
numerous along the coasts of central and northern Peru, mainly at Foca island in
the middle of the biogeographic transition area. It is very abundant at this locality,
covering most of the rock surfaces. To the north, where echinoderm diversity
increases, this sea urchin is rare. It is also very numerous in the bays of Sechura,
Lobos de Tierra island, Ancash (mainly Samanco) and Pucusana. This species,
which extends north to Colombia, seems to have its higher concentrations not only
in the area of its biogeographic transition but in the northern bays of the Warm
Temperate South Eastern Province. These are all closed and therefore warmer than
the surrounding oceanic waters, environments similar to that found in the transi-
tion area.
In deeper areas, usually below 10 m, the sea urchin Arbacia spatuligera
becomes abundant. It prefers organic remains of decomposing algae and deposited
organic matter as food, especially in the interface between rocks and sand bottoms
(Fig. 8.3f). It is also numerous in soft bottoms with organic matter. Loxechinus
albus becomes abundant from Paracas southwards (Fig. 8.3e). It is found mainly in
areas where kelps are present since it feeds on pieces of algae that fall to bottom
and that are picked with the tube feet. Old fishermen of the area affirm that L. albus
used to cover extended areas and gave a reddish color to the bottom where they
lived. When commercial exploitation of the species began, it became scarce in
most of its distribution area, remaining abundant only in remote localities or in
protected places, such as Punta San Juan in Marcona.
The seastar S. striatus prefers the intertidal and wave exposed well oxygenated
shallow waters, while H. helianthus and Luidia magellanica are found in deeper
waters (Fig. 8.3c). The last two species are active, voracious predators. Heliaster
helianthus is an effective biological control of turbinid snails and mussels, as well
as other small animals. Luidia magellanica may also feed on larger prey, including
A. spatuligera and T. niger. Old fishermen say T. niger was not as abundant as it is
today and that now it is out of control, covering large areas areas in some loca-
tions. In the 90s T. niger was a major prey of the wrasse Semicossiphus darwini (Y.
Hooker personal observations). This fish is now almost extinct. This may be one of
the causes of the overpopulation of the sea urchin.
Patiria chilensis, a small seastar, occurs along the entire Warm Temperate
South Eastern Province of Peruvian waters (Fig. 8.3a). The ophiuroid Ophiactis
kroeyeri, is a species that inhabits a wide diversity of habitats and depths in high
concentrations. High amounts of organic matter are required to support its vast
aggregations. It is usually found inside colonies of intertidal mussels, below rocks
with accumulations of detritus, inside the exhalant canals of large sponges, and
mainly at depths greater than 60 m because decrease oxygen reduces its abun-
dance. North of Lima to Paita Bay (Ancash), Ophiothrix magnifica dominates
muddy bottoms. This species is a northern element of the Warm Temperate South
Eastern Province as well as the transition zone to the Tropical Province. The sea
cucumber Pseudocnus dubiosus (Fig. 8.3g) dominates muddy, hypoxic, unstable
bottoms, forming large aggregations. It is the most representative member of this
habitat.
There are many doubts about the taxonomy of some Peruvian brittle stars like
O. kroeyeri. It probably includes at least two species that have not yet been
distinguished. More detailed studies are required.
8.3.2 Tropical Peruvian Eastern Pacific
Seventy five percent of the intertidal Peruvian echinoderm species are concen-
trated at the northern tip of Peru along less than 150 km of coastline. North of
Cabo Blanco (4°150 S; Fig. 8.1), all species are typical of the Tropical Eastern
Pacific, with the exception of C. gibbosus, Pentamera chierchia and P. chiloensis,
which are widely distributed. Pentamera spp. are even found in cold waters and
cannot be considered members of any given province.
On the other hand, Ophiothrix magnifica, a brittle star common at north Peru
(Fig. 8.4f), enters the Peru Current to Paracas and is successfully established in
enclosed shallow bays where warmer waters allow the temporary presence popu-
lations. This species is not considered widely distributed since it appears only in
refugia where environmental conditions differ from the surrounding areas. The
region between El Ñuro and Punta Sal is where the best example of the Tropical
Eastern Pacific echinoderm community is found in Peru. Rocky reefs are scarce in
northern Peru, represented mainly by flat strata of sedimentary rocks. In this area
some taller reefs appear, mainly at Punta Sal. Further north, the diversity is reduced
because of the sediment load and turbidity caused by the discharge of adjacent rivers.
In the intertidal, the most common species are Ophioderma panamensis,
Ophiocoma aethiops (Fig. 8.4d), O. magnifica, Arbacia incisa and Echinometra
vanbrunti, mainly in tide pools and below rocks. In the subtidal zone the most
abundant species are Eucidaris thouarsii (Fig. 8.5g), and Phataria unifascialis.
Pentaceraster cumingi used to be abundant, but it has become rare since it is col-
lected to be sold as souvenirs. The most common sea cucumber is Cucumaria flamma
(Fig. 8.5f) which populates the higher portions of reefs. Under the rocks, Holothuria
arenicola and H. impatiens are frequent species, together with O. panamensis and
O. aethiops (Fig. 8.4d). Another common species, O. magnifica, may be found in all
available cryptic habitats, including rocks, algae, sponge, ascidians, and gorgonians.
Ophiothela mirabilis is a brittle star that is always associated with gorgonians, where
it is attached to their branches in large numbers (Fig. 8.4e).
Astropyga pulvinata (Fig. 8.4h), a large sized sea urchin, is a very rare species in
Peru. It has been reported in two localities: north of Punta Sal, where there is a rather
large population composed mostly by large adults, and at Lobos de Afuera, where a
large group of juveniles on a rocky reef were found (Hooker et al. 2005). Some other
rare species are Toxopneustes roseus, known in Peru only from scarce reports at El
Ñuro, Isla Foca, and Islas Lobos de Afuera. Paulia horrida is a rare seastar found
through its whole distribution, reported in Peru only from Punta Sal (Fig. 8.4c).
This differs from the status of the seastar Nidorellia armata, which was frequent
but has become close to extinction due to extraction for the souvenir market.
Holothuria cf. inhabilis (Fig. 8.5g), a remarkable large sized sea cucumber
(about 30 cm in length in life), is frequent on the soft bottoms surrounding rocky
reefs, where it feeds on organic sediments. This species, reported as a possible new
species when it was collected at Islas Lobos de Afuera (Hooker et al. 2005), should
be reevaluated in order to clarify doubts concerning its taxonomy identity.
On shallow sandy bottoms the dominant echinoderms are the seastars Astro-
pecten spp., with A. armatus as the most frequent (Fig. 8.4a). The sand dollar
Encope micropora has also been reported from this habitat, but seems to be more
frequent in the biogeographic transition region. Luidia bellonae is rarely reported
in deeper waters (10–20 m) along the north Peruvian coast. However, it is fairly
common in Islas Lobos de Afuera. There are no records of this species south of
Islas Lobos de Afuera. Nevertheless, Clark (1910), Anonymous (2010) and
Morales-Montecinos (2011) reported it from central and southern Peru. In waters
below 50 m depth, L. superba is the dominant seastar, together with Clypeaster
europacificus. Both species are frequently caught in artisanal fishing nets.
8.3.3 Tumbes Mangroves
Northern Peruvian mangroves possess a typical Tropical Eastern Pacific fauna but
the reported faunal diversity is low. The most representative species is Mellitella
stokesii, a very common sand dollar in sandy tide pools and intertidal beaches in
the interface between the mangroves and the sea (Fig. 8.4d). Another abundant
species is the sea urchin Agassizia scrobiculata. Dense populations occupy the
bottom of tidal creeks at depths between 0.5 and 2 m at low tide. The brittle star
O. magnifica is also frequent, mainly between mangrove roots and inside sponges
and zoanthids. Another ophiuran, an unidentified species of the genus Ophio-
grammus, has been recorded in the muddy sand bottoms. A bright-orange morph of
Luidia brevispina is common on sandy beaches around mangroves, different from
the grey specimens known from Lobos de Afuera.
8.3.4 Biogeographic Transition Area Temperate South Eastern

Pacific-Tropical Eastern Pacific
As mentioned earlier, the transition area between these biogeographic provinces is

between Cabo Blanco (4°150 S) and Punta Aguja (6°S). This also includes the
islands Lobos de Tierra and Lobos de Afuera. Echinoderms of both provinces are
found in this refuge area as well as clear differences in abundance. For example,
O. magnifica is extremely abundant in almost every habitat, with the exception of
mud and fine sand bottoms. Another abundant species is Neothyone gibber,
especially at Lobos de Tierra where it covers 100 % of the vertical walls of large
rocks (Fig. 8.5e). Intertidal cliffs are dominated by H. helianthus. The sea
cucumber Psolidium dorsipes is found in the intertidal zone. In the subtidal,

P. unifascialis (Fig. 8.4b) has been rarely found on rocks. Athyonidium chilensis
appears occasionally below them.
Isostichopus fuscus (Fig. 8.4h) has been found throughout the area with low
densities. This sea cucumber is not as affected here by over-exploitation as it has
been in northern Peru. On the other hand, Lobos de Afuera is the only location
where Tripneustes depressus has been reported. This species was very abundant
there until 2005. It is said that fishermen working in the island extracted them for
their own use and for commercialization. This has decreased the populations
below commercial levels.
The islands of Lobos de Afuera possess species from both provinces. They may
be considered relatively tropical. The relative abundance of the species differs
from coastal waters (Hooker et al. 2005). Knowledge about the Lobos de Afuera
echinoderm fauna (Hooker et al. 2005), fishes (Hooker 2009) and sponges are
more related to the Galapagos Islands than to the coastal tropical waters of
northern Peru (Fig. 8.6).
8.3.5 Deep Sea Waters
The diversity of echinoderms in Peruvian deep waters is very high, reaching 46 %

of the total reported species for Peru. Richness has been reported mostly by old
marine cruises. The information was collected and analyzed by Maluf (1988). The
most recent reports come from deep water cruises organized by the Instituto del
Mar de Peru (IMARPE). These cruises collected a large number of species that
should increase our knowledge of the diversity of Peruvian deep waters.
Prieto-Rios (2010) published the first report for Peru of the sea cucumbers
Molpadia and Caudina californica at depths of 1,412 and 996 m, respectively.
Morales-Montecinos (in preparation) has the first collection for Peru of the sea
stars Thrissacanthias penicillatus, Ctenodiscus crispatus, Pectinaster agassizi,
Ceramaster leptoceramus, Lophaster furcilliger, and Hymenaster quadrispinosus.
Recently, Prieto-Rios et al. (2011) published the first report of a crinoids for Peru,
adding F. magellanica (Bell, 1882) (360–814 m) to the biodiversity list of deep
sea echinoderms. Some other species of sea cucumbers and seastar are now under
revision and brittle stars and more crinoid species are still awaiting a proper
taxonomic revision.
Historically echinoderms were of interest to Prehispanic Peruvian cultures. The sea

cucumber A. chilensis, locally called ‘‘ancoco’’, has been exploited and locally
consumed by the Mochica culture in the Lambayeque area (*6°500 S) for at least
Fig. 8.6 Bathymetric ranges and abundance (to 20 m deep) of the Lobos de Afuera echinoderms
in June 1999. R Rare (1–2 individuals); F Frequent (3–10 individuals); C Common (11–50
individuals); A Abundant (more than 50 individuals) (from Hooker et al. 2005)
2,000 years. Starfish were also important in the ancient Peruvian cultures, as ritual
offerings to the gods. This practice is still kept in the rites carried by the local
chamanes. Evidence for this is seen in their representation in different artistic
objects like pottery and paintings from the old Moche culture (Fig. 8.7a, b).
Fig. 8.7 a Painted Moche ceramic (2000 years old) depicting a seven armed seastar (from
Vergara and Sánchez 1996); b Moche iconography depicting a fishing scene including some
seastar; c dry seastar included in different handcrafts; d and e Pattallus mollis landing at
Chimbote Pier; f Ancoco (Pattallus mollis) processed, and g Sea urchin (Loxechinus albus)
processed and ready to sell at the markets in Lima and Ilo (Pictures by Y. Hooker)
Table 8.1 Landing information, total catch (t) of echinoderms per area in the region of Ancash-
Tacna, Peru, from 2001 to 2007 (from Sueiro 2009)
Common name Scientific name/year 2001 2002 2003 2004 2005 2006 2007 Total
Sea urchin Loxechinus albus 2111 1948 2059 1387 3010 80 1365 11960
Sea cucumbers Pattalus mollis- 5 190 8 1 21 86 567 978
(‘‘mixture of Holothuria sp.
species’’)
Seastar Stichaster sp. 5 0 0 0 0 0 0 5
Dry specimens of Stichaster striatus and Heliaster helianthus are used in these
rites. Both species are still sold as souvenirs (Fig. 8.7c) or in herb and mystic
objects (chamaneria) markets.
Exploitation of echinoderms in Peru is not older than 20 years. It began with
exportation of the red sea urchin Lytechinus albus gonads and later with some
species of sea cucumbers to Japan. The first sea cucumber formally and legally
exploited was I. fuscus in 1995, possibly when the fishing of the species began to
be controlled or even sold in some areas of the Galapagos Islands. Sueiro (2009)
mentioned the problem of exploitation of I. fuscus in the Galapagos and the
regulation measures taken for its protection. Although I. fuscus is an economically
important marine product in Peru (class ‘‘A’’ product), no protection measures are
taken in its fishery. Catches are sent to Ecuador immediately with no information
on the catch provided to authorities. Local fishing divers have provided infor-
mation that I. fuscus was eliminated in less than 3 years after fishing began from
areas in Peru, Lobos de Afuera and the region between Cabo Blanco and Punta
Mero, north of Punta Sal. Legal exportation of sea cucumbers began officially at
the end of the 1990s. Catches were principally composed of P. mollis and
Holothuria theeli. The last species became common after El Niño 1997–1998, with
large landings in Huacho (Elliott et al. 2005) and Chimbote. The last year this sea
cucumber was reported was 2002. Since that it has been replaced in fishing by
P. mollis captures (Fig. 8.7d–f).
Toral-Granda et al. (2003) and Sueiro (2009) published the catch and expor-
tation statistics of Peruvian sea cucumbers with a ‘‘mixture of species’’, probably
including A. chilensis (Tables 8.1 and 8.2).
Sueiro (2009) published data that included landings of the sea urchin L. albus, a
species that was always important for local consumption (Fig. 8.7g). It has been
taken to the edge of collapse, being very scarce nowadays. The Minister of Pro-
duction has established bans several times in order to protect and restore this
resource.
Tripnesutes depressus is a sea urchin that reaches large sizes at Lobos de
Afuera. Local fishermen reported that it was abundant. However, the presence of
purchasers resulted in its illegal and accelerated fishing, with the consequent
depletion of its populations. IMARPE’s landing records from Lambayeque report
it as ‘‘L. albus’’. However that species does not occur there, being restricted to
south of Paracas.
Table 8.2 Total export data (t) of Peruvian sea cucumbers to Hong Kong (dry, salted or pickled
product) 1999-September 2005 (from Toral-Granda 2008)
1999 2000 2001 2002 2003 2004 2005 Total (t)
Peru 4.1 7.3 3.8 1.8 8.3 19.9 31.0 76.2
Another kind of commerce for the Peruvian echinoderms that is usually not
taken into account which affects seriously populations of some species is the
souvenir market (Fig. 8.7c). Commerce of dry specimens of S. striatus and
H. helianthus in some tourist places of central and southern Peru is common.
These species are also used to make handcrafts. Some species are also sold dry on
the tropical north beaches, mainly Mancora. The list includes N. armata,
P. cumingi, P. pyramidatus, P. unifascialis, C. europacificus and E. micropora.
The first two species have become close to extinction because they were never
abundant and the reefs where they live are not common. C. europacificus is
accidentally caught by artisanal trawling boats. The effect of this on its populations
is unknown.
Sueiro (2009) mentions a trade exportation of 5 tons of S. striatus in the year
2001, but the objective of this commerce is unknown. The populations of this
seastar have been reduced due to its multiple usages given (ornaments, souvenirs,
mystic rituals, etc.). More recently most local universities and schools use them for
their teaching laboratories because they are easily collected in the intertidal.
No special effort has been made to develop echinoderm aquaculture techniques
in Peru. Nevertheless, the IMARPE’s Ilo Aquaculture Laboratory has been
working successfully on reproduction of L. albus. It is possible that in a short time
the aquaculture of this sea urchin will be developed for both market and scientific
interests.
8.5 Echinoderm Threats
There is no national strategy for the conservation and management of echinoderms

except for the sea urchin L. albus. Because new Marine Protected Areas (MPA) in
Peru were created, in 2010 and 2011, the country is now faces new challenges. The
Servicio Nacional de Areas Naturales Protegidas del Peru currently preparing Red
List files of threatened species in Peruvian MPAs. It is expected that several
echinoderm species will be included. The following list shows some of the pro-
posed species and their status following the criteria of the International Union for
Conservation of Nature (IUCN).
1. Nidorellia armata: Critically endangered.
2. Pharia pyramidatus: Endangered.
3. Phataria unifascialis: Almost threatened.
4. Stichaster striatus: Vulnerable.
5. Clyepaster europacificus: Insufficient data.

6. Loxechinus albus: Vulnerable.
7. Tripneustes depressus: Endangered.
8. Athyonidium chilensis: Almost threatened.
9. Isostichopus fuscus: Endangered.
10. Pattalus mollis: Almost threatened.
8.6 Recommendations and Concluding Remarks
Peru has 215 species of echinoderms inhabiting its territorial sea. Crinoidea (1
species), Asteroidea (50 species), Ophiuroidea (32 species), Echinoidea (23 spe-
cies) and Holothuroidea (53 species). This compilation is important to make
recommendations: It is necessary to do more ecological studies. The situation
previously described regarding publications about ecology of echinoderms in Peru
provides evidence of an unequal and limited development of ecological studies
related echinoderms. There are no papers that analyze echinoderm communities in
Peru. It is important to have more information about their density and other
population parameters, including growth, reproduction, feeding and their relation
to biotic and abiotic conditions in the environment. This would make it possible to
evaluation of environmental factors that could affect their presence in a certain
location and indicate their vulnerability to climate change. This information will
be of great importance for the basis for developing strategies leading towards
sustainable production of echinoderms as a resource and their administration, as
well as the search of alternatives to increase their production through aquaculture.
One of the main problems in Peru, as well as in neighboring countries, is the
lack of taxonomists who can work full time reviewing and identifying specimens
in biological collections. Deep water specimens as well as brittle stars should be
reviewed. This should increase greatly the number of species reported from Peru.
Complete knowledge of the echinoderm fauna will allow ecological research
directed towards understanding the function of each species in the ecosystem allow
researchers to monitor climate change associated with El Niño, and the distribution
and biogeography of the species.
Controlled production of commercial species should be developed in order to
reduce the impact of fishing on natural resources. Creation of new MPAs should
guarantee conservation of exploited species.
Acknowledgments To Dr. Tina Kameya Kameya, Unidad de Investigaciones en Biodiversidad,

Instituto del Mar del Peru (IMARPE), for providing access to IMARPE’s echinoderm collection.
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Chapter 9
Echinoderm from Brazil: Historical
Research and the Current State
of Biodiversity Knowledge
Carlos R. R. Ventura, Michela Borges, Lúcia S. Campos,

Leticia V. Costa-Lotufo, Carolina A. Freire, Valéria F. Hadel,
Cynthia L. C. Manso, José R. M. C. Silva, Yara Tavares
and Cláudio G. Tiago
9.1 Introduction
9.1.1 General Geographic and Oceanographic Features

of the Brazilian Coast
Brazil has a long coastal zone which extends from 4°300 N to 33°440 S with a total
area of approximately 1,550,000 km2. It is divided in 16 marine sedimentary
basins (Fig. 9.1). One half of this area (about 770,000 km2) extends up to 400 m
C. R. R. Ventura (&)
Museu Nacional, Universidade Federal do Rio de Janeiro,
Rio de Janeiro- RJ, 20940-040, Brazil
e-mail: ventura@acd.ufrj.br
M. Borges
Museu de Zoologia, Universidade Estadual de Campinas,
Campinas-SP, 13083-863, Brazil
e-mail: borgesm@unicamp.br
L. S. Campos
Departamento de Zoologia, Instituto de Biologia,
Universidade Federal do Rio de Janeiro, Rio de Janeiro-RJ, 21941-971, Brazil
e-mail: campos-lucia@biologia.ufrj.br
L. V. Costa-Lotufo
Departamento de Fisiologia e Farmacologia,
Universidade Federal do Ceará, Fortaleza-CE, 60430-270, Brazil
e-mail: lvcosta@secrel.com.br
C. A. Freire
Departamento de Fisiologia, Universidade Federal do Paraná,
Curitiba-PR, 81531-990, Brazil
e-mail: cafreire@ufpr.br

302 C. R. R. Ventura et al.
depth, while the other half (about 780,000 km2) ranges from depths of 400 to
3,000 m (IBAMA 2008). Along its 8,698 km of shoreline, including all
geographic features like indentations (e.g. bays) and capes, it comprises tropical
and subtropical environments (IBAMA 2008).
Brazilian’s continental shelf is variable in extension. It is larger in the north,
ranging from 148 km at the coast of Amapá State to 293.3 km at the mouth of the
Amazon River. It narrows in the northeastern region, ranging from 37 to 55 km.
From the southeast to the south it expands gradually to 185 km (IBAMA 2008). The
Brazilian Exclusive Economic Zone has about 4.2 9 106 km2 which includes dif-
ferent estuarine and marine ecosystems, such as mangroves, rocky shores, coral
reefs, sandy and muddy bottoms in shallow water and deep sea environments. Rocky
and calcareous bottoms are common along the northeastern coast while sandy and
muddy bottoms prevail along the southeastern and southern coasts (IBAMA 2008).
Two main marine currents flow on the continental margins of Brazil: the North
Brazil Current (NBC) and the Brazil Current (BC) (Stramma 1991; Silveira et al.
2000). Both currents are formed at 10oS and are associated with the western border
of the South Atlantic Subtropical Gyre (SASG) as two branches of the South
Atlantic Equatorial Current (SAEC) (Silveira et al. 2000). The NBC flows
northwards while the BC flows southwards along the continental margin up to the
Argentine Basin (33° to 38°S), where it meets the Malvinas Current (Brazil-
Malvinas Confluence) (Gordon 1988; Silveira et al. 2000). Therefore, the BC
prevails on the Brazilian coast.
The first 3 km of the water column of the BC is formed by five water masses:
the Tropical Water (TW), the South Atlantic Central Water (SACW), the Antarctic
V. F. Hadel
Centro de Biologia Marinha, Universidade de São Paulo,
São Sebastião-SP, 11600-000, Brazil
e-mail: vafhadel@usp.br
Cynthia L. C. Manso
Laboratório de Invertebrados Marinhos, Universidade Federal de Sergipe,
Itabaiana-SE, 49500-00, Brazil
e-mail: cynthia@phoenix.org.br
J. R. M. C. Silva
Departamento de Biologia Celular e do Desenvolvimento,
Instituto de Ciências Biomédicas, Universidade de São Paulo, São Paulo-SP,
05508-99, Brazil
e-mail: jrmcs@usp.br
Y. Tavares
Faculdade de Ciências, Filosofia e Letras de Paranaguá,
Universidade Estadual do Paraná, Pontal do Paraná-PR, 83203-280, Brazil
e-mail: ytavares@ufpr.br
C. G. Tiago
Centro de Biologia Marinha, Universidade de São Paulo,
São Sebastião-SP, 11600-000, Brazil
e-mail: clgtiago@usp.br
9 Echinoderm from Brazil 303
Fig. 9.1 Principal marine currents and Brazilian sedimentary basins in the South Atlantic Ocean
Intermediate Water (AIW), the North Atlantic Deep Water (NADW), and the
Circumpolar Upper Water (CUW) (Silveira et al. 2000).
The TW is superficial, warm and salty due to the strong solar radiation and
evaporation which characterizes the Tropical Atlantic (Silveira et al. 2000). While
the TW is carried to the south by the BC, it is mixed with the Coastal Water (CW)
current which is less warm and salty. Consequently, the TW is characterized by
temperatures higher than 20 °C and salinities higher than 36 % along the south-
eastern coast (Silveira et al. 2000).
The SACW is characterized by temperatures between 6 and 20 °C and salinities
between 34.6 and 36 %. It is formed by the sinking of a water mass at the Atlantic
Subtropical Convergence. When it meets the southern border of the SASG, it
spreads to the northwest forming the South Atlantic and Benguela Currents.
Afterwards it returns to the South America continental margin, carried by the
SAEC. Part of the SACW flows to the north towards the equator while the other
branch flows southwards below the TW (Silveira et al. 2000).
The AIW flows between depths of 700 and 750 m and is characterized by
temperatures ranging from 3 to 6 °C and salinities ranging from 34.2 to 34.5 %
(Silveira et al. 2000). The AIW divides into two branches that flow northwards at
28°S and southwards at 25°S (Silveira et al. 2000).
The NADW flows between depths of 1,500 and 3,000 m. It is a cold water mass
(3 to 4 °C) with salinities ranging from 34.6 to 35 %. It flows southwards up to 32°S,
but part flows back northwards at higher latitudes (Silveira et al. 2000). The CUW
may be considered as part of the BC at the Brazil-Malvinas Confluence, although the
exact definition of the BC at this latitude is controversial (Gordon 1988).
In summary, the BC shows different features along the Brazilian coast. It is
shallow, warm and salty at low latitudes (from 10 to 20°S) because it is composed
basically by the TW. The SACW is incorporated into the BC at 20°S and ascends
at 28 to about 750 m. Actually, from 25°S southwards, the BC is well-consolidated
and is composed by the TW, the SACW, the AIW and the NADW, forming a
water column 3 km in depth (Silveira et al. 2000).
Along the southeast continental shelf, from 22 to 23°S, a seasonal upwelling
phenomenon, the Cabo Frio Upwelling System, occurs. This provides nutrient
enrichment onshore (Franchito et al. 1998; Mahiques et al. 2005; Sumida et al.
2005). This upwelling is ecologically important because the surface layers of water
on the coast are formed by the CW and the TW with low levels of productivity
(Franchito et al. 1998; Silveira et al. 2000). Some geographic features of the
coastline and continental shelf favor upwelling in the Cabo Frio region. The ori-
entation of the shoreline changes abruptly from N–S to E–W and the continental
shelf narrows (Valentin 1984; Franchito et al. 1998). When strong northeast winds
prevail for several days, usually from September to March during spring-summer,
the surface water mass is moved offshore and is replaced onshore by the cold and
enriched SACW. When cold fronts reach this region from June to August in the
winter, the S-SW surface winds blow for a few days and the upwelling cycle is
interrupted. Therefore, the seasonality of the upwelling at Cabo Frio area is
associated with the movements of the SACW from offshore to onshore and vice
versa (Valentin 1984; Franchito et al. 1998).
9.2.1 Historical Remarks of Echinoderm Research in Brazil
Studies on echinoderms in Brazil have always been characterized by a general

approach in which they are considered only as part of the benthic fauna in a given
area. The history of such studies can be traced back to 1648 when Marcgrave (1942)
reported the first echinoderm species of the Brazilian coast. In his ‘‘Brazilian
Natural History’’ the echinoderms were included in the fourth book, which regis-
tered the fishes, crustaceans and zoophytes. As zoophytes he listed two sea stars,
probably Luidia senegalensis (Lamarck, 1816) and Oreaster reticulatus (Linnaeus,
1758), a brittle star, probably of the genus Gorgonocephalus Leach, 1815, and two
sand dollars, both probably Mellita quinquiesperforata (Leske, 1778).
The following records were made in the nineteenth century between 1867 and
1898. Among them, three publications must be highlighted: the catalog written by
Rathbun in 1879, the report written by Ludwig in 1882, and, 16 years later, the
check-list of echinoderms from the southern coast of Brazil (Santa Catarina State)
written by Müller in 1898 (Tiago 1998; Hadel et al. 1999; Tiago and Ditadi 2001;
Ventura et al. 2006).
The first studies on Brazilian echinoderm fossils were also published in the
nineteenth century. In 1878, Derby described a Devonian ophiuroid from the Ponta
Grossa Formation (Paraná Basin—southern Brazil), and in 1887, White described
13 species of Cretaceous echinoids from the northeast region, at Sergipe State.
In the twentieth century, paleontological research was extended to other basins:
in the middle-west region, at Goiás and Mato Grosso States (Clarke 1913;
Löfgreen 1937; Erichsen and Löfgreen 1940; Barbosa 1949; Almeida 1954;
Beurlen 1957; Del‘Arco et al. 1982; Bosetti and Quadros 1996), and in the north
region, in the Amazon Basin (Ferreira and Fernandes 1989).
Taxonomic studies on Brazilian coastal echinoderms were improved after the
second half of the twentieth century, especially due to the efforts of Luiz Roberto
Tommasi from the Oceanographic Institute of the University of São Paulo
(Instituto Oceanográfico da Universidade de São Paulo). Tommasi has studied the
five classes of echinoderms morphologically, and provided the first identification
keys (Tommasi 1957, 1958a, b, 1960, 1962, 1963, 1964a, b, c, 1965a, b, c, d,
1966a, b, 1967a, b, 1968, 1969a, b, c, 1970a, b, c, d, 1971a, b, c, d, 1972, 1974a, b,
1985, Tommasi and Lima-Verde 1970; Tommasi and Abreu 1974; Tommasi and
Oliveira 1976; Tommasi and Aron 1987, 1988; Tommasi and Hecht 1988;
Tommasi et al. 1988a, b). Due to his pioneer contributions, his studies were the
basis for subsequent taxonomic research in Brazil. In this period, some other
studies are also relevant, such as those published by Brito (1959, 1960a, b, c, d, e,
f, 1962, 1968, 1971, 1979, 1982), and, afterwards, those by Ávila-Pires (1983);
Albuquerque (1986); Monteiro (1987); Monteiro et al. (1992) and Manso (1988a,
b, c, d, e, 1989, 1993).
From 1990 onwards there was an increase in echinoderm knowledge, such as
studies on fauna (Hadel et al. 1999; Netto et al. 2005; Ventura et al. 2007),
community structure (Ventura 1991; Castro et al. 1995; Ventura and Fernandes
1995; Sumida and Pires-Vanin 1997), reproduction and feeding (Ventura et al.
1994, 1997, 1998, 2001; Hadel 1997; Junqueira 1998; Ventura 1998; Guerrazzi
1999, Alves et al. 2002, Carvalho and Ventura 2002, Tavares et al. 2004, Ventura
and Barcellos 2004; MacCord and Ventura 2004); development (Hadel 1997;
Vellutini and Migotto 2010; Contins and Ventura 2011), growth (Freire et al.
1992), local distribution (Mac Cord and Duarte 2002), environmental education
(Hadel and Tiago 2004) and taxonomy (Tiago 1998; Tiago and Ditadi 2001;
Borges et al. 2002a; Hopkins et al. 2003).
In 2009, Brazilian researchers gathered for the 1st Brazilian Symposium on
Echinoderms (Ventura et al. 2009) and again in 2012 for the second symposium.
On these two occasions, information on several current and ongoing research was
presented, as well as needs and suggestions to improve the knowledge of Brazilian

echinoderms.
In short, research on echinoderms in Brazil was developed on several subjects,
mostly on species of the classes Asteroidea, Echinoidea, Ophiuroidea and Holo-
thuroidea. Few studies have been made on crinoids, probably due to their low
diversity along the Brazilian coast.
9.2.2 Echinoderm Paleontological Research
Brazilian echinoderms are found from the Palaeozoic to the Tertiary basins. Ten
species of Crinoidea, two of Stylophora, three of Blastoidea, 63 of Echinoidea,
three of Asteroidea, one of Ophiuroidea and two of Holothuroidea have been
recorded (Tommasi 1999; Scheffler 2007). There are difficulties in increasing
knowledge of extinct echinoderms, particularly due to the large distances among
the basins and the few paleontologists devoted to the study of echinoderm fossils.
Besides, very little is known of the origins and geological history of macro and
megafaunal biodiversity in the deep-sea, including echinoderms (Campos et al.
2010). Even though it is recognized today that species richness of deep-sea is
comparable to, or even exceeds, that of continental shelves, the difficulties in
finding identifiable macro-fossils in deep-sea cores limits any assessment of the
geological history of specific benthic groups, including echinoderms (Gray et al.
1997, Smith and Stockley 2005). Smith and Stockley (2005) combined evidence
from echinoid phylogenetic trees and molecular studies and suggested deep-sea
nutrient enhancement over a long geological time has had a key role in the
development of its biodiversity.
According to Scheffler (2007), research on Devonian echinoderms (Ponta
Grossa Formation, Paraná Basin) started at the end of the nineteenth century with
the studies published by Derby (1878) on the Ophiuroidea, and the monograph of
Clarke (1913) on the Asteroidea. The first references on Crinoidea were made by
Löfgreen (1937) with findings from Goiás State, and by Erichsen and Löfgreen
(1940) with material from Mato Grosso State. Lange (1943) identified the first
crinoid from Paraná Basin while Petri (1948a, b) described 18 levels of the cri-
noids and cystoids from Ponta Grossa Formation in the same Basin. Caster (1954a,
b) described two new species of Stylophora. Crinoid fragments were recorded by
Barbosa (1949), Almeida (1954), Del‘Arco et al. (1982) and Bosetti and Quadros
(1996) in Mato Grosso State, and Beurlen (1957) in Goiás State. Ferreira and
Fernandes (1985) reported the crinoids from the Devonian of Brazil and Ferreira
and Fernandes (1989) recorded the crinoid species Erisocrinus sp. from the Car-
boniferous and the genera Ctenocrinus, Exaesyodiscus, Laudonomphalus and
Monstrocrinus from the Devonian of the Amazon Basin.
In the twenty-first century studies on the Devonian echinoderms were intensified
with new descriptions and taxonomic reviews such as those published by Alves and
Fernandes (2001) about Encrinaster pontis, and Scheffler et al. (2002), who
described parts of some crinoid. In addition, several papers describing the crinoids
and blastoids from the Ponta Grossa Formation were published by Scheffler (2003,
2004, 2006) and Scheffler and Fernandes (2003, 2007a, b). Descriptions of crinoid
species from the Maecuru Formation (Devonian of the Amazon Basin) were pre-
sented by Scheffler et al. (2006).
The Cretaceous echinoderms from Brazil were identified for the first time by
White (1887) with the description of 13 echinoid species collected by the Geologic
Commission of the Empire of Brazil from the Cretaceous of the Sergipe-Alagoas
Basin. Derby (1907) reported a little echinoid near the Maraú Peninsula, in the
Camamu Basin, Bahia State. In the 1920s and 1930s, Maury (1925, 1930, 1934a,
b, 1937) studied the basins from the northeast of Brazil. She described echinoids
from the Sergipe-Alagoas, Potiguar and Pernambuco-Paraíba Basins, and the
asteroid Uraster janettae from the Sergipe-Alagoas Basin. She also revised the
paper published by White in 1887 adding two new species. Oliveira (1957)
described a new species of Hemiaster from Pernambuco-Paraíba Basin.
Few decades later, some authors studied the echinoid fauna from the sedi-
mentary basins of the northeast and north of Brazil from the 1950s to the 1980s,
e.g. Santos (1958a, b) who recorded one species of Salenia sp. in the Camamu
Basin; Santos and Cunha (1959) Santos (1960a, b) who found new species in the
Potiguar Basin; Beurlen (1961, 1964a, b, 1966), who made systematic reviews,
identified two new echinoids species from the Araripe Basin and a new echinoid
species from the Pernambuco-Paraíba Basin. Tinoco (1963) reported fragments of
ophiuroids, echinoids, asteroids and holothuroids from Maria Farinha Formation,
Tertiary from Pernambuco-Paraíba Basin. Brito (1964, 1980, 1981a, b, c) gave
descriptions of the echinoid fauna from the Cretaceous and the Tertiary.
Smith and Bengtson (1991), who published a review with 39 species of echi-
noids from the Sergipe-Alagoas Basin, made some observations about echinoids
from the other Cretaceous Basins of northeastern Brazil. They described two new
genera (Colliclypeus and Acriaster) and a new species (Acriaster sergipensis) from
the Sergipe-Alagoas Basin. They also concluded that the Brazilian echinoid fauna
has great affinities with the West African fauna and a connection with the Texan
and Mexican fauna.
Ferré et al. (1996) described species of the roveocrinids crinoids from the
Turonian (Cretaceous) of the Sergipe-Alagoas Basin, and Ferré and Bengtson
(1997) recorded Roveocrinus spinosus from the Turonian for the same basin.
According to Morais (1998), Kraatz-Korschlau was the first paleontologist to
publish, in 1900, the presence of the echinoid fauna from the Oligo-Miocene of the
Pirabas Formation, in the north of Pará State. Carvalho (1926) pointed out simi-
larities between echinoid fauna of Fortaleza Island and Bragança County, Pará
State. Santos (1958a, b) recognized an endemic fauna in the Oligo-Miocene of the
Pirabas Formation. This fauna consisted of clyperasteroids and cassidulids, and
great similarities were found with the Oligo-Miocene Caribbean fauna, suggesting
echinoid dispersion to northern Brazil before the Miocene. Santos (1957) was the
first to describe cidaroids and spatangoids from the Pirabas Formation and
commented on the similarity of the cidarids with the genera from Indo-Pacific
Province.
Brito (1979) studied the Brazilian Cenozoic clyperasteroids and described a
new species from the Pirabas Formation, Clypeaster paraensis. Brito (1981d)
reported a new species of clyperasteroid, Abertella complanata, from the Pirabas
Formation and suggested the presence of Cidaris antarctica (Ortmann, 1900).
Brito (1986) redescribed Abertella complanata in greater detail. Fernandes and
Morais (1994) reported Rhyncholampas candidoi from the Pirabas Formation in
Salinópolis, Pará State. Martinez and Mooi (1997) transfered Karlaster pirabensis
to the genus Abertella.
Morais and Fernandes (1997) recorded for the first time a regular echinoid in
Brazilian Cenozoic rocks. Morais (1998) studied the taxonomy and paleoecology
of regular echinoids from Pirabas Formation.
Collections in the twenty first century resulted in several publications on tax-
onomy, paleoecology and paleobiogeography of the echinoderms fauna from the
Cretaceous and Tertiary Brazilian Basins (Cooke 1955; Seeling et al. 2000; Manso
2001, 2003a, b, 2006; Souza-Lima and Manso 2000; Manso and Souza-Lima
2003a, b, c, 2005, 2007, 2010; Rodovalho et al. 2003; Manso and Hessel 2007;
Manso and Andrade 2008; Manso and Lemos 2008). All this fossil material was
deposited at the Phoenix Paleontological Foundation (Fundação Paleontológica
Phoenix) in Sergipe State.
9.2.3 Echinoderm Physiological Research
Echinoderm physiology has been the focus of intensive research in the last cen-
tury. However, interest in this area of research has decreased significantly in the
last decades.
The first studies on the effect of acetylcholine on holothuroid muscles were
conducted at the Oswaldo Cruz Institute (Rio de Janeiro State) by Moussatché in
1949. Classical studies conducted at the University of São Paulo focused on muscle
biochemistry in holothuroids (Sawaya 1951; Ancona-Lopez 1964) and echinoids
(Sawaya and Ancona-Lopez 1964; Bianconcini et al. 1982, 1985), and on gas
exchange in ophiuroids (Petersen 1976). In the last decade, the physiology of
excretion and ionic regulation has been investigated at the Federal University of
Paraná (Vidolin et al. 2002, 2007; Santos-Gouvea and Freire 2007). The main focus
of this research has been on the interspecific differences in the homeostasis of ions
of the perivisceral coelomic fluid of three echinoids (Echinometra lucunter,
Lytechinus variegatus and Arbacia lixula), one holothuroid (Holothuria grisea) and
one asteroid (Asterina stellifera). Another target of research has been the investi-
gation of how echinoderms deal with the different ions. The current evidence that
echinoderm epithelia treat different ions in distinct ways led to the conclusion that
they are not very homogeneous and identical in their osmo-ionic-conforming
strategies. These studies pointed out that E. lucunter shows reasonable euryhanility
because it can tolerate at least a 30 % reduction in seawater salinity for 5 days;

however it is not able to maintain large ion gradient, especially of Na+ and Cl-, with
respect to external seawater (Freire et al. 2011). On the other hand, L. variegatus is
able to take up Na+ by active transport, showing selective permeability or ion
regulation, somewhat different from A. lixula. Therefore, L. variegatus was termed
as ‘‘hyper-conformer’’ and A. lixula as ‘‘iso-conformer’’ (Vidolin et al. 2007). In
situ experiments with the holothuroid H. grisea and the asteroid A. stellifera
indicated that these intertidal species may have a mechanism to detect exposure to
air, possibly reducing body wall permeability to avoid intense water loss (Vidolin
et al. 2002).
Studies on the immunological response of sea urchins have been conducted at the
University of São Paulo (e.g., Silva et al. 2001; Shimada-Borges et al. 2005),
described later in this chapter. In addition, research on the ecology and biology of
echinoderms yielded many contributions on the function of reproduction and
development (e.g., Alves et al. 2002; Carvalho and Ventura 2002; MacCord and
Ventura 2004; Tavares and Borzone 2006; Cinelli et al. 2007; Lima et al. 2009;
Vellutini and Migotto 2010; Contins and Ventura 2011). For instance, there is a
significant imbalance in sex ratio in a population of the fissiparous sea star Coscin-
asterias tenuispina where no females have been found (Alves et al. 2002). Although
fission occurred throughout the year, it was most frequent in the winter (July, August
and September) when the extremely low tides occur during the daytime.
Echinoderms have also been studied in Brazil in the sister-area of physiology,
biochemistry, and are also common subjects of toxicological studies in environ-
mental agencies. The biochemistry of sulfated oligo- and polysaccharides from
echinoderms with anticoagulant and antithrombotic activities (Mourão et al. 1998)
and the reproductive features of these organic substances (e.g., Cinelli et al. 2007,
2010) have been the subject of intensive research for more than two decades at the
Federal University of Rio de Janeiro. The antithrombotic activities of a polysac-
charide (a fucosylate, chondroitin-sulfate-like polysaccharide) extracted from the
body wall of the holothuroid Holothuria grisea and its chemically modified
derivatives were tested using a static thrombosis model in rabbits. The native
organic extract has an effective antithrombotic action. It prevents thrombosis
completely at a dose of 1.5 mg kg-1 after 10 min stasis, and also, in a dose-
dependent manner, it reduces thrombosis. The anticoagulant or antithrombotic
effects are suppressed by partial defucosylation and desulphation of the polysac-
charide (Mourão et al. 1998). The dynamic of polysaccharide sulfated presented in
the jelly egg coats of the sea urchin E. lucunter shows that: (1) a great amount of
sulfated polysaccharide is present in mature oocytes; (2) a disulfated galactans
(2, 6- disulfated galactan) is produced in the ovary firstly and (3) the 6-sulfate ester
is removed when the polysaccharide concentration increases in the egg jelly coat.
In conclusion, the ovary of E. lucunter synthesizes a specific sulfated galactan
isotype composed of two subunits (2, 6-disulfated units), but its egg jelly coat has
only 2-sulfated residues (Cinelli et al. 2010). These compounds (i.e. sulfated
polysaccharides) are important due to their species specificity in inducing the
acrosome reaction in the fertilization process.
In conclusion, few investigators are devoted to the study of echinoderm

physiology in Brazil. This is a field that should receive more attention in the future.
Physiological knowledge of echinoderm species, both adults and larvae, is
essential for any initiative for the conservation of these invertebrates in a global
changing world.
9.2.4 Echinoderm Immunological Research
Brazilian research with echinoids includes physiological and innate immune

response of the tropical sea urchins L. variegatus and E. lucunter, as well as the
Antarctic Sterechinus neumayeri. General reviews of the innate immune response
in Brazilian echinoderms can be found in Silva et al. (2007) and Tajima et al.
(2007).
The stage at which phagocytosis can first be characterized in the embryos of the
sea urchin L. variegatus was investigated by microinjection of the yeast Saccha-
romyces cerevisiae into its blastocoele. The starting point of phagocytic activity
reflects a biological capacity for distinguishing between self and non-self. Thus the
phagocytosis of yeast by mesenchymal cells beginning in the mid-gastrula stage in
L. variegatus may indicate the moment of acquisition of ‘‘identity’’ (self) in this
organism. Comparative aspects of embryo and adult phagocytes in L. variegatus
have been discussed (Silva 2000). A review on the role of the phagocytes on
embryos was carried out focusing on the following topics: (1) the search for the
origin and first identification of well known adult phagocytes in embryos,
including their role after induced injuries; (2) the search for the occurrence of
phagocytosis in embryos and its role during their physiological development; (3)
the search for phagocytosis in embryos, as a tool to study identity and self rec-
ognition. It is possible to verify that different cell types are able to undertake
phagocytosis under a variety of different stimuli and that the nature of phagocy-
tosis also varies widely (Silva 2002).
Induced inflammatory processes in the sea urchins L. variegatus and E. lucunter
were demonstrated in vitro and in vivo studies using either yeast or India ink
confirming endocytic activity of free ferritin labeled perivisceral phagocytic
amoebocytes (Mangiaterra and Silva 2001; Faria and Silva 2008).
In recent years, studies on effects of oil contamination on sea urchins have been
conducted on Antarctic and tropical species. They showed that crystalloids can be
biomarkers for the analyses of low levels of water-soluble fractions of oil (WSF).
The Antarctic sea urchin S. neumayeri from regions free of pollution was exposed
for 2, 5, 10 and 15 days to different levels of WSF. This study characterized
histological alterations in crystalloids of S. neumayeri that could be used as
biomarkers for oil contaminants with a simple and inexpensive protocol (Borges
et al. 2010).
Global warming is a reality and its effects have been evaluated on the effect of
temperature increase on the innate immune system of the Antarctic sea urchin
S. neumayeri. This study demonstrated the use of red spherule cells as biomarkers
for heat stress because they increase in numbers under acute stress. In addition,
a significant increase in phagocytic indexes was observed at 2 °C coinciding with a
significant increase of iron intranuclear crystalloids at the same temperature.
Moreover, alterations in cell adhesion and spreading were observed. Hsp70
expression was not found in this study and ultrastructural analysis of coelomocytes
did not show any significant difference. This was the first study of the innate
immune response alterations with an increase in temperature (Branco et al. 2011).
Some investigations compared Antarctic echinoderms with tropical species.
One of these studies determined the oxygen consumption in two species of sea
urchins, the tropical L. variegatus and the polar S. neumayeri. In sealed respi-
rometry L. variegatus had a higher consumption of oxygen than S. neumayeri. This
is probably due to diet and temperature. The relation of O:N between S. neumayeri
and L. variegatus was the same, showing that the substrate metabolized of the two
animals is similar (Borges et al. 2012).
Phagocytosis was studied in vitro at 0 °C using the coelomic fluid of the
Antarctic sea star Odontaster validus and the types of amoebocytes involved were
identified. This was the first study of this kind on an Antarctic invertebrate. Rates
of phagocytosis were similar to, or higher than, previously reported for temperate
sea stars. However, this conclusion must be considered with caution because of the
scarcity of data and the different methods used. On the other hand, the data do
suggest that one of the animals’ main defenses against infection may be well
adapted to low temperatures in O. validus. The phagocytosis and germicide
capacity of S. cerevisiae by phagocytic amoebocytes (PA) of the Antarctic sea star
O. validus was also studied in vivo at 0 °C (Silva and Peck 2000; Silva et al.
2001).
9.2.5 Echinoderm Ecotoxicological Research
Bioassays are a valuable approach to evaluate and predict harmful effects of

pollutants on the biota at different levels of organization (Environmental Canada
1992, 2011; Nascimento et al. 2002; Losso and Volpi-Ghirardini 2010). In this
context, echinoderms have a great importance on ecotoxicological and environ-
mental risk assessment studies.
Toxicity tests for environmental quality analyses in Brazil began in the 1970s
with the standardization by the São Paulo State Environmental Agency (Com-
panhia Ambiental do Estado de São Paulo—CETESB) of tests using freshwater
organisms (Resgalla and Laitano 2002; Zagatto 2006). Afterwards, standard pro-
tocols for marine organisms were published (CETESB 1991, 1992, 1999; Resgalla
and Laitano 2002; Zagatto 2006). Since then, toxicity tests with sea urchin
embryos have been used extensively in environmental pollution monitoring
(Prósperi 1993, 2002; Mastroti 1997; Abessa et al. 2003; Nilin et al. 2007).
Among the most common sea urchin species of the Brazilian coast are
E. lucunter and L. variegatus. Both are commonly used in ecotoxicological assays
(Prósperi 1993, 2002; Mastroti 1997; Nascimento et al. 2002; Prósperi and Araújo
2002; Abessa et al. 2003; Nilin et al. 2007; Perina et al. 2011). For instance, Perina
et al. (2011) showed that embryotoxicity experiments using L. variegatus have
shown different effects of two antifouling agents (tributyltin or TBT, and tri-
phenyltin or TPT) and two booster biocides (Irgarol and Diuron). Embryonic-
larval effects of these two antifouling agents were quite different, i.e. development
was interrupted at the blastula and gastrula stages at two TBT concentrations (1.25
and 2.5 lg l-1, respectively) while the pluteus stage was reached at the same
concentrations of TPT. Different ecotoxicological effects were found depending on
concentrations of the two biocides: higher concentrations of Irgarol interrupted
embryonic development at the blastula-gastrula stages while the pluteus stage was
reached under exposure of Diuron at same concentrations. Other studies showed
that embryonic-larval development of E. lucunter and L. variegatus are affected by
chronic toxicity of a chemical mixture of light-stick flags (Cesar-Ribeiro and
Palanch-Hans 2010) and that the chronic toxicity of the sediment in the Guanabara
Bay (Rio de Janeiro) affects the development of L. variegatus (Maranho et al.
2010). Short term chronic toxicity tests with embryos of those species were
standardized by CETESB (1999) and more recently by the Brazilian Association
for Technical Standards (Associação Brasileira de Normas Técnicas—ABNT)
(ABNT 2006, 2012).
Acute and chronic toxicity assays employing aquatic organisms at different life
stages have been extensively used for environmental monitoring programs.
Gametogenesis and embryonic development are some of the aspects that can be
measured. They deserve special attention due to the effects they can cause at the
population and community levels (Cherr et al. 1992, 1993; Kroeker et al. 2010;
Stumpp et al. 2011).
The ecological relevance of echinoderm communities and the vast knowledge
on their embryology make these animals the best choice for laboratory and field
studies on environmental hazards (Bowmer et al. 1986; Briggs and Wessel 2006;
Stumpp et al. 2011). Several species of Echinoidea, Ophiuroidea and Asteroidea
have been used, especially sea urchins, which account for the higher number of
species and assays described and validated by environmental regulatory agencies
(Environmental Canada 1992, 2011; EPA 1995; ABNT 2006, 2012; Nilin et al.
2008). Among the most studied sea urchin species are Paracentrotus lividus,
A. lixula, L. variegatus, E. lucunter and Strongylocentrotus purpuratus.
The use of sea urchin embryos in short-chronic toxicity assays for environ-
mental monitoring and toxicity assessment has been very significant all over the
world. More recently this model proved to be suitable also for the evaluation of the
biological effects of global warming and ocean acidification (Clark et al. 2009;
Dupont et al. 2010; Kroeker et al. 2010; Stumpp et al. 2011). Investigations of the
effects of global warming on development of echinoderms with different devel-
opmental strategies (i.e. planktothrophic and lecitothrophic development) are
currently starting in Brazil (C.R.R. Ventura, pers. comm.). Sea-urchin embryos
have been used also for the evaluation of cytotoxicity, genotoxicity, teratogenicity
and antimitotic activity of natural and synthetic compounds and on the develop-
ment of new medicines and pharmacological tools (Kobayashi 1973; Fusetani
1987; Zúñiga et al. 1995; Saotome et al. 1999; Costa-Lotufo et al. 2002; Hansen
et al. 2003; Cummings and Kavlock 2004). For instance, the potential cytotoxicity
of a diterpene (kaurenoic acid), a natural compound isolated from the wood of a
tropical forest tree (i.e. oleo-resin of Copaifera langsdorffii), shows nonspecific
cytotoxicity. This kaurenoic acid disrupts the embryonic-larval development of
L. variegatus, inhibits tumor cell growth in mammals and causes hemolysis of
mouse and human erythrocytes (Costa-Lotufo et al. 2002).
Many factors contribute for the increasing use of sea urchin embryos as test-
organisms, including their worldwide distribution, their key role on intertidal and
subtidal communities and the ease with which adults of many species can be
collected in their natural habitat. It is also easy to obtain large amounts of ova and
sperm for in vitro fertilization, and the embryonic development is rapid and highly
successful (Nilin el al. 2008). However, for environmental quality assessment
there are some additional factors that have to be considered, such as salinity,
ammonia and sulfite concentrations, pH and temperature of the water (Nipper
2000; Carr et al. 2006).
9.2.6 Brazilian Echinoderm Deep-Sea Research
Echinoderms live in a wide variety of marine habitats, exert a considerable

influence on benthic communities, especially in the deep-sea where they may
represent more than 80 % of the total megafaunal biomass in certain areas, playing
an important role in complex food webs (Haedrich et al. 1980; Sibuet and Law-
rence 1981; Smith and Hamilton 1983; Gage et al. 1983, 1985; Gooday and Turley
1990; Amon and Herndl 1991; Gage and Tyler 1991; Campos et al. 1994; Hendler
et al. 1995; Borges et al. 2002a; Campos et al. 2010).
During the past 23 years, substantial deep-sea research has been carried out in
Brazil (Tavares 1999; Lavrado 2006; Campos et al. 2010). The first records of the
Brazilian bathyal and abyssal fauna were compiled at the end of the nineteenth
century as a result of the Hassler, Challenger and Albatross expeditions (Tavares
1999). In 1987, the French-Brazilian Marion Dufresne (MD55) expedition sam-
pled in the Vitoria-Trindade seamounts region down to a depth of 5,500 m in order
to investigate the faunistic transition from Cabo Frio (where the sub-Antarctic
SACW upwelling occurs) to the Abrolhos slope (Guille and Ramos 1987; Tavares
1999). Several echinoderm species, especially ophiuroids, sea stars and
sea cucumbers, have been sampled during the MD55 campaign (Guille and
Ramos 1987).
Other efforts provided valuable deep-sea material from the Brazilian conti-
nental margin towards the end of the twentieth century. The INTEGRADO Project
sampled from 60 to 600 m depth at São Paulo State coast, and later the Evaluation
Program of the Sustainable Yield of Living Resources from the Brazilian Exclu-
sive Economic Zone (Programa de Avaliação do Potencial Sustentável de
Recursos Vivos na Zona Econômica Exclusiva—REVIZEE) sampled down to
800 m in the South region and deeper than 1,000 m at the Southeast one (Pires-
Vanin 1993; Lana et al. 1996; Sumida and Pires-Vanin 1997; Amaral et al. 2004;
Ventura et al. 2006). Diversity of echinoderm fauna from the Brazilian South-
eastern coast is higher between depths of 20 and 100 m and decreases gradually up
to 400 m. However, higher frequencies of species are found between depths of 400
and 600 m and diversity increases to similar values at depths of 100–200 m
(Ventura et al. 2006). Borges et al. (2002b, 2004a, b) explored ophiuroid diversity
and Ventura et al. (2006) expanded the list to other echinoderm classes found at
depths down to 2,000 m. Most biodiversity data originated from the South and
Southeast upper slope, some stations from the lower slope and a few from the
abyss, although a few records were also established in the Northeast (Sumida and
Pires-Vanin 1997; Amaral et al. 2004; Amaral and Rossi-Wongtschowski 2004;
Lavrado 2006; Lavrado and Brasil 2010; Lavrado et al. 2010a, b; Amaral and
Jablonsky 2005; Capítoli and Bemvenuti 2006; Capítoli and Bonilha 2010).
Studies on the effect of anthropogenic activities at the Brazilian continental
margin have enhanced our knowledge on benthic communities, especially at
Campos Basin down to nearly 3,000 m (Santos et al. 2009; Campos et al. 2010;
Lavrado et al. 2010a, b). Echinoderms have been one of the most abundant
invertebrate groups sampled during the megabenthic campaigns at Campos Basin
(Rio de Janeiro State), representing approximately 45 % of the total megafauna
(Lavrado et al. 2010a, b). During February (summer) and August (winter) 2003,
4,392 specimens were collected by trawling. The Ophiuroidea were the most
abundant amongst the echinoderms (n = 3,281) followed by the Asteroidea
(n = 936) and Holothuroidea (n = 158). The Echinoidea were very rare (n = 17),
and no Crinoidea were collected. The abundance of ophiuroids and asteroids was
higher at a depth of 1,100 m, and had the tendency to decrease with increasing
depth. But this essentially happened because of the presence of a few dominant
species from both classes [Ophiuroidea (Ophiura ljungmani, Ophiophycis sp.1)
and Asteroidea (Plutonaster bifrons and Nymphaster arenatus)]. The higher
expected species values were found at a depth of 1,300 m.
According to Campos et al. (2010), 39 species (Appendix) were identified at
Campos Basin (CB) during the Campos Basin Deep-Sea Environmental Project/
PETROBRAS. (Caracterização Ambiental de Águas Profundas a Bacia de Cam-
pos/CENPES/PETROBRAS). Amongst the ophiuroids, the species Asteroschema
(A. arenosum), Histampica (H. duplicata), Ophiuraster (Ophiuraster sp.),
Ophiosphalma (O. armigerum), Ophiacantha (O. aristata, O. metallacta and
O. anomala) were new records for Brazilian waters. Eight asteroid species from
five families (Astropectinidae, Benthopectinidae, Goniasteridae, Zoroasteridae and
Brisingidae) were identified: Astropecten sp. 1; Plutonaster bifrons; Cheiraster
(Cheiraster) sepitus; Ceramaster sp. 1; Nymphaster arenatus; Pseudarchaster
gracilis gracilis; Zoroaster fulgens; and Brisinga sp. 1. Pseudarchaster gracilis
gracilis and Cheiraster (Cheiraster) sepitus were new records for Brazil (Campos
et al. 2010). Sea cucumbers represented 3.6 % of the total echinoderms collected.
Most were identified to species level, except for a morpho-species classified as
Synallactidae morphotype 1. The most abundant species were Mesothuria verrilli,
Molpadia blakei, Deima validum validum and Molpadiodemas porphyrus.
Barboza et al. (2011) used metadata from several echinoderm studies and data
from the Brazilian continental margin in order to evaluate the potential relation-
ship between the faunas from South America and the Antarctic region, taking into
account the proximity of both continents and the fact that they were the last to
separated from Gondwana around 34 million years ago. These authors recorded
602 echinoderm species from both regions, 101 of them (*17 %) were shared
between the continents. Around 47 % of these are typically from the deep-sea,
most deep-sea records being from Brazil.
More recently, other wide spread programs have been carried out by the energy
companies in different regions off the Brazilian continental margin. Many new
echinoderm records are expected in the near future. Taking into account the length
of the Brazilean coastline (*8,000 km), and the fact that around 70 % of its
exclusive economic zone, defined between 19.3 km (12 NM) and 321.8 km (200
NM) off the coast, is within the slope and abyss zones (Miloslavich et al. 2011),
much work is still needed in order to enhance our knowledge on Brazilian deep sea
echinoderms.
9.2.7 Ecology and reproduction
Ecological studies of echinoderms from the Brazilian shallow waters concern

spatial distribution patterns (Borzone 1991; Ventura and Fernandes 1995; San-
chez-Jérez et al. 2001; MacCord and Duarte 2002; Capítoli and Benvenutti 2006),
population size structure (Ventura and Fernandes 1995; Junqueira et al. 1997),
feeding biology (Lima-Verde and Matthews 1969; Ventura et al. 1994; 2001; Calil
et al. 2009) and ecological association (Ribeiro et al. 2003; Clavico et al. 2006).
MacCord and Duarte (2002) described the dispersion pattern and the size
structure of the crinoid Tropiometra carinata along the southeastern coast of
Brazil (São Sebastião Channel, São Paulo). This species shows a clumped distri-
bution (especially for smaller individuals), with a preference for attachment to the
lateral surface of rocks. Adults are predominant in rocky environment and recruits
among algae. The size structure of the population was stable throughout one year.
Regarding sea stars, Ventura and Fernandes (1995) studied the bathymetrical
distribution and population size structure of paxillosid species in the upwelling
environment of the continental shelf of Cabo Frio. Five species were found but
three were mostly frequent for three years: Astropecten cingulatus, A. brasiliensis
and Luidia ludwigi scotti. Their bathymetric distribution in this soft-bottom
environment is very distinct, according to the grain size of the sediments:
A. brasiliensis occurs at a depth of 30 m (from very well to well-sorted medium
sand) and a depth of 45 m (from well to moderated-well sorted medium sand);
A. cingulatus and L. ludwigi scotti are most frequent at a depth of 60 m (from well
to moderated-well sorted fine sand) and also at a depth of 45 m. Analysis of
population size structure of these three species showed that individuals of the same
size range did not occur at the same depth, especially for Astropecten species.
Studies on feeding biology are more frequent for paxillosid sea stars, particu-
larly the genera Astropecten and Luidia because these species show intra-oral
digestion and the stomach content analysis provides reliable information about
their diet. In general, studies have shown that species of Astropecten (A. cingulatus,
A. marginatus and A. brasiliensis) and Luidia (L. senegalensisi and L. ludwigi
scotti) play a wide range of trophic roles in soft-bottom environments along the
Brazilian coast. They can be non-selective deposit feeders (feeding on organic
matter from sediments), non-selective predators (feeding on the most frequent prey
in the environment, generally mollusks and crustaceans) and also selective pre-
dators (showing preferences for some species of prey, especially when there are
competitors like other sea stars). Ventura et al. (2001) reported that A. brasiliensis
and A. cingulatus can shift their food niche breadth when they occur together. Also
specimens of both species of same size range have broader food niche at depths
where they do not occur together.
Calil et al. (2009) carried out field experiments to test the effectiveness of
predation of the sea star Asterina stellifera in a rocky-shore environment in the
south of Brazil. They increased the density of A. stellifera in order to evaluate its
predation pressure on the community. Their results showed that this starfish has a
small effect on the barnacle population (Chthamalus bisinuatus) and has a minimal
effect on this rocky-shore community of a subtropical region, in contrast to other
asteroid species.
Studies on reproduction of echinoderms, especially of common sea urchins and
sea stars species from Brazilian shallow waters, have been carried out. Regarding
echinoids, in general, continuous gametogenesis occurs throughout the year, but in
some cases a long spawning period of three or four months has been detected. That
is the case for E. lucunter, whose spawning time occurs from December to June in
populations from northeastern, southeastern and southern coasts (Tavares et al.
2004; Ventura et al. 2004; Lima et al. 2009).
9.3 Echinoderm biodiversity
9.3.1 Distribution Patterns and Biogeographic Remarks
Echinoderm fauna in Brazilian waters is composed essentially of species with

three principal distribution patterns: 1) species ranging from North–South Carolina
(USA) or from Caribbean to Argentine; 2) species ranging from North–South
Carolina or Florida (USA) to Cabo Frio or São Paulo (Southeastern Brazilian
coast); and 3) species ranging from Cabo Frio to the Rio de la Plata mouth
(Uruguay/Argentine). However, very few studies on biogeography of echinoderms
have been done in Brazil, and even in South America or worldwide (Ghiold and
Hoffman 1986; Price et al. 1999; O’Hara and Poore 2000; Martinez 2008). This is
probably due to the relatively low number of researchers dedicated to this kind of
study, particularly in Brazil, and lack of reliable information on taxonomy and
distribution ranges. Although information on the occurrence and distribution of
echinoderm species in Brazil still seems to be insufficient regarding the great
length of the country’s coastline. New records and new species have been
described recently as emphasized later in this chapter.
These three patterns of distribution are much correlated with the hydrographical
features (i.e. marine currents and water masses) of the Brazilian coast, as described
above (see Introduction). Changes in species composition occur markedly at the
Cabo Frio region, where upwelling often brings sub-Antarctic water (the South
Atlantic Central Water, SAWC) to shallow depths. Therefore, there are features of
a temperate environment at a such usually tropical latitude (22°S), which could
explain the distribution pattern of some species from Cabo Frio to Uruguay (Rio de
la Plata) and the occurrence of tropical and subtropical fauna at Cabo Frio, such as
the sea stars Astropecten cingulatus (tropical), Astropecten brasiliensis (subtrop-
ical), Echinaster (Othilia) guyanensis (tropical), Echinaster (Othilia) brasiliensis
(subtropical), Asterina stellifera (subtropical) and the sea urchin Paracentrotus
gaimardi (subtropical).
Historically, the South Atlantic Ocean was formed during the Cretaceous
(ca. 135 My) when South America was separating from Africa and Antarctica/
Australia (the so-called Gondwana paleo-continent). Therefore, species composi-
tion on the Brazilian continental shelf should be similar to those at the margins of
Africa and Antarctica/Australia and different from the marine fauna of north of the
South America (‘‘Tethyan’’), because there is no marine connection between the
south and the north regions of the Atlantic (Floeter et al. 2009). South America and
Africa became effectively separated by a deep-sea barrier during the Paleocene (ca.
60 My). About the same time, a connection between the marine biotas of Brazil and
the Caribbean was formed. During the Eocene (ca. 50 My), Australia and Antarctic
became separated and also the latter and South America. Each step of the history of
South American west coast might mark the evolution of the Brazilian marine biota,
at least in shallow water (Floeter et al. 2009). In general, this seems to be true for
echinoderms and other marine organisms because the three patterns of species
distribution are similar for different taxa. Furthermore, the echinoderm fauna of
Brazilian waters is currently very similar to those of the Caribbean, as for other
groups of marine invertebrates. This similarity may also be due to the effects of
relatively recent dispersal capacity and colonization, as well as ancient extinctions
of species at the Brazilian continental shelf.
Comparing the number of species described worldwide and for Brazilian
waters, one can notice that there are not as many species of Brazilian echinoderms
as expected from the great length of the country’s coastline. This is probably due
to the low number of researchers dedicated to the study of echinoderms in Brazil.
Therefore, information on the occurrence and distribution of echinoderm species
in Brazil is insufficient.
There are about 700 extant species of Crinoidea, of which only 16 have been
reported for Brazilian waters. Of the 1,800 extant species of Asteroidea, only 77
reported for Brazil. Of the 2,000 extant species of Ophiuroidea, 153 have been
reported for the Brazilian coast. Of the 900 extant species of Echinoidea, 52 have
been reported for the Brazilian coast. Of the approximately 1,250 extant species of
Holothuroidea, 49 reported for Brazil.
9.3.2 Crinoidea, Asteroidea and Echinoidea
The remarkable low diversity of echinoderms of Brazilian waters may be due to

the low number of researchers who have been dedicated to the study of the
phylum. Considering the great length of the country’s coastline, information on the
occurrence and distribution of extant echinoderm species in Brazil seems to be
insufficient, especially for crinoids, asteroids and echinoids. Little information
about earlier studies on crinoids, asteroids and echinoids are available. However,
some information about ophiuroids and holothuroids are emphasized on the fol-
lowing topics.
9.3.3 Ophiuroidea
The first report about ophiuroids in Brazil appeared in the 17th century, when
Marcgrave (1942) recorded a brittle star, probably from the genus Gorgonocephalus
Leach, 1815. Ljungman (1867) established the genus Amphipholis based on the
species Amphipholis januarii from the coast of Rio de Janeiro State. Several other
studies were carried out in Brazil, mainly on taxonomy and geographical distribu-
tion (Brito 1960e; Tommasi 1965a, b, 1967, 1970b, c, d, 1971a, b, c, d; Lima-Verde
1969; Manso 1988a, b, 1989; Absalão 1990). Tommasi (1970b, c, d) published a
survey including 65 species of ophiuroids known to that time. Up to the end of the
1980s, eight new species from the Brazilian coast were described and new occur-
rences were recorded (Tommasi 1971a, b, c, d, Tommasi and Abreu 1974; Tommasi
and Oliveira 1976; Albuquerque 1986; Monteiro 1987; Manso 1988a, b, c). In 1999
Tommasi published an identification key for 98 species and 38 genera of Brazilian
ophiuroids.
More recently, several other studies were carried out in different regions of
Brazil. Alves and Cerqueira (2000) investigated the echinoderms found on beaches
of Salvador (Bahia State), recording 15 species of ophiuroids. Albuquerque et al.
(2001) described two new species of the family Amphiuridae: Amphiura callida
and Amphiodia habilis. Borges et al. (2002a), in a study about the ophiuroids of the
continental shelf off southern and southeastern Brazil, recorded 25 species,
including two new records, at depths down to 800 m. Manso (2004) recorded 10
species of ophiuroids in Camamu Bay (Bahia State), at depths down to 50 m.
Ventura et al. (2006) recorded the occurrence of the family Asteronychidae and
reported a total of 15 species from the coast of Rio de Janeiro to Bahia States from
depths down to 1,800 m. Borges et al. (2006) described the species Ophiomisidium
tommasii (Ophiuridae) collected at depths down to 800 m, and Borges and Amaral
(2007) recorded four new occurrences for Brazil, including the first record of the
family Ophiochitonidae. Manso et al. (2008) collected 23 species at Todos os
Santos Bay and Aratú Bay (Bahia State), at depths down to 62 m. Gondim et al.
(2008) recorded 13 species at Cabo Branco Beach (Paraíba State). Barboza (2010)
and Gondim et al. (2010) recorded for the first time the occurrence of the genus
Gorgonocephalus (G. chilensis) and the species Ophionereis dolabriformis,
respectively.
At the present time, 153 species have been recorded (Appendix), despite the
fact that knowledge on Brazilian Ophiuroidea, especially from the continental
margin and the deep sea, is yet quite scarce (Tommasi 1999).
Ophiuroids were one of the most abundant macrofauna groups collected at
different bathymetric ranges, from the intertidal down to a depth of 1,600 m.
Especially in regions deeper than 200 m, new species and many new records of
occurrence have been reported (Borges et al. 2006, Borges and Amaral 2007,
Campos et al. 2010).
One of the most recent programs of faunal survey was REVIZEE. Between
Cabo de São Tomé (Rio de Janeiro State) and Arroio Chuí (Rio Grande do Sul
State), more than 16,000 Ophiuroidea specimens were collected between depths of
60 and 810 m with 32 species recorded (Amaral et al. 2004). On the Biota Marine
Benthos project (Biodiversidade Marinha do Estado de São Paulo), which sampled
from the intertidal down to a depth of 50 m, 29 species of Ophiuroidea were
recorded and approximately 2,500 specimens were collected (Borges et al. 2011).
Species of the family Amphiuridae predominate in waters shallower than 50 m,
although banks of ophiuroids are scarcely found at this bathymetric range. These
studies recorded herds of ophiuroids deeper than 500 m for at least three species of
the family Ophiuridae: Ophimisidium tommasii, Ophiura clemens and Ophiura
ljungmani (Monteiro 1987, 1990; Tommasi et al. 1988a, b; Absalão 1990; Sumida
1994; Heitor 1996; Petti 1997; Capítoli and Monteiro 2000; Borges 2001).
Although the high diversity of Amphiuridae in shallow and deeper waters, pop-
ulation densities are low when compared to those of Ophiuridae in deeper regions.
Some genera, such as Ophiura, Ophiomusium, Ophiomastus, and Ophiomisidium,
are dominant in these regions.
On the inner continental shelf, Hemipholis elongata (Ophiactidae), an infaunal
and gregarious species, was frequently found together with other species such as
Microphiopholis atra and Amphiodia planispina, from the beach down to a depth
of about 50 m, similarly to the findings reported by Hendler et al. (1995) in the
Caribbean region. These authors stated that the species occur frequently on poorly
oxygenated unconsolidated bottoms, but also on oyster banks and coral reefs. Boffi
(1972) recorded only young individuals of H. elongata in the phytal of the
northern coast of São Paulo State, and reported that adults probably live prefer-
entially on sublittoral soft bottoms.
Several studies in the southeastern and southern regions stated that Amphipholis
squamata, Ophiactis lymani, Ophiactis savignyi, and Ophiothrix angulata occur-
red associated with algae, on rocky shores, and in the unconsolidated sublittoral.
According to Chao and Tsai (1995), these species preferentially inhabit biological
substrates such as sponges, algae, and corals. Morgado and Tanaka (2001)
recorded them associated with colonies of the bryozoan Schizoporella errata, on
the Brazilian southeastern coast. Boffi (1972) analyzed the ophiuroids associated
with 23 species of macroalgae on the northern coast of São Paulo State, and found
a predominance of Ophiactis lymani and Amphipholis squamata, which occurred
on almost all the algae and were the most frequent and highest-density species.
According to Boffi (1972), Ophiactis savignyi and Ophiothrix angulata prefer-
entially inhabit sponges, although not exclusively. Ophiothrix angulata is a fre-
quent species in shallow depths on the Brazilian coast, and may extend to a depth
of about 540 m. Amphipholis squamata is a cosmopolitan and well-studied species
that inhabits several types of substrates, from sand to crevices, macroalgae,
sponges, corals, and aggregations of polychaetes. Boffi (1972) reported that this
species occurred from the intertidal zone to a depth of 5 m.
Epifaunal species such as Ophioplocus januarii and Ophioderma januarii,
collected in the unconsolidated sublittoral, such as muddy, sandy, and gravel
bottoms, are common along the northern coast of São Paulo State from the
intertidal to depths of 180 and 100 m, respectively (Monteiro et al. 1992).
Ophioplocus januarii occurs in the transition zone between the inner and outer
continental shelf along the São Paulo State coast (Monteiro et al. 1992; Pires-
Vanin 1993).
Among the Amphiuridae, some species were most commonly collected during
the REVIZEE/South Score and Biota/FAPESP programs, such as A. squamata
(coast and algae), Microphiopholis atra and Amphiura joubini (unconsolidated
sublittoral). These last two are typically infaunal species, with morphological
adaptations, such as a soft disc and long slender arms, to live in unconsolidated
bottoms. Microphiopholis atra is common along the Brazilian southeastern coast,
in depths to 100 m. Amphiura joubini, although frequent on the southeastern-
southern coast, is a subantarctic species with a wide bathymetric distribution (from
shallow water to a depth of 3,000 m). Its occurrence in this region is possibly a
result of the penetration of colder water masses, such as the South Atlantic Central
Water (SACW) (Tommasi 1985). According to Pires-Vanin (1993), A. joubini is
characteristic of an environmental gradient in the transition zone between the inner
and outer continental shelf. Also, Amphiura flexuosa is present in the zone of
environmental transition between the inner and outer shelf, as recorded by findings
of REVIZEE and Biota/FAPESP programs.
Capítoli and Monteiro (2000) reported that Hemipholis elongata, Amphiodia
planispina, Ophiactis lymani, and Amphioplus lucyae form an association of
shallow-water species (\30 m), along the extreme southern coast of Brazil. On the
other hand, Amphiura joubini and Amphiura complanata form an association
between depths of 30 and 100 m.
Species of the family Ophiuridae are most commonly collected above a depth
of 100 m. Sumida and Pires-Vanin (1997) reported that Ophiura ljungmani has a
wide distribution, although it occurs preferentially at depths above 100 m.
According to these authors, there is a clear relationship between depth and species
composition, which changes significantly at depths between 300 and 500 m off
southeastern Brazil. At the shallower depths, changes in species composition occur
more gradually. This differentiation in species composition along the bathymetric
gradient was observed by several investigators on the southeastern coast of Brazil,
for different taxonomic groups such as Ophiuroidea (Borges 2001), Mollusca
(Miyaji 2001; Quast 2003; Arruda 2005), and benthic macrofauna in general
(Sumida and Pires-Vanin 1997; Amaral et al. 2004). Borges (2001), analyzing the
ophiuroids of the REVIZEE/South Score program, found a group of species at
depths to 100 m, another between 100 and 300 m, and a third below 300 m.
Several studies in the southeastern and southern continental shelf of Brazil
(Monteiro 1987; Pires-Vanin 1993; Sumida 1994; Petti 1997; Borges 2001)
highlighted some groups of species of Ophiuroidea frequently found: (1) species
such as O. lymani, O. savignyi, A. squamata and O. angulata that are common
along the coast, especially ocuring associated with other invertebrate (sponges and
octocorals); (2) species such as O. januarii, H. elongata, M. atra, and Ophio-
phragmus luetkeni that occur at depths between 5 and 50 m; (3) species such as
Amphiura flexuosa, A. joubini and Ophiothrix rathbuni that occur below a depth of
30 m, although more frequently near depths of 100–200 m; and (4) species of
greater depths ([200 m), such as Ophiomastus satelitae, Ophiura clemens,
O. ljungmani, Ophiomusium acuferum, Ophiomusium eburneum, Ophiostriatus
striatus and Ophiomisidium tommasii.
9.3.4 Holothuroidea
Data on Brazilian Holothuroidea appeared at the beginning of the nineteenth

century, published by European and North American scientists who identified
echinoderms from the Brazilian coastal waters collected on scientific expeditions
(Tiago and Ditadi 2001). However, this group has been the most neglected among
the echinoderms and even intertidal species known for over 40 years remain
poorly studied.
According to Clark (1907), Selenka’s monograph (1867) was the pioneering
study of the anatomy and modern systematics of the Holothuroidea. Selenka listed
212 species known to that time, 49 of which were new to science. One year later
Selenka (1868) added seven other species to his list, although none of the 219
species were reported for Brazilian waters. Indeed, the first record of Brazilian
Holothuroidea is in Verrill (1868). He published an article on the Radiata
Collection sampled by the geologist Charles Frederick Hartt, who accompanied
Louis Agassiz on his expedition to Brazil. Hartt explored the coral reefs at Santa
Cruz and Porto Seguro region (Bahia State) and, among the marine specimens he
sampled, were two species of sea cucumbers. One of them was Thyone brazili-
ensis, described by Verril as Thyone (Sclerodactyla) braziliensis. The other one
was Chiridota rotifera, recorded for the first time for Brazil by Verril as Chirodota
rotiferum.
Later Rathbun (1879) provided the first list of Brazilian echinoderms, with four
species of sea cucumbers, 11 species of sea urchins, 13 species of sea stars, 32
species of brittle stars, and four species of sea lilies (Tiago and Ditadi 2001). He
gathered all the species collected by the Geological Commission of Brazil from
1875 to 1877. He also added the species previously recorded by other scientists,
like Ljungman and Fritz Müller. When he recorded the geographic distribution of
the sea star Echinaster brasiliensis, he stressed that the term ‘‘Rio de Janeiro’’ was
commonly used to designate Brazil in general at that time, rendering some con-
fusion about the distribution of many species. The Holothuroidea he recorded
were: C. rotifera (as C. rotiferum), T. braziliensis (as T. (Sclerodactyla) brazili-
ensis), Thyonella sp. and Holothuria sp.
In 1881, Ludwig examined the animals collected at the Brazilian coast by
E. van Beneden, recording the occurrence of C. rotifera and describing two new
species of Holothuroidea: Phyllophorus parvus (as Thyonidium parvum) and
Protankyra benedeni (as Synapta benedeni). In 1882, he also published a list of the
echinoderms collected by E. van Beneden at the Brazilian coast (Tommasi 1957).
The specimens collected by the Challenger Expedition along the coast of Bahia
State were studied by Théel (1882, 1886a), who described Thyone pervicax,
Pentacta pygmaeus (as Colochirus pygmaeus), and Lissothuria braziliensis (as
Psolus braziliensis).
Ludwig (1881) and Théel (1886a, b) contributed by describing new species and
findings of sea cucumbers in Brazil. However, it was only in 1897 that Hermann
von Ihering listed the first marine invertebrates, among them echinoderms, for the
coast of the State of São Paulo, specifically at São Sebastião Island. However,
Ihering did not identify a single species. That task was performed by Luederwaldt
(1929a, b), who cited the occurrence of Holothuria grisea at that island (Tiago and
Ditadi 2001).
Deichmann (1930) described Holothuroidea sampled over a huge area, from
southern Brazil to Cape Cod (Massachussetts, USA). She gave detailed informa-
tion about the 103 species found, with 12 reported for the Brazilian coast:
Holothuria grisea, Phyllophorus seguroensis, P. occidentalis, P. parvus, Thyone
cognita, T. suspecta, T. pervicax, T. belli, Pentacta pygmaeus, Thyonepsolus
braziliensis, Synaptula hydriformis and Protankyra benedeni.
Although mentioned by earlier authors such as Verrill (1868), Ludwig (1881),
and Théel (1886a, b), the Apoda Chiridota rotifera was not reported for Brazil in
Deichmann’s monograph (Tiago and Ditadi 2001). Later on Luederwaldt (1929a,
b) recorded the occurrence of H. grisea at São Sebastião Island (São Paulo State).
Systematic research on Holothuroidea by Brazilian biologists is somewhat

recent, beginning with Ancona-Lopez (1957), on specimens sampled at Pernam-
buco, Rio de Janeiro and São Paulo States. In her publication she listed H. grisea,
Stichopus badionotus, Thyone braziliensis, Chiridota rotifera and Synaptula
secreta. Tommasi (1957) also published a list of Holothuroidea sampled down to a
depth of 15 m along the coast of São Paulo State, including Synaptula hydriformis,
H. grisea and S. badionotus.
In 1969, Tommasi also published a list with 31 species of Holothuroidea, as
well as a survey of one hundred years of the literature about the group (Tommasi,
1969a). Nine new species were added to the Brazilian fauna in the following thirty
years (Tiago and Ditadi 2001).
Other authors also published lists on the Brazilian Holothuroidea, among them
Sluiter (1910), Krau (1950), Heding and Panning (1954), Ancona-Lopez and
Sawaya (1955), Ancona-Lopez (1962, 1964, 1965a, b), Tommasi (1959, 1969a, b,
1971b, d, 1972, 1974a), Brito (1960f, 1962), Deichmann (1963), Lima-Verde
(1969), Freitas et al. (1973), Tommasi and Oliveira (1976), Tommasi and Aron
(1988), Tommasi et al. (1988a,b), Freire and Grohmann (1989), Hendler et al.
(1995), Hadel (1997), Pires-Vanin et al. (1997), Hadel et al. (1999) and Brites et al.
(2011).
Our knowledge of the southeastern Atlantic Holothuroidea is poor, particularly
for deep-sea species. Hyman (1955) observed that ‘‘interest in this group appears
relegated to taxonomic specialists’’, while Hendler et al. (1995) pointed out that
holothurians are far from being attractive animals. Besides, their densities in the
Atlantic waters are low compared to the tropical Indo-Pacific. Moreover, the
Atlantic species are generally small, dull in colour and are not as easily seen as are
coral reef’s species.
From Verrill (1868) to the present, 49 valid names of Holothuroidea species
have been ascribed to the 7,367 km of the Brazilian coast (see Appendix). At first
this may seem to indicate a low occurrence of sea cucumbers on the Brazilian
coast, but it properly reflects the lack of systematic studies of these echinoderms
by Brazilian biologists (Tiago and Ditadi 2001).
Species with a wide geographical distribution predominate among the Brazilian
Holothuroidea, with most species reaching at least to the Caribbean. Endemic
species are also well-represented: Psolus vitoriae, Thyone montoucheti,
Euthyonidiella dentata, Cucumaria manoelina, Ocnus braziliensis, Trachytione
crassipeda, Leptosynapta brasiliensis and Synaptula secreta (Ancona-Lopez 1957;
Tiago and Ditadi 2001).
Although São Sebastião (São Paulo State) is recognized as the southern limit of
the geographical distribution of many Brazilian sea cucumbers, some species, like
I. badionotus and H. grisea, have been found at Santa Catarina State, some 750 km
south of São Sebastião (Tiago and Ditadi 2001). However, the number of species
described for the Brazilian littoral, as well as their geographical distribution,
should be reviewed taking into account possible synonymies.
Although the collection of echinoderms for human consumption is illegal (Hadel

et al. 1999), aquaculture enterprises to culture Brazilian echinoderms are absent in
the country (Campos and Lima 2000; Ventura et al. 2006). This may be attributed
to a cultural heritage from the first colonizers, mainly of Portuguese and Spanish
origins. In the first centuries of European occupation of the Brazilian territory, the
main purpose was to collect as much natural resources as possible and take them
back to their countries of origin. This attitude towards nature still persists and the
only marine and brackish animals cultured commercially are shrimp. Therefore,
despite their high economic value on foreign markets, echinoderms are still
ignored in Brazil as a potential food resource (Campos and Lima 2000).
Nineteen of the 627 threatened species listed in the Brazilian Red List (Machado
et al. 2008) are echinoderms: fourteen species of Asteroidea (Asterina stellifera,
Astropecten brasiliensis, A. cingulatus, A. marginatus, Coscinasterias tenuispina,
Echinaster (Othilia) brasiliensis, E. (Othilia) echinophorus, E. (Othilia) guyan-
ensis, Linckia guildingii, Luidia clathrata, L. ludwig scotti, L. senegalensis,
Narcissia trigonaria, Oreaster reticulatus), three species of Echinoidea (Cassid-
ulus mitis, Eucidaris tribuloides, Paracentrotus gaimardi) and two species of
Holothuroidea (Isostichopus badionotus, Synaptula secreta). Most of these species
live in shallow waters and are easily collected.
Machado et al. (2005, 2008) pointed out several threats that menace Brazilian
echinoderms, among them habitat destruction or alteration, eutrophication, and
erosion. Anthropogenic impacts are caused by the increased occupation of coastal
regions. Landfills, for instance, cause sediment transport to shallow sea waters,
changing the physical and chemical properties of the substrate. The sea lily
Tropiometra carinata, for instance, was once abundant on the exposed rocky
shores of the coast of São Paulo State. Due to anthropogenic impacts, however,
population densities of these animals are decreasing steadily (Hadel et al. 1999).
Oil spills and discharge of domestic or industrial sewage are also side effects of
coastal municipaltie’s development (Machado et al. 2008).
The impact caused by touristic enterprises, including diving, is recognized as
threats to several species of benthic animals, including sea stars, sea urchins and
crinoids (Machado et al. 2008).
Accidental by-catch by trawling boats fishing for shrimp is also listed as one of
the major menaces for echinoderms, especially for asteroids and echinoids that live
on sandy and muddy bottoms (e.g. A. brasiliensis, A. cingulatus. A. marginatus,
L. clathrata, L. ludwigi scotti, L. senegalensis and Encope emarginata). The
number of benthic species accidentally captured by trawl-nets with doors may
amount to 96 %, with A. marginatus always present in the nets (Branco 1999;

Branco and Verani 2006a, b). Fishing gear also damages the animals indirectly as
they pass over them and disturb the substrate physically (Micael et al. 2009). In the
southeast region, fishing of sea-bob-shrimp (Xiphopenaeus kroyeri) with bottom
trawl nets results in heavy by-catch due to its low selectivity (Coelho et al. 1986;
Branco 1999; Branco and Fracasso 2004; Carniel 2008; Bail et al. 2009).
According to Bail et al. (2009), 0.04 kg of echinoderm fauna are captured by trawl
nets for each kg of sea-bob-shrimp. Branco (1999) estimates that 7.13 ton d-1 of
by-catch fauna, including fishes, crustaceans, mollusks and echinoderms are
captured on the southern Brazilian coast, amounting to 1,711.7 ton y-1.
Either caught or not by the nets, echinoderms may be severely damaged by
them. Among the damage listed by Micael et al. (2009) are: injuries caused by
demersal trawls passing over the bottom, damage or autotomy of body parts due to
physical contact with fishing gear, interactions with other species in the catch or
handling by the crew, crushing when dropped on to the deck, punctures in the
epidermis, and bacterial infection due to trauma. Echinoderms are menaced also
by the impacts caused by trawls and dredges by scraping or resuspension of the
substratum (Jennings and Kaiser 1998).
Sea urchins and sea cucumbers have been fished commercially all over the
world (Micael et al. 2009) but in Brazil it is illegal to collect native fauna except
fishes, mollusks and crustaceans captured for human consumption. Even for
research projects, special permission from environmental agencies is needed.
Predatory collection for aquarium trade, souvenirs, and decorative pieces, food
markets, and other commercial purposes are threats listed for several species of
Brazilian echinoderms in addition to those listed by Machado et al. (2008),
including the sea star Asterina stellifera, the sea urchins L. variegatus and
E. lucunter and the sea cucumber H. grisea. Sea stars, O. reticulatus for instance,
is illegally and extensively collected all over the Brazilian coast and sold as
decorative pieces, souvenirs or religious objects used in some local rituals (Hadel
et al. 1999; Campos and Lima 2000). Alves et al. (2006, 2009/2010) created the
term zoohandicraft to designate workmanship products made of marine macro-
fauna. When interviewed by the authors, the craftsmen claimed no knowledge on
the threats the animals were under. Along the northeast coast of Brazil, Alves et al.
(2009/2010) described zoohandicraft production as the only or the main economic
source of many people with strong familiar hereditary tradition.
Because it is illegal to collect echinoderms, except for research purposes with a
special permission, the capture of those animals occurs in an artisanal way. This is
particularly true for sea urchins whose mature female gonads, known as roe, are
prepared as a delicacy. Although not menaced at present, it is known that con-
siderable numbers of E. lucunter, L. variegatus and P. gaimardi have been
collected by hand during low tides or by divers at localities like Ilhabela Island
(São Paulo State). Because there are no external differences between males and
females, and between mature and immature females, a large number of individuals
are killed before the consumers find the desired specimens, resulting in high levels
of mortality. Moreover, killing of mature females, the ones that would produce
new generations, endangers even more those species’ populations. These threats
apply also to the sea cucumbers I. badionotus and H. grisea. Those echinoderms
are reported as a food source for some local populations on southern, southeastern
and northeastern Brazilian coasts (Hadel et al. 1999; Campos and Lima 2000;
Mendes et al. 2006).
About 17 % of the animals in the aquarium global trade are echinoderms
(Micael et al. 2009). Among the species commonly found in Brazilian home
aquaria, and at public or private ones, are A. stellifera, E. (Othilia) brasiliensis,
O. reticulatus, E. tribuloides and I. badionotus. Most of those animals die before
they reach the stores. Even after that, most of them survive only for short periods
and have to be replaced. Therefore, more and more specimens have been collected,
endangering even more the species already menaced by other threats.
Some echinoderm are threatened by low density populations, limited or dis-
continuous distribution, or because they occur on vulnerable environments. The
endemic sea cucumber Synaptula secreta was described for one locality only, at
Segredo Beach (São Sebastião, São Paulo State). There are no data on the
reproductive strategy of this species, but its distribution pattern suggests that they
brood their young in the coelom, like other Apoda species such as Synaptula
hydriformis and Chiridota rotifera. The brooding sea urchin C. mitis, that keeps
their short-term larvae on the female’s body, is also threatened by their low dis-
persion capacity and, in consequence, restricted distribution (Amaral et al. 2008).
The sea stars L. guildingii and C. tenuispina have discontinuous distribution
patterns and low population densities, many of which probably persist only by
asexual reproduction. This reproductive strategy decreases genetic variability and
gene flow among populations, rendering them even more vulnerable to environ-
mental impacts.
Besides the threats already stated here there are others worth mentioning. As
benthic animals echinoderms are susceptible to pollutants stored in marine sedi-
ments and to biomagnification processes (Camargo 1982; Hadel et al. 1999;
Carnevali et al. 2001; Sugni et al. 2007; Micael et al. 2009). Some species, like
L. variegatus and E. lucunter, have been used for ecotoxicological tests according
to the standard protocol ABNT-NBR 15350 (Brazilian Association for Technical
Standards; ABNT 2006, 2012) because their embryos are a reliable source of
material for biochemical analyses (Micael et al. 2009). Arbacia lixula, another sea
urchin species, has been suggested also as potential test organism for ecotoxico-
logical analysis by Máximo et al. (2008). Although the fact that they are not
regarded as severely threatened species, the use of their embryos and larvae as
experimental tools is leading to an increasing depletion of wild populations in
some regions along the Brazilian coast.
Studies on the reproductive biology, population dynamics and genetics of most
species are needed as well as on the taxonomy of those with high morphological
variations, such as Echinaster species and P. gaimardi. In order to achieve such
knowledge, which depends on long term studies, it is essential to protect these
species through governmental laws that restrict and control collection.
9.6 Concluding Remarks
Although research on echinoderms from Brazilian waters is still scarce, current

advances in several fields of knowledge are remarkable. The principal bottlenecks
seem to be the current low number of researchers dedicated to the phylum and the
lack of integrative actions to promote information exchange and collaborative
studies. However, there has been a relevant increase in the number of researchers
working in different subject areas during the last two decades.
In fact, an optimistic feeling is corroborated by some evidences. For instance,
although there are few taxonomists in Brazil, the zoological collections were
improved during the last ten years due to some governmental programs of bio-
diversity inventory and protective laws that obligate petroleum companies to
provide environmental analyses before any exploitation. These facts have induced
the needs for more qualified people, which stimulated the academic institutions to
have more students working on projects of biodiversity, ecotoxicology and
physiology in their Post-graduate Programs. This interest has also generated
increasing studies on immunology, biochemistry, ecology, reproduction and
development of echinoderms.
In addition, the Brazilian Symposium of Echinoderms has convened twice in
the past five years (the first in 2008 and the second in 2012). The second Latin
American meeting on echinoderms (II Congreso Latinoamericano de Equinoder-
mos, CLE) will take place in Brazil in 2013. These are important initiatives to
enhance exchange of knowledge and to stimulate collaborative studies.
In summary, it is well-known that much time and effort are necessary to train
new researchers and to produce knowledge, but some robust steps have taken place
as described earlier in this chapter. Echinoderm studies in Brazil have to be
stimulated more and integrative research has to be encouraged. The state of
echinoderm knowledge seems to have an exciting and successful future in Brazil,
like in the whole Latin America.
Acknowledgments We are grateful to the editors of this volume (Juan José Alvarado and
Francisco Solís Marín) for the invitation to take part in this volume. We are also indebted to John
M Lawrence and the editors for their critical comments and suggestions that improved this
chapter.
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Chapter 10
Echinoderm Research in Uruguay
Sergio Martínez
10.1 Introduction
The eastern 220 km of the Uruguayan coastline are considered marine (salinities
around 30–35 %), and the remaining ca. 350 km correspond to the Río de la Plata
estuary. The Río de la Plata is the second largest river system in South America
(drainage basin: 3.1 9 106 km2, sediment load: 91 9 106 t y-1, discharge of an
average of 22,000 m3s-1 water to the ocean) (Framiñan and Brown 1996;
Guerrero et al. 1997; Syvitski et al. 2005). Along the coast there is a remarkable
salinity gradient, with changing fronts, caused by the discharge of fresh water from
the Paraná and Uruguay rivers into the Río de la Plata and from it into the Atlantic
Ocean (Fig. 10.1a). Winds change the location of the salinity and turbidity fronts
(Larrañaga 1894; Nagy et al. 1987). The Río de la Plata is extremely muddy
because of mud derived mainly by the Paraná River.
The Atlantic area is influenced by the warm coastal N–S Brazilian Current and
by the cold S–N Malvinas (Falkland) Current (Boltovskoy 1966; Podestá et al.
1991; Lentini et al. 2000; Piola et al. 2000). On the platform there are superficial
warm waters and deeper cold ones (Fig. 10.1b). The sub-tropical warm waters
converge and mix with the cold subantartic waters at about the isobaths of 100 and
200 m. Other factors such as topography, seasonality, and El Niño-Southern
Oscillation (ENSO) phenomena also contribute to the littoral environmental
conditions (Olson et al. 1988; Podestá et al. 1991; Lentini et al. 2000; Ortega and
Martínez 2007; Raicich 2008). In brief, the estuary and the adjunct platform have
complex horizontal and vertical structures complicated by a high degree of
S. Martínez (&)
Departamento Evolución de Cuencas, Facultad de Ciencias,
Iguá 4225, 11400, Montevideo, Uruguay
e-mail: smart@fcien.edu.uy

Springer-Verlag Berlin Heidelberg 2013
346 S. Martínez
Fig. 10.1 a Examples of distribution of salinity (%) in the Río de la Plata and adjacent platform
(from Ortega and Martínez 2007). b Examples of distribution of temperature (C) in the Río de la
Plata and adjacent platform (from Ortega and Martínez 2007)
seasonal and inter annual variability. The substrate is dominated by soft sediments,
with few areas consolidated or constituted by crystalline rocks (Correia et al.
1996).
The Uruguayan Atlantic coast shows arched (sometimes rather straight), wave
dominated sandy beaches of variable size, delimitated by rocky points (Fig. 10.2).
They have a wide range of morphodynamic grades, from dissipative to reflective.
Coastal morphology is conditioned more by its geological background than by the
present dynamics (Brazeiro et al. 2003; Gómez Pivel 2006). Astronomic tides are
between 0.4 and 0.6 m, sea level changes induced by the wind being more
important (Wells and Daborn 1997; SOHMA 2002).
Jurisdictional issues are regulated by the ‘‘Treaty between Uruguay and
Argentina concerning the Río de la Plata and the Corresponding Maritime
Boundary’’, signed November 19th 1973 (English version can be consulted in UN
2002, Spanish one in FREPLATA 2011). Apart from the Río de la Plata and
Atlantic coastal boundaries, it establishes a ‘‘common fishing zone, beyond 12
nautical miles measured from the corresponding coastal baselines, for duly reg-
istered vessels flying their flag. Such zone shall be that determined by two arcs of
circumferences of a radius of 200 nautical miles, the centers of which are located
at Punta del Este (Eastern Republic of Uruguay) and Punta Rasa del Cabo San
10 Echinoderm Research in Uruguay 347
Fig. 10.2 Examples of coastal morphology of Uruguay. a Lagoon, rocky point, and rather
straight beaches at each side. b Rocky points and arched beaches in between
Fig. 10.3 Political limits of

Uruguayan waters (modified
from FREPLATA)
Antonio (Argentine Republic) respectively’’ (Fig. 10.3). It also provides a ‘‘Joint

Technical Commission …which shall be responsible for conducting studies and
adopting and co-coordinating plans and measures for the conservation, preserva-
tion and rational exploitation of living resources and the protection of the marine
environment in the common zone binational organisms…’’
Uruguay has a National System of Protected Areas (SNAP). In the Atlantic
zone of influence, there are two coastal marine protected areas (from the terrestial
coast seaward some miles: Laguna de Rocha and Cabo Polonio). Two additional
areas will be formalized soon (Laguna Garzón and Cerro Verde). There are not
exclusively marine Protected Areas (SNAP 2010).
348 S. Martínez
In such a variable and unpredictable context as described above, it is not

surprising that recent echinoderm faunas are rather poor. But, as mentioned below,
the lack of research must be taken into account. Fossil echinoderms, although also
not diverse, include key representatives of the extinct echinoid family Mono-
phorasteridae, the sister group of the abundant and widespread Mellitidae. The aim
of this chapter is give an overview of the knowledge about extinct and extant
echinoderms of Uruguay.
10.2 Research
Echinoderm research is not yet systematically developed in Uruguay. There is

little knowledge of the modern fauna. Evidence of this situation is the few papers
that have been devoted to Uruguayan echinoderms.
Although Mortensen (1917) mentioned the presence of Isometra vivipara in
Uruguay, the given coordinates (338000 S; 518100 W) correspond to Brazilian
waters.
Barattini (1938) mentioned one ophiuroid species (Ophioceramis januarii,
presently Ophioplocus januarii), two asteroid species (Asterina marginata, today
classified as Asterina stellifera, and Astropecten brasiliensis) and two echinoid
species (Mellita quinquesperforata and Encope emarginata; Fig. 10.4b). The same
species are included in Barattini and Ureta (1961), a book for the general public.
A similar book by Klappenbach and Scarabino (1969) mentions only the sand
dollar species.
Bernasconi (1941a, b, 1947, 1953, 1955, 1966) and Bernasconi and D’Agostino
(1977) made important contributions to the knowledge of the echinoderms of the
Southwestern Atlantic, adding to the Uruguayan fauna one asteroid species
(Astropecten cingulatus; Fig. 10.4a), three ophiuroid species and an echinoid spe-
cies. This material was included in papers focused on museum collections or about
the fauna collected in vessel campaigns at a regional scale (e.g. ‘‘Vema’’ from the
Lamont Geological Observatory). Bernasconi (1964) include the species of asteroids
and echinoids of Southern South America, including the Uruguayan species, to
already known biogeographical entities (e.g. Argentine and Magellan provinces).
In the framework of a faunistic survey, Milstein et al. (1976) added five
ophiuroid species to the Uruguayan fauna, and indicated some already known
asteroid and echinoid species. Also in a report of a faunistic survey and ecologic
study, Juanicó and Rodríguez-Moyano (1976) added three asteroid and two regular
echinoid species to the Uruguayan fauna. Parts of these papers include environ-
mental data.
Tommasi (1970a, b, 1974) and Tommasi et al. (1988a, b) published monographs
or detailed surveys concerning Brazilian fauna, but included some Uruguayan
species found near the borderline between the two countries. In fact, the only
Uruguayan holothuroids identified at the species level can be found in these works.
Fig. 10.4 a Astropecten cingulatus, dorsal view. b A. cingulatus ventral view. c Amphiodia sp.,
dorsal view (from Carranza et al. 2007). d Encope emarginata, adapical view. e E. emarginata,
oral view. f Amphiodia sp., ventral view (from Carranza et al. 2007). Specimens of A. cingulatus
and E. emarginata from S. Martínez’s personal collection
Lucchi (1985) identified 17 species of ophiuroids in samples obtained between

10 and 800 m. (continental shelf and slope), being five of them reported for the
first time for Uruguay. He and Martínez (2008, with an updated database and
including asteroids) made some biogeographical considerations about ophiuroids
(see below).
Martínez and Mooi (2005) published a bibliographical review about the fossil
and recent sand dollars of Uruguay, recognizing three recent species, two
Quaternary, and four Miocene ones. They stressed the importance of the extinct
Miocene family Monophorasteridae in the phylogeny of Mellitidae and included
some biogeographical observations (see Sect. 3.1).
Scarabino (2006) presents an updated list of the Uruguayan echinoderms
reported to a depth of 50 m. He provided a brief account of the characteristics of
their habitats. There is only one new record (Carranza et al. 2007) of Amphiodia
sp. (Fig. 10.4c), an unnamed ophiuroid species different from the other Uruguayan
representatives of the genus.
Amaro (1974), Riestra et al. (1992, 1998), Roux and Bremec (1996), Obenat
et al. (2001), Giberto et al. (2004), Bremec and Giberto (2004), or Demichelli and
Scarabino (2006) have considered Uruguayan echinoderms, adding new localities
or habitat characteristics, but without adding new taxa.
350 S. Martínez
Table 10.1 Numbers of echinoderms per taxonomic group in Uruguay

Crinoidea Asteroidea Ophiuroidea Echinoidea Holothuroidea Total
Order 0 3 2 3 3 11
Family 0 4 7 3 3 17
Genus 0 4 15 5 3 27
Species 0 5 24 5 3 37
Puig (1986) found three unidentified holothuroid species and three species of
Ophiuroidea in the stomach content of the fish Micropogonias opercularis.
Fossil material has been more intensively studied (Martínez 1984, 1994;
Martínez and Durham 1988; Mooi et al. 2000; Martínez and Mooi 2005). It is
relatively well known but is restricted almost exclusively to Cenozoic echinoids
because of their best preservation potential and of the characteristics of the sedi-
mentary units of Uruguay.
10.3.1 Extant Echinoderm Fauna
The echinoderm fauna of Uruguay (Table 10.1) is composed of 37 species.

Ophiuroidea is by far the richest class with 24 species. Asteroidea has five species;
Echinoidea five species; and Holothuroidea three species. There are no species of
Crinoidea recorded for Uruguayan waters. Is this species proportion among classes
reliable? As commented below, there has been no systematic sampling of the
Uruguayan waters, so this question remains open.
Twenty-four of these species have been collected from a depth of less than
50 m, four from between 50 and 100 m, two from between 100 and 200 m, four
from 400 m, two from 700–800 m, and one at a wide range of depth (Amphiura
eugeniae, 46–800 m).
Considering an average depth of 200 m for the shelf break, there are thus only
seven species beyond the shelf. Again, it is not clear if this distributional pattern,
with species concentrated in the littoral, is reliable, or if it is an artifact caused by
the proximity of the coast (easier to collect), and/or by the fact that some
collections are a side consequence of commercial fishing (more intense at these
depths). Research-oriented faunistic surveys of the Uruguayan waters have been
scarce, except for recent years and the collections have not been studied.
Biogeographic considerations have been few. Tommasi (1970a, b) indicated the
marine area surrounding the mouth of the Río de la Plata (Rio Grande do Sul State,
Brazil, to Buenos Aires Province, Argentina) is an area with reduced diversity.
He related this to the latitudinal decrease of temperature, decrease in salinity
caused by the discharge of the Río de la Plata, and the relatively cold water mass
along the coasts of Rio Grande do Sul.
Lucchi (1985) identified two groups within the ophiuroids she studied, one
related to the Malvinas (Falkland) Current and living at a depth greater than 60 m,
and the other related to warm waters and living at less a depth than 60 m.
Martínez (2008) identified two distributional patterns of ophiouroid and aster-
oid species. The first one comprises species with the southern end of their dis-
tributional range near Uruguay. This was divided in two sub-groups, depending on
whether their distribution extended across the Río de la Plata or not. Species whose
distribution is limited by the Río de la Plata, are affected by two main factors in
this area: temperature variation of the Subtropical Confluence Zone (ca. 308 to
458S, see Boltovskoy 1966 for example), where the cold Malvinas (Falkland) and
the warm Brazilian currents meet, and the strong variation in salinity and the
increase of turbidity of waters caused by fresh water discharge from the Rio de la
Plata (Piola et al. 2000, González-Silvera et al. 2004). The ‘‘salinity barrier’’ of the
Río de La Plata has been considered an explanation for the truncation of the
distribution of other organisms at the Rio de la Plata area, such as mollusks
(Scarabino 1977) or sand dollars (Martínez and Mooi 2005). The second group is
composed of species distributed within the area of the Argentinean or Patagonian
Province (ca. 438S and ca. 288S; Scarabino 1977; Boschi 2000), a unit charac-
terized by a complex faunal composition. Tommasi (1970b; ophiuroids), Scara-
bino (1977; mollusks) and Martínez and del Río (2002; mollusks) suggested this
zone may be a true transition zone or ‘‘Provinciatone’’ between the adjacent
Magellanic and Brazilian Provinces.
The distribution of sand dollars was studied by Martínez and Mooi (2005) with
results and explanations comparable to the sub-groups of asterozoans that do not
cross the Rio de la Plata barrier.
10.3.2 Extinct Echinoderm Fauna
There are only eight species of echinoderms in the Uruguayan fossil record, one
crinoid species, one ophiuroid species, and six echinoid species. The crinoid and
ophiuroid species are from the Devonian.The echinoid species are from the
Cenozoic, being abundant in the Late Miocene. During those times (ca. 10 Ma.
ago) the Monophorasteridae (sister group of the Mellitidae) were particularly
abundant and diverse in what are today Uruguay and Argentina. Two species
(Mellita sp. and Encope sp.) are recorded only for the Quaternary and are still
extant in the area.
The following taxonomic arrangement of the fossil species. Suprageneric
classification is based on: Crinoidea, Ausich (1998); Holothuroidea, Pawson et al.
(2010); Ophiuroidea, Smith et al. (1995) and Shackleton (2005); Echinoidea: Kroh
and Smith (2010).
Class Crinoidea J.S.Miller, 1821
Subclass Camerata Wachsmuth and Springer, 1885
352 S. Martínez
Fig. 10.5 Miocene sand dollars. a Monophoraster duboisi (x 1.3), FCDP 2359; b Amplaster
coloniensis (x 0.9); holotype MNA-CPO 3426; c Amplaster ellipticus (x 0.9), paratype MNA-
CPO 3425; d Amplaster alatus (x 1.3), FCDP 2831. From Martínez and Mooi (2005)
Order Monobathrida Moore and Laudon, 1943

Family Melocrinitidae d’Orbigny, 1852
Genus Ctenocrinus Bronn, 1840
Ctenocrinus sp.
References: Daners (1990).

Age and Stratigraphic Unit: Early Devonian, Cordobés Fm.
Comments: identified on the basis of columnar plates.
Class Ophiuroidea Gray, 1840
Family Encrinasteridae Schuchert, 1914
Genus Encrinaster Haeckel, 1866
Encrinaster pontis (Clarke, 1913)
References: Méndez-Alzola (1938).

Age and Stratigraphic Unit: Early Devonian, Cordobés Fm.
Comments: a single disk. The specimen is lost.
Class Echinoidea Leske, 1778
Order Clypeasteroida L. Agassiz, 1835
Crown group Clypeasteroida
Crown group Scutelliformes
Family Monophorasteridae Lahille, 1896
Genus Monophoraster Lambert and Thiéry, 1921
Monophoraster duboisi (Cotteau, 1884)
(Fig. 10.5a)
References: Martínez (1994), Mooi et al. (2000).

Age and Stratigrapic unit: Late Miocene, Camacho Fm.
Comments: Martínez and Mooi (2005) stated. ‘‘Prior to these records, all
authors (e. g., Goso and Bossi 1966, Figueiras and Broggi 1971) mentioned the
presence of Monophoraster darwini in Uruguay. In fact, this species is not present
in the Miocene of Uruguay and all the known specimens are referable to
M. duboisi. The species was originally described from the contemporaneous Pa-
raná Formation of Argentina (Cotteau 1884, Mooi et al. 2000).’’
Genus Amplaster Martínez, 1984
Amplaster coloniensis Martínez, 1984
(Fig. 10.5b)
References: Martínez (1984, 1994) (partim), Martínez and Durham (1988),

Mooi et al. (2000).
Comments: This species is endemic to Uruguay.
Amplaster ellipticus Mooi, Martínez and Parma, 2000
(Fig. 10.5c)
References: Mooi et al. (2000).

Comments: This species is endemic to Uruguay.
Amplaster alatus (Rossi de García and Levy, 1989)
(Fig. 10.5d)
References: Mooi et al. (2000).

Comments: This species was described from the ‘‘Patagoniense’’ strata from
Chubut Province (Rossi de García and Levy 1989), but with a controversial age
assignment ranging from? Middle Eocene to the? Late Oligocene-Early Miocene
(see Mooi et al. 2000).
Family Mellitidae Stefanini, 1912
Genus Mellita L. Agassiz, 1841
Mellita sp.
References: Figueiras (1962), Figueiras and Broggi (1967); Martínez and Mooi
(2005).
Age and Stratigrapic unit: Holocene, Villa Soriano Formation.
Genus Encope L. Agassiz, 1841
Encope sp.
References: Martínez and Mooi (2005)

Age and Stratigrapic unit: Quaternary.
354 S. Martínez
There are no echinoderm fisheries or aquaculture currently in Uruguay (see in

example Defeo et al. 2009). The effect of fishing of other species (fishes, shrimps)
on echinoderm populations has not been evaluated.
10.5 Threats
For most of the Uruguayan coast, baseline studies include echinoderms only
incidentally. Hence, it is unknown if species are being affected by anthropogenic
activity (e.g., fishing, coastal urbanization), or if the echinoderms are affecting the
environment to some extent.
There are not Uruguayan echinoderms considered to be charismatic or
bioengineers.
10.6 Recommendations and Concluding Remarks
As shown in this chapter, knowledge of the Uruguayan echinoderm fauna is very

poor. Therefore, the priority is to increase substantially all types of research on this
group. A reliable inventory of the species must be developed, including basic data
such as georeferences, depth, salinity, temperature, sedimentological characteris-
tics, and other species associated with the echinoderms. The lack of national
experts on the group undoubtedly has retarded the progress of the knowledge of
echinoderms in Uruguay.
Acknowledgments Juan José Alvarado kindly invited me to participate in this book and, along
with three anonymous reviewers, made important suggestions on earlier versions of the manu-
script. Alejandro Brazeiro provided useful information about the National System of Protected
Areas.
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Chapter 11
Echinoderms from Argentina:
Biodiversity, Distribution and Current
State of Knowledge
Martín I. Brogger, Damián G. Gil, Tamara Rubilar, Mariano I.

Martinez, María Enriqueta Díaz de Vivar, Mariana Escolar,
Lucía Epherra, Analía F. Pérez and Alejandro Tablado
Martín I. Brogger, Damián G. Gil, Tamara Rubilar These authors contributed equally to this
work.
M. I. Brogger (&) M. I. Martinez A. Tablado

Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’-CONICET,
Buenos Aires, Argentina
e-mail: mbrogger@bg.fcen.uba.ar, mmartinez@macn.gov.ar, tablado@macn.gov.ar
D. G. Gil
Instituto de Desarrollo Costero, Universidad Nacional
de la Patagonia San Juan Bosco, Comodoro Rivadavia, Argentina
e-mail: gil_damian@hotmail.com
T. Rubilar L. Epherra
Centro Nacional Patagónico-CONICET, Puerto Madryn, Argentina
e-mail: rubilar@cenpat.edu.arepherra@cenpat.edu.ar
L. Epherra
e-mail: epherra@cenpat.edu.ar
M. E. Díaz de Vivar
Facultad de Ciencias Naturales, Universidad Nacional
de la Patagonia San Juan Bosco, Puerto Madryn, Argentina
e-mail: mediazdevivar@gmail.com
M. Escolar
Instituto Nacional de Investigación y Desarrollo Pesquero,
Mar del Plata, Argentina
e-mail: marianaescolar@inidep.edu.ar
A. F. Pérez
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires,
e-mail: analiafperez1@gmail.com

360 M. I. Brogger et al.
11.1 Introduction
Echinoderm research in Argentina began with Professor Irene Bernasconi

(1896–1989; Fig. 11.1), at the Argentine Museum of Natural Sciences ‘‘Bernardino
Rivadavia’’ (MACN), who dedicated 55 years of her life to study the echinoderms
inhabiting the Argentine Sea. Her first studies were focused on Echinoidea of Tierra
del Fuego (54°S, 68°W) and Ophiuroidea and Crinoidea of South Georgia islands.
In 1924, she published the first work with pictures and drawings of the most
common sea urchins (Bernasconi 1924). In 1935, she published her first taxonomic
publication about Asteroidea describing new species of the genus Pteraster
(Bernasconi 1935). From that date, Asteroidea became the focus of her research,
describing two new species of the genus Luidia (e.g. Luidia patriae) (Bernasconi
1941a). She revised the taxonomy of seven families: Pterasteridae (Bernasconi
1937), Luidiidae (Bernasconi 1943), Odontasteridae (Bernasconi 1962a), Gonias-
teridae (Bernasconi 1963), Ganeriidae (Bernasconi 1964a), Asterinidae (Bernas-
coni 1973a) and Echinasteridae (Bernasconi 1980). The 1960’s were the most
productive of her life. She published 15 papers (e.g. Bernasconi 1961, 1962a, b,
1963, 1964a, b, c, 1965a, b, c, 1966a, b), described one new genus (Vemaster;
Bernasconi 1965a) and four new species (e.g. Vemaster sudatlanticus). In 1964, she
published the first key for orders, families and genera of Asteroidea from the
Argentine Sea (Bernasconi 1964b). Although her major contribution was with sea
stars, she also made significant advances in taxonomic research of sea urchins (e.g.
Bernasconi 1953) and brittle stars (e.g. Bernasconi and D’Agostino 1977).
Irene Bernasconi was a pioneer in echinoderm research in Argentina. At the age
of 72 she travelled on an expedition to Antarctica (1968–1969), being one of the
first Argentinean female scientists to explore the white continent (Fig. 11.1). Her
work was and still is invaluable for all the research on echinoderms in Argentina.
She retired in 1984 and passed away in 1989. We would like to dedicate this
chapter to Professor Irene Bernasconi, the first echinoderm specialist in our
country.
This chapter reviews all knowledge of Argentine echinoderms up to the present
time, available in published papers, meetings, and Licentiate and PhD theses
focused on this Phylum. Several studies that focus on Antarctic echinoderms are
beyond the scope of this chapter, which is restricted to the Argentine Sea (from the
Río de la Plata to Tierra del Fuego and Malvinas Islands). Current PhD projects
and soon to be published data about chemical, biological, ecological and taxo-
nomic research conducted by the authors and other colleagues are not included in
this chapter. Information about depth range, habitat and taxonomic accounts of the
species is limited to published information.
11 Echinoderms from Argentina 361
Fig. 11.1 Professor Irene Bernasconi, on an expedition to Antarctica (1968–69) and working at
the Museum (MACN)
11.1.1 Oceanography and Marine Environments
The Argentine Continental Shelf (ACS; Fig. 11.2) is one of the largest and flattest
in the world. This shelf extends from Río de la Plata (RP) (36°000 S, 56°110 W) to
Tierra del Fuego (TF) (54°380 S, 64°450 W) (Martos and Piccolo 1988). The ACS
has several major coastal embayments: Bahía Blanca bay (BBB) (38°550 S,
62°120 W), San Matías gulf (SMG) (41°290 S, 64°340 W), San José gulf (SJG)
(42°190 S, 64°210 W) and Nuevo gulf (NG) (42°780 S, 64°950 W), gulfs which border
the headland of Península Valdés (PV) (42°260 S, 63°320 W), and San Jorge gulf
(SJOG) (46°190 S, 66°150 W). The ACS (ca. 1,000,000 km2) is an exceptionally
large Neritic Province (Acha et al. 2004). The width of the ACS varies between a
few kilometers at 55°S to 850 km along 51°S (Martos and Piccolo 1988). The
principal circulation is characterized by the confluence of the cold north-flowing
Malvinas Current and the warm south-flowing Brazil Current (Garzoli and Garraffo
1989; Bastida et al. 1992; Brandini et al. 2000; Acha et al. 2004; Lucas et al. 2005).
The Malvinas Current is an energetic north-flowing branch of the Subantarctic
Cabo de Hornos Current that affects both coastal and offshore areas. As it moves
farther north, it affects only offshore waters (Garzoli and Garraffo 1989; Bastida
et al. 1992). The Malvinas Current has seasonal variations, with temperature and
Fig. 11.2 The Argentine Continental Shelf (ACS). Scheme of major upper level currents,
Malvinas and Brazil currents and the Brazil-Malvinas confluence (BMC) (adapted from Brandini
et al. 2000). It shows the Río de la Plata (RP), San Matías gulf (SMG), San José gulf (SJG),
Nuevo gulf (NG), San Jorge gulf (SJOG), the Malvinas Islands (MI) and Tierra del Fuego (TF)
salinity ranging seasonally from 4 to 11 °C and from 33.8 to 34.4 %, respectively

(Bastida et al. 1992). On the other hand, the Brazil Current is a warm oligotrophic
south-flowing branch of the South Equatorial Current (Garzoli and Garraffo 1989;
Bastida et al. 1992). Nutrient concentration increases gradually from north to
south, with lowest values in the subtropical region coincident with the lowest value
of chlorophyll-a (Brandini et al. 2000).
The Brazil and the Malvinas Currents converge along the continental margin of
South America within latitudes 35°–45°S as the Brazil-Malvinas confluence (BMC)
(Garzoli and Garraffo 1989; García et al. 2004). The front shows a substantial
spatial and temporal variation forming a large area of intense mixing of subtropical
and subantarctic waters (Garzoli and Garraffo 1989; Brandini et al. 2000; Rivas
et al. 2006). Maximum annual values of chlorophyll-a at the ACS occur in spring
and summer, with minimum values in winter (Rivas et al. 2006). This high primary
productivity supports a great number of fisheries in the Southwestern Atlantic
Ocean (Bremec et al. 2000; Schejter et al. 2002; Acha et al. 2004).
In order to characterize the different coastal regions, we will describe the
physical oceanographic features of the coastal fronts (Fig. 11.2).
The Río de la Plata estuary is an extensive and shallow coastal plain estuary at
35°–36°S that receives freshwater from two major rivers (Paraná and Uruguay).
The system is characterized by strong vertical stratification. Freshwater flows
seaward on the surface while denser shelf water intrudes along the bottom, taking
the shape of a salt wedge (Guerrero et al. 1997; Acha et al. 2004; Lucas et al.
2005). Due to lack of osmoregulation in echinoderms, this frontal zone might
represent an important geographic barrier for the distribution of coastal echino-
derms. Many echinoderm species are distributed off RP estuary, but only in denser
and deeper shelf waters of the front.
The front at the coastal zone of El Rincón (39°–41°S, depth \ 40 m) encloses
an area of 10,000 km2, showing weak seasonality. The front is characterized by
vertical homogeneity due to tidal forcing. A coastal front separates diluted coastal
water, from Negro and Colorado rivers, and shelf waters. Salinity gradients
increase on the continental shelf due to high saline waters originated in SMG
(Guerrero and Piola 1997; Acha et al. 2004; Lucas et al. 2005).
The Península Valdés tidal mixing front is a mesoscale (100–1,000 km) thermal
front. The front is observed in spring and summer, which establishes the boundary
between stratified waters offshore and a coastal vertically mixed body of water (Acha
et al. 2004). The structure of the front is maintained until autumn when stratification
of shelf waters decays. The front begins to form in spring (Acha et al. 2004).
Rivas and Ripa (1989) analyzed the temperature and salinity of the Nuevo gulf.
They observed no stratification of temperature during winter. However, this
condition was not observed during summer when a layer of warmed water over-
layed cooler, deeper water. San José gulf is influenced by winds and tidal currents.
Therefore, for a great part of the year, this gulf does not show stratification. Only a
weak vertical structure occurs during summer (Rivas 1990).
Off southern Patagonia, water masses on the northern extreme of the Drake
Passage are diluted. This dilution is caused by an excess of rainfall in the SE Pacific
and the continental discharge along the west coast of South America. The flow
towards the Atlantic, known as Cabo de Hornos Current, enters onto the continental
shelf and contributes to lowering the salinity of the Magellan Strait. Several basins
drain along the strait, collecting abundant rainfall during summer and discharging
further diluted waters into the ACS. A subsequent diluted plume is then traced 200 km
off-shore (100 m depth) and 800 km northward. The plume reaches the southern limit
of SJOG and the coastal tidal front (Krepper and Rivas 1979; Acha et al. 2004).
Near the ACS break, the Subantarctic shelf waters of the Malvinas Current
produce an important thermohaline front (Martos and Piccolo 1988; Lutz and
Carreto 1991). The shelf-break front is a permanent feature that characterizes the
border of the shelf. This front has mild gradients in density controlled mainly by
temperature in summer and by salinity in winter. The dynamics of the Malvinas
Current affects the latitudinal and seasonal variation of the front, moving offshore
during summer and onshore during spring and autumn (Olson et al. 1988; Fedulov
et al. 1990; Carreto et al. 1995; Acha et al. 2004). The front extends from the
Burwood Bank along the shelf-break and around Malvinas Islands (Guerrero et al.
1999) to the Brazil/Malvinas confluence in the North, producing a large region of
interrelated fronts (Acha et al. 2004).
The ACS, particularly the Patagonian shelf, is identified as a high primary
productivity area (Rivas et al. 2006). The ecology and reproduction of echino-
derms inhabiting the coastal, shelf and shelf-break region of the ACS are affected
by variations of the primary productivity and oceanographic variables (e.g.
Brogger et al. 2004, 2010; Rubilar et al. 2005, 2008a; Marzinelli et al. 2006; Gil
and Zaixso 2007; Martinez 2008; Pérez et al. 2008; Gil et al. 2009).
The ACS is characterized by a homogeneous layer of unimodal fine and
medium size sand, which is the main component of the sediment. In some areas,
this layer is up to 2 m thick (Urien and Ewing 1974; Bastida and Urien 1981).
Although some places show a high percentage of gravel, other areas associated
with estuaries, gulfs and bays have fine sediments (mud-sand). Hard-bottom areas
with rocky shores seem to be scarce (Bastida et al. 1992).
The fauna of the Argentine shelf can be separated into two biogeographic
provinces: the Argentine and the Magellanic. These are part of the Subtropical and
Subantarctic Regions (Boschi 1979; Balech and Ehrlich 2008). The Argentine
province is a warm-temperate subregion extending from the littoral of Buenos
Aires southward to the coastal waters of northern Chubut (43–44°S). The depths of
the external boundaries range between 40 and 60 m. Water temperature varies
latitudinally and seasonally from 8 to 23 °C (Boschi 1979). This subregion also
reaches northward to Cabo Frio, (Brazil; 23°S, 42°W) having seasonal fluctuations
in the southern area. On the other hand, the Magellanic province is a cold-tem-
perate subregion. At latitudes of 44–45°S it leaves the coastal region following the
Malvinas Current, upon reaching 37°S. This province extends about 150-200 km
from the continent, with depths greater than 80 m. The northern boundary of this
subregion can be tentatively placed at 34–35°S. Water temperature varies sea-
sonally from 4 to 16 °C. This subregion extends to the Pacific Ocean along the
southern coastline of Chile (Boschi 1979).
11.2 Diversity and Distribution of Echinoderms
11.2.1 Expeditions to the Argentine Sea
The first campaign in the Argentine Sea was the Eugenie (1851–1853), which
explored the Río de la Plata and the Magellan Strait. From the collections of this
expedition, Ljungman (1867) studied the ophiuroids and Ekman (1925) studied the
holothuroids. The Hassler Expedition (1871–1872) explored the Brazilian coast

southward to the Magellan Strait. Lyman (1875) described the ophiuroids and
A. Agassiz (1873) the echinoids of this expedition. One of the most important
expeditions was that of the Challenger (1872–1876), which established knowledge
of the deep sea echinoderms, as well as other deep sea groups in the region. The
echinoderms from this famous expedition were described by A. Agassiz (1881;
Echinoidea), Lyman (1882; Ophiuroidea), Théel (1882, 1886; Holothuroidea),
Carpenter (1884, 1888; Crinoidea), and Sladen (1889; Asteroidea). From 1874 to
1876 the Gazelle explored the same region. From the collections of this expedition,
Studer examined the ophiuroids (1882), asteroids (1883) and echinoids (1880),
while Lampert (1889) examined the holothuroids. The Albatross (1888–1904)
explored a great part of the world, including the Argentina Sea. Agassiz (1904)
studied the echinoids, Ohshima (1915) studied the holothuroids and Clark (1915,
1947) the crinoids.
The vessels Vettor Pisani and Alert, the expedition Hamburger Magalhaensi-
sche Sammelreise (1892–1893), and the Mission scientifique du Cap Horn
(1882–1883) explored the southern region around the Magellan Strait, the coast of
Patagonia and the Beagle Channel. The ophiuroids and holothuroids were studied
by Ludwig (1886, 1898, 1899) and echinoderms in general by Bell (1881) and
Perrier (1891).
Between 1897 and 1910, Antarctic expeditions from different countries
explored the Argentine waters and Subantarctic Islands. The Belgian Antarctic
Expedition 1897–1899, the Scottish National Antarctic Expedition on the SY
Scotia 1903–1904, the Swedish Antarctic Expedition 1901–1903, the German
Antarctic Expedition 1901–1903. Koehler (1902, 1908, 1923) studied the ophiu-
roids, Hérouard (1906), Ekman (1925) and Vaney (1909) the holothuroids, An-
dersson (1904), Mortensen (1918), Vaney and John (1939) the crinoids, Ludwig
(1903) and Koehler (1923) the asteroids, and Koehler (1902) and Mortensen
(1920a) the echinoids. The French Antarctic expeditions during 1903–1905 (on the
Français vessel) and during 1908–1910 (on the Pourquoi-Pas? IV vessel) col-
lected invertebrates. The holothuroids from this campaign were identified by
Vaney (1906, 1914). Two British campaigns: RV Scoresby (1926–1936) and RV
Discovery (1927–1932) explored the Southern Ocean. Mortensen (1936) identified
the ophiuroids and echinoids, John (1938) identified the crinoids, Fisher (1940)
identified the asteroids, and Ahearn (1994) identified the holothuroids.
The R/V Vema (1958–1962), from the Lamont Geological Observatory,
explored the Argentine Basin. Bernasconi (1966b) studied both echinoids and
asteroids. The Campagne de la Calypso in 1961–1962, explored 29 stations on the
shelf off Uruguay and northern Argentina (33–39°S) and a few samples from
Nuevo gulf. Most results were published in special issues of the Annales de
l’Institut Oceanographique (Monaco), vols. 44 (1966), 45 (1967), 47 (1969–1970),
49 (1973), and 55 (1979).
The US Antarctic Research Program onboard the USNS Eltanin from 1961 to
1966 studied the Magellan region and Malvinas Islands. Agatep (1967), Pawson
(1977) and Speel and Dearborn (1983) studied the ophiuroids, holothuroids and
crinoids respectively. The FRV Walter Herwig in 1966 and 1971 studied the
Uruguayean and Argentine Break, and the Malvinas Islands. From this expedition
Bernasconi (1973b) studied the ophiuroids, echinoids and asteroids, and Hernán-
dez (1982) studied the holothuroids. For the second campaign (1971) of the Walter
Herwig, only the ophiuroids were studied by Bartsch (1982).
The USNS Hero (1969–1971) from the US Antarctic Research Program explored
the Argentine shelf. Tommasi et al. (1993) studied the ophiuroids and Larrain Prat
(1975) studied the echinoids. There were no reports of other echinoderm classes.
Since 1955, Russian vessels such as RV Ob (1955–1958), RV Akademik
Kurchatov (1971), RV Dmitriy Mendeleev (1989) and RV Akademik Sergei
Vavilov explored the western coast of South America. Abyssal echinoderms
associated with the waters of the Argentine Sea were collected (Gebruk 1988,
1990, 1993; Gebruk and Shirshov 1994; Malyutina 2004).
From 1965 to 1972 a Brazilian vessel Almirante Saldanha explored the
Brazilian, Uruguayean and Argentine waters, from 20°S to 38°S. Tommasi (1965,
1966, 1968, 1969a, b, 1970, 1971, 1974) and Tommasi et al. (1988a, b) described
all of the echinoderms collected.
The Shinkai Maru, from Japan, made twelve expeditions to the Argentine Sea
between 1978 and 1979. Four of these campaigns collected echinoderms from 46
benthic stations. Only the results of holothuroids have been published (Hernández
1982, 1985).
Many Argentinian vessels have explored the Argentine Sea: ARA Rosales,
ARA Azopardo, ARA Patria, ARA San Luis, CAP Undine, ARA Independencia,
ARA Pueyrredón, CAP Maneco, ARA Comodoro Rivadavia, ARA Bahía Blanca,
ARA Capitán Cánepa, ARA Zapiola, Hero, Sta. Maria Magdalena, Lamatra,
Torre Blanca, ARA San Juan, ARA Madryn, B/O Puerto Deseado and multiple
expeditions from the Instituto Nacional de Investigación y Desarrollo Pesquero
(INIDEP) (Bernasconi 1937, 1941b, 1953, 1962b, 1964a, b, c, 1972; Bernasconi
and D’Agostino 1977; Hernández 1981, 1982, 1987).
11.2.2 Diversity
The echinoderm diversity of Argentina (Table 11.1; Appendix) is composed by

134 species, being Asteroidea the most speciose with 58, followed by Ophiuroidea
(32), Holothuroidea (26), Echinoidea (16), and Crinoidea (2). Nevertheless, the
taxonomic status of some groups (i.e. Asteroidea, Ophiuroidea and Holothuroidea)
needs to be reviewed.
Crinoidea
The stalked crinoids, predominantly associated with bathyal zones (Messing 1997;
Améziane and Roux 1997), have not been found in the Argentine Sea. Mortensen
Table 11.1 Numbers of echinoderms from Argentina by taxonomic classification

Orders Families Genera Species
Crinoidea 1 1 2 2
Total 17 42 94 134
(1917, 1920b) described two viviparous comatulids, Phrixometra nutrix and

Isometra vivipara, from the family Antedonidae. These species are the only two
crinoids reported until now from Argentina.
Asteroidea
The systematics and distribution of the class Asteroidea were first studied by
Bernasconi (1934, 1935, 1937, 1941a, 1943, 1961, 1962a, b, 1963, 1964a, b, c,
1965a, 1966a, 1972, 1973a, 1980) and later by Tablado (1982) and Tablado and
Maytia (1988). These authors reported a total of 58 species of sea stars inhabiting
the region. Twenty-eight species are found exclusively along the Magellanic
Province, 13 along the Argentine Province, while the others are found in both
provinces. Six orders are represented: Paxillosida, Notomyotida, Valvatida,
Velatida, Spinulosida and Forcipulatida. The order Valvatida is the most numerous
one, with 19 species. Forcipulatida has 14 species and Paxillosida has 11 species.
In the order Forcipulatida, Hernández and Tablado (1985) identified three species
of the genus Anasterias: A. antarctica, A. minuta and A. pedicellaris, off Ría
Deseado (*48°S). However, according to Clark and Downey (1992) this remains
to be settled. Although they recognized that there are morphologically distinct
forms among the genus Anasterias, the sea stars can belong to different species or
they may comprise a very variable species. In the same order, Allostichaster has
one representative in Argentine waters. According to Clark and Downey (1992)
this species is a synonym of Allostichaster capensis from South African waters,
but this is not certain. The order Velatida is represented by nine species, followed
by Spinulosida and Notomyotida with three and two species, respectively.
Ophiuroidea
Knowledge about diversity of brittle stars from Argentina is based on Bernasconi

and D’Agostino (1971, 1974, 1977). Thirty-two species of brittle stars have been
reported from the region. Only one is an endemic species, Amphilepis sanmatiensis
from the San Matías gulf (Bernasconi and D’Agostino 1975). The order Euryalida
is represented by the family Gorgonocephalidae with two species,
Gorgonocephalus chilensis and Astrotoma agassizzi. The order Ophiurida is well

represented, with eight families and 18 genera. Family Amphiuridae is the most
speciose with 13, of which seven in the genus Amphiura. A complete taxonomic
revision, based upon additional benthic samples, is needed (Brogger et al. 2008).
Echinoidea
Sixteen known species of sea urchins are present (Mortensen 1910, 1936;
Bernasconi 1924, 1925, 1941c, 1947, 1953, 1964c, 1966b, 1970). Two are
members of the Subclass Cidaroidea and 12 belong to the Subclass Euechinoidea.
Spatangoida is well represented, with eight species, followed by Camarodonta (4),
Cidaroida (2), Clypeasteroida (1) and Arbacioida (1). There is a marked pre-
dominance of Magellanic species. Some Magellanic species have a wide distri-
bution (e.g., Pseudechinus magellanicus and Arbacia dusfresnii) reaching low
latitudes (*35°S) but only in deeper waters (Bernasconi 1947, 1957).
Holothuroidea
Twenty-six species of holothurians occur in the Argentine Sea (Martinez et al.

2009). Records of holothurians from South of Argentina are scarce, but they
increase northward in the Magellanic Province (Pawson 1969). There are a great
number of species from cold waters that also occur off Buenos Aires (35°–38°S).
At these latitudes, holothurians are found only in deep waters (Bernasconi 1941b;
Hernández 1981; Martinez 2008). Holothurians from the Argentine Province are
restricted to the North (Deichmann 1947). Overall, cold water species
predominate.
Dendrochirotida is well representated by 18 species, followed by Apodida (5)
and Molpadiida (3). The fauna has the closest similarity with those of Chile
(Pawson 1969) and Antarctica (Deichmann 1947).
The diversity of substrates in ACS provides suitable habitats for several species
of holothurians. Some are infaunal, and others are epifaunal on rocky bottoms,
while others live on Macrocystis sp. holdfasts and fronds or on hard substrata
provided by other invertebrates (Bernasconi 1941b; Deichmann 1947; Hernández
1981; Zaixso and Lizarralde 2000; Martinez 2008).
11.3 Research
11.3.1 Biological Studies
Crinoidea
Crinoids are the least species-rich class of the Argentine echinoderms and there-
fore the least studied among them. Only two species are known from the region.
Andersson (1904) and Mortensen (1917, 1920b) studied the brooding habits of
Isometra vivipara on the Argentine shelf using samples collected by the Sweden
South Polar Expedition. Austin H. Clark (1915 et seq.) covered the Argentine
species in his monograph. Speel and Dearborn (1983) provided important tools for
the identification of crinoids from the Southern Ocean.
Tommasi described crinoids from Argentina and Brazil, collected by the
expeditions of the Brazilian vessel N/Oc. Almirante Saldanha between 26–38°S
(Tommasi 1969b; Tommasi et al. 1988a, 1988b). Only one publication (Bernas-
coni 1934) deals exclusively with Argentine taxa.
Asteroidea
Research has been focused mainly on eight species of sea stars: Allostichaster
capensis, Anasterias minuta, Asterina stellifera, Cosmasterias lurida, Astropecten
brasiliensis, Luidia ludwigi scotti, Ctenodiscus australis and Diplopteraster clarki.
Allostichaster capensis (= A. inaequalis; Fig. 11.3D) is a fissiparous sea star
inhabiting South Atlantic Ocean coasts (Clark and Downey 1992). This sea star
has been extensively studied. Pucheta and Urban (1989) focused on trophic
behavior. The authors carried out laboratory and in situ observations by SCUBA
diving in NG. Allostichaster capensis is a relatively specialized predator, feeding
on the gastropod Tegula patagonica and on the mussels Lithophaga patagonica
and Aulacomya atra atra, selecting this last species over the other prey regardless
of their abundance. Rubilar et al. (2005) found that A. capensis has a very low
percentage of females and displays an annual sexual and asexual reproductive
cycle. Fission is most frequent in summer (November-January) and arm regen-
eration occurs during the rest of the year. In March, gametogenesis begins when
gonads appear in the regenerating arms. In August, sexual maturity is achieved.
Shedding of gametes does not take place until the arms are almost fully regen-
erated and individuals are ready to undergo fission. Spawning occurs during
September and October followed by a long recovery phase from November to
March while arms are in the first stages of regeneration. The pyloric caecum index
displays a clear annual cycle and a reciprocal relationship with gonad index,
demonstrating the storage function of pyloric caeca and suggesting nutrient
transfer. Biochemical analysis revealed that only lipids are translocated from
pyloric caeca to gonads at the beginning of gametogenesis (Rubilar et al. 2008a).
Fig. 11.3 Common species of echinoderms from Argentina. a The sea star Cosmasterias lurida.
b The brooding sea star Anasterias sp. (probably Anasterias minuta). c The brittle star
Ophioplocus januarii. d The sea urchin Arbacia dufresnii and the fissiparous sea star
Allostichaster capensis. All scale bars = 10 mm
The sea star population appears to be maintained primarily by fission, as only two
females of 840 individuals examined were found. Fission in A. capensis lasts
approximately eight hours. At the beginning of the process the body wall begins to
lose its stiffness just before the two halves start to walk away from each other in
opposite directions opening a wound across the disc. The oral tissue splits easily,
in about four hours. The two halves remain joined only by the aboral tissues and
they continue moving apart until the tissue completely breaks. After fission, the
two new individuals are usually immobile while the wound heals. The body wall
contracts to seal the wound. The tips of the new arms can be observed one week
after fission. Individuals had a variable regeneration rate related to body size and
number of regenerating arms. Larger individuals showed higher regeneration rates
than smaller individuals. Accordingly, individuals regenerating four or five arms
had faster regeneration rates than those regenerating three arms (Rubilar et al.
2006).
The effect of fission and regeneration on feeding, energy storage and gonad
production was evaluated during the different stages of regeneration in the
laboratory, and under differing feeding regimens (Villares 2008; Villares et al.
2009). Split individuals showed the lowest feeding rates, lipids, energy in the
pyloric caeca, and gonad production. Regenerating individuals had the highest
feeding rate, lipids were allocated to the pyloric caeca and gametogenesis was
carried out. Individuals with arms in an advance regeneration state showed an
intermediate feeding rate and no allocation of lipids to the pyloric caeca. These
individuals spawned, demonstrating that the gonads were mature. Availability of
food and healthy nutritional state before fission seem to play key roles in survival.
The sea star appears to be extremely well adapted to carry out sexual and asexual
reproduction by increasing feeding rates. Regenerating individuals increase their
feeding rates to store energy for regeneration, gamete production and nutrient
storage. When gonads are mature, individuals have arms in an advanced state of
regeneration, and feeding rates decrease since spawning may inhibit feeding. After
fission, when capacity to attack prey is reduced, the stored energy is used for
survival. Only those individuals that were able to store energy through high
feeding rates can survive. During this period, individuals do not spend energy in
gamete production, but invest all energy resources in regenerating new arms,
which are necessary to increase feeding capacity for the following sexual repro-
duction period.
The great regenerative capacity of this sea star led Rubilar (2009) to study the
cellular mechanism involved in arm regeneration. The sea star regenerates through
a morphallatic mechanism (i.e. cellular migration, dedifferentiation and transdif-
ferentiation), followed by a morphogenetic process producing a miniature arm.
Then new structures arise between the arm tip and the stump. The morphology of
the radial nerve cord and circumoral ring of the sea star was examined by Rubilar
et al. (2008b), using antisera raised against the gastropod neuropeptide Tritonia
Pedal Ganglia (TPep) (Lloyd et al. 1996). This peptide has been found in mo-
torneurons as well as in neurons located in the pedal ganglia of the gastropod
Tritonia diomedea and appears to be involved in motor functions. In the sea star,
TPep-like immunoreactivity was detected in the hyponeural, ectoneural systems
and innervations of the tube feet. In the hyponeural system, the label was observed
in numerous neurons and groups of cell bodies in the radial nerve cord as well as in
the circumoral ring. In the ectoneural systems, the TPep-like immunoreactivity
was observed in isolated neurons as well as in the neuroepithelium.
The oral-brooding sea star Anasterias minuta (Fig. 11.3b) is the most abundant
intertidal sea star in Patagonia (Salvat 1985; Gil and Zaixso 2007). This species is
widely distributed along the Patagonian coast, including the Malvinas Islands, and
occurs from the middle intertidal zone to 80 m depth (Bernasconi 1964c).
Brooding females bend their arms in order to create a chamber to protect the
young. In Ría Deseado (RD; 47°450 S, 65°550 W), brooding occurs during the
coldest months of the year, from March to October (Salvat 1985; Gil and Zaixso
2007). Salvat (1985) suggested a biennial oogenic cycle. Gonads are in a recovery
phase from April to June after eggs are laid. Gametogenesis begins in July and
vitellogenesis continues during the second austral winter; maturity is reached in
January. Female gonads might take around 20 months to mature. On the contrary,
males display an annual cycle. Spawning occurs once a year during March and
April, following a short recovery period in May. Gametogenesis begins again in
June, and maturity is reached in February (Salvat 1985). The pyloric caecum index
displays a clear annual cycle, with maximum index values occurring before the
maximum gonad weight, suggesting nutrient translocation. Except in males, in
which a one month lag was observed, no inverse relationship was found between
pyloric caeca and gonads (Gil and Zaixso 2007). In Punta Maqueda (46°010 S,
67°010 W), the brooding season of A. minuta shows the same pattern (Escudero
1999). The number of juveniles follows a linear regression with female size
(Escudero and Zaixso 2003). This sea star is a common predator and prominent
member of intertidal and shallow sublittoral communities (Gil 2002; Zaixso 2004;
Bertness et al. 2006; Gil and Zaixso 2007). The relationship between predatory and
brooding activities in RD was studied by Gil (2002) and by Gil and Zaixso (2007).
They found a seasonal feeding cycle with the maximum feeding frequency during
the austral summer that was related to the female brooding habits. During brooding
season, females with juveniles did not feed. The feeding frequency was also
positively correlated with seawater temperature and sea star size, but no differ-
ences were found along a shore-level gradient. Anasterias minuta consumes a wide
range of prey, including mollusks and crustaceans. The diet composition differed
among the study sites (GSJO and RD), and it varied with depth and body size. The
diversity of prey consumed by this sea star was highest on a semi-exposed coast
during the warmest season on the infralittoral fringe. Abundance of prey beneath
rocks within tide pools at the infralittoral fringe does not seem to determine diet
composition. Generally, the sea star consumes the purple mussel Perumytilus
purpuratus and the gastropod Pareuthria plumbea more than other prey, even
when they are not the most abundant prey present. However, at the midlittoral
zone of semi-exposed coast, it feeds on prey species according to their abundance.
A significant positive correlation was found between the size of the sea star and the
size of the most frequent prey. The authors suggested that A. minuta could have a
variable impact on the community structure since there appears to be an oppor-
tunistic feeding strategy, taking unattached, wave-washed mussels when available
rather that attached mussels (Gil and Zaixso 2008). An ascothoracid parasite
(Dendrogaster argentinensis) of A. minuta was found at RD (Grygier and Salvat
1984). It is the first ascothoracid described from the South American coast.
The sea star Asterina stellifera (= Patiria stellifer) is distributed from Cabo Frio
(Brazil) to Mar del Plata (MDP) (Argentina; 38°100 S, 57°100 W) in the West
Atlantic Ocean (Clark and Downey 1992) and is a very common sea star on MDP
shores (Salvat 1975a). The embryonic and larval development was studied by
Salvat (1975a). The dipleurula larvae appear 32 h after fertilization and the larva is
completely formed after one week. The size of the sea star varied with depth; the
largest individuals were found in deeper waters (Salvat 1975b). This sea star has
been defined as an omnivore and scavenging feeder (Meretta et al. 2009a). It feeds
on macroalgae, bryozoans, tunicates, polyps and biofilm. Between 2006 and 2009,
the size structure distribution changed from bimodal to unimodal, thus suggesting
that the younger cohort had grown to reach the maximum size and that there was
no recruitment during this period. The high density (35 ind m-2) and the trophic
range suggest a potential role in the hard bottom community (Meretta et al.
2009b).
Stomach contents of Astropecten brasiliensis and Luidia ludwigi scotti have
been analyzed by Brogger and Penchaszadeh (2008) at Puerto Quequén coast
(38°370 S, 58°500 W) between 40 and 60 m depth. Astropecten brasiliensis was the
most abundant sea star sampled and showed the most diversified diet, with a total
of *28 different items. Bivalves and gastropods were dominant. In contrast,
L. ludwigi scotti was found in lower numbers and preyed upon eight species.
Bivalves, foraminiferans and ophiuroids were the most common prey. There were
differences not only in items eaten by the two species, but also in the size of the
common prey. This suggests that differences in diet may allow the coexistence of
the two predators. Penchaszadeh (1973) also observed mollusks as main prey items
for A. brasiliensis off Mar del Plata.
The sea star Cosmasterias lurida (Fig. 11.3a) shows a continuous distribution
from the Argentine shelf (38°S, 57°W) to the Chilean coastline (29°S, 78°W)
(Bremec et al. 2000). It is usually found on subtidal coarse bottoms from 0 to
650 m depth (Clark and Downey 1992) in very low densities (Bremec et al. 2000).
Reproduction has been studied on pier piles of Almirante Storni in NG (Pastor de
Ward et al. 2006). This highly modified habitat had an unusually supply of mussels
and large aggregations of sea stars. Gametogenesis takes place from June to
January, sexual maturity occurs during the summer, and spawning in April. The
gonadal index and pyloric caecum index tended to have an inverse relationship,
but this was statistically significant only in males. However, biochemical analyses
indicated no transfer of nutrients between pyloric caeca and gonads in either sex.
Pastor de Ward et al. (2006) suggested that the high availability of trophic
resources in the study area could be the clue to the high allocation of nutrients, the
absence of nutrient transfer from the pyloric caeca to the gonads during prolif-
eration and the continuous proliferation of oocytes in C. lurida at the Almirante
Storni pier.
The brooding sea star Ctenodiscus australis is distributed along the east coast of
South America, from southern Brazil (23°000 S, 41°400 W) to TF and the Malvinas
Islands, between 70 and 700 m depth. It is usually found on sand bottoms,
occasionally on coarse sand or sand with stones (Clark and Downey 1992). This is
a conspicuous species in the community associated with the shelf-break front
(Bremec and Lasta 2002). While brooding, the young (at different stages of
development) are hidden under the paxillae, mainly at the base of the arms near the
interradius. The youngest stages were entirely hidden by the paxillae. In advanced
stages, parts of their bodies protruded between the spines and are easily seen. The
youngest stages show a larva typical of an abbreviated development (e.g. a yolky,
nonfeeding larva) (Lieberkind 1926). The larvae lack arms and ciliated bands.
Nothing is known about internal morphology of the larvae, but they are probably
yolky, or has a direct-development mesogen (McEdward and Miner 2001).
The pterasterid Diplopteraster clarki has a continuous distribution along the
coast of Argentina, from RP to waters off SJOG, between 82 and 600 m depth
(Bernasconi 1973a). This is a brooding sea star with an aboral nidamental

chamber, where fifty embryos can be brooded. The size of embryos ranges
between 1.5 and 7 mm. Inside the nidamental chamber, embryos are located in
specific areas: eggs and smallest embryos are among the paxillae, close to the
osculum; the intermediate embryos are, following the edge of the arms, between
paxillae and resting on the actinolateral membrane; and the largest embryos
(radius = 4–6 mm) are located between paxillae in the interradii. When embry-
onic development is completed, the juveniles leave the aboral chamber by
breaking the suture between the supradorsal and actinolateral membranes. The
estimated brooding season is between September and May, from spring until
autumn (Bernasconi 1937).
Ophiuroidea
Few publications focus exclusively on this group in Argentina. Bernasconi (1965b,

c), Tommasi (1968), and Bernasconi and D’Agostino (1971, 1974, 1975, 1977)
published a taxonomic revision for the Argentine region. Bartsch (1982) analyzed
benthic samples from the Patagonian shelf and described external and internal
characters of 11 species of brittle stars. The first reports of Argentinian brittle stars
were provided by Bernasconi (1926, 1934). She documented the presence of
Gorgonocephalus chilensis and Amphipholis squamata in the region.
Zaixso and Lizarralde (2000) examined the distribution of the most frequent
echinoderm species of the SJG and southern SMG, referring to four brittle star
species. They analyzed species densities by using a point kriging method, and
related them to environmental parameters (depth, substrata granulometry, currents
and bottom temperature). Amphiura crassipes (1–25 ind m-2) was found close to
80 m depth on substrates with high percentages of silt and clay. The increase of
abundance was associated with an increase of fine sand substrates and no corre-
lation was found with temperature and currents. Ophiactis asperula was the most
abundant ophiuroid (25–3,125 ind m-2) in the study area. The density of this
species was not related to any particular depth, but the density decreased with
increasing summer temperatures and the increase of silt–clay and fine sand sub-
strates. Higher densities were correlated with faster surface currents. Amphiura
magellanica (5–125 ind m-2) was associated with coarse sand substrates. The
abundance was not related to any particular depth or temperature. However,
abundance was positively related to faster surface currents. The abundance of
Ophioplocus januarii (5–125 ind.m-2) was highest at shallow depths and on
coarse sand substrates, and increased with summer temperature.
Several aspects of the biology of the brittle star Ophioplocus januarii
(Fig. 11.3c) in a shallow water population from SJG were studied. Arribas et al.
(2008) analyzed presence and abundance of the fauna associated with this ophi-
uroid species throughout two years. The authors observed that the fauna associ-
ated, mainly mollusks, arthropods and annelids, remained stable over the study
period (2005–2007). The number of species did not show annual variation. This is
consistent with a biocenosis previously described by Zaixso et al. (1998). Growth

studies were conducted by Rodríguez et al. (2008, 2009) using calcein as a
chemical marker. Growth was observed through a distinct green fluorescent label
on the jaws. Coulin et al. (2009) described jaw growth and related it to individual
body growth. The authors proposed a power function relationship between the
growth of the teeth and dental plate and the body size.
Echinoidea
Sixteen species of sea urchins have been reported from the region. However, there
is information available on the general biology and ecology of only six species:
Arbacia dufresnii, Pseudechinus magellanicus, Loxechinus albus, Tripylaster
philippii, Abatus cavernosus and Encope emarginata.
The sea urchin Arbacia dufresnii (= A. dufresnei; Fig. 11.3d) is widely dis-
tributed around the southern tip of South America, from RP on the Atlantic coast
to the north of Puerto Montt (Chile; 41°300 S, 72°560 W) on the Pacific coast, at a
depth range of 0–315 m (Bernasconi 1953). This species was associated with
shallow waters and areas with coarse sediments (sand) and hard substrates in SJG
and NG (Carriquiriborde et al. 1983; Zaixso and Lizarralde 2000; Brogger 2005;
Epherra 2009). A new turbellarian species (Syndesmis patagonica) was found and
described in the gut of A. dufresnii from NG. This is the first record of a sea urchin
of the genus Arbacia as host of an umagillid (Brogger and Ivanov 2010). The diet
of A. dufresnii was studied by using a qualitative analysis of prey items in gut
contents at different sites and across different depths from Buenos Aires to GSJ.
Arbacia dufresnii is primarily carnivorous (Vásquez et al. 1984), and its diet is
determined by the local composition of potential prey (Penchaszadeh and Law-
rence 1999). Some aspects of the embryonic development were studied by Ber-
nasconi (1942) and Brogger et al. (2004). Reproduction of A. dufresnii was studied
at NG (Brogger 2005; Brogger et al. 2010). There are two spawning periods, a first
spawning event occurs during spring and a second in summer. Gametogenesis is
synchronous between males and females and spawning is synchronized. Epherra
et al. (2009a) and Epherra (2009) found that A. dufresnii has a biochemical cycle
directly related to the reproductive cycle at NG and shows differences between
sexes. A similar prolonged spawning period during spring and summer was also
observed. The overall biochemical composition, in different organs, was similar to
that of other sea urchin species, with a high proportion of protein, followed by
lipids and carbohydrates. The population of A. dufresnii at NG displays a good
nutritional state, as both the gut and the gonads act as storage organs. The
reproduction of this species in southern populations (SJOG) has some inter-
population differences in the duration of spawning events and gonadal indices
(Epherra et al. 2009b).
Pseudechinus magellanicus (Fig. 11.4b) is the most common sea urchin in
southern South America. This species is distributed from 35°S on the Atlantic
Ocean coast through the Magellan Strait and throughout TF, to Puerto Montt,
Fig. 11.4 Common species of echinoderms from Argentina. a The brooding heart urchin Abatus
cavernosus. b The sea urchin Pseudechinus magellanicus. c The sea cucumber Psolidium
dorsipes. d The sea cucumber Pseudocnus dubiosus leoninus. All scale bars = 10 mm
Chile, on the Pacific Ocean coast. The sea urchin occurs from intertidal to 340 m
depth (Bernasconi 1953) and lives in a wide variety of habitats, including tidepools
and rocky and mixed gravel-sand bottoms. It can also be found as associated biota
on Macrocystis kelp and in mussel beds (Carriquiriborde et al. 1983; Bremec and
Roux 1997; Adami and Gordillo 1999; Penchaszadeh et al. 2004; Marzinelli et al.
2006). At SJG P. magellanicus has a wide distribution, and its abundance appears
to be related primarily to sites with strong surface currents, dominated by coarse
substrates at intermediate depths (Zaixso and Lizarralde 2000). This distribution
seems different from that of A. dufresnii and may be related to the consumption of
both live food (algae and animals) and drift-material produced by currents (Zaixso
and Lizarralde 2000). Pseudechinus magellanicus has been noted as a member of
various major macrozoobenthos associations in SJG (Carriquiriborde et al. 1983;
Zaixso et al. 1998) and SJOG (Roux et al. 1995). A parasitic gastropod (Pelse-
neeria sudamericana) is attached to the test in Isla de los Estados (54°450 S,
63°520 W) (Pastorino and Zelaya 2001). In the Beagle Channel (54°500 S) this
species was sexual mature at a diameter of 12 mm. The relative fecundity
fluctuated between 70,000 and 100,000 eggs/gonad. The reproductive cycle pro-
duced a fully synchronous spawning event between late winter and early spring.
A new gametic proliferation may occur during the spring with subsequent partial
spawning in the summer (Orler 1992). At NG a main spawning event occurrs in
winter and a second minor event during the summer (Marzinelli et al. 2006). The
gonad index at NG was negatively correlated with photoperiod and water tem-
perature with a lag period of 3 mo. Egg strain values are high in eggs spawned
by small individuals. According to Marzinelli et al. (2006) the jelly coat of eggs of
P. magellanicus may protect them from the forces experienced during spawning,
thus allowing reproductive success.
Diet was studied by Penchaszadeh et al. (2004). The authors suggested an
omnivorous behaviour related to different ecological conditions. The diet spectrum
of P. magellanicus depends mostly on the habitat and biotic community in which
they occur. The species has been identified as a grazer in the benthos, associated
with Macrocystis pyrifera forests in Beagle Channel (Ojeda and Santelices 1984;
Vásquez et al. 1984; Adami and Gordillo 1999), and as a secondary consumer in
circalittoral mussel banks, feeding on barnacles, polychaetes, bivalve recruits,
small gastropods and crustaceans (Penchaszadeh 1979). It can also feed on drifting
or detached algae (Castilla and Moreno 1982; Penchaszadeh et al. 2004).
According to Penchaszadeh et al. (2004) this species can play an important role in
the community structure of the circalittoral beds of Mytilus edulis platensis,
mainly through intense predation on bivalve spat.
Due to the invasion of the kelp Undaria pinnatifida in Argentine Patagonia, the
potential role of invertebrate grazers, such as the sea urchins P. magellanicus and
A. dufresnii were preliminarily analyzed by Teso et al. (2009). They found that
both species of sea urchins fed on the alga at a low rate. In this context, they are
unlikely to play a role in the control of the seaweed because of the high rates of
recruitment and growth of U. pinnatifida.
The edible sea urchin Loxechinus albus has a wide geographic distribution
along the Pacific coast, from Ecuador (6°S) to south of TF including the Magellan
Strait and Beagle Channel (BC) (Bernasconi 1947, 1953) and Malvinas Islands
(Schuhbauer et al. 2010). Growth was analysed by Schuhbauer et al. (2010) in the
Malvinas Islands and growth curves were found to be asymptotic with the von
Bertalanffy growth model as the best predictor. The authors argue that the rela-
tively short period of optimal water temperatures limited this species’ final size
compared to populations in regions with higher seasonal temperatures (e.g. central
Chilean coasts). Reproductive aspects of L. albus populations have been studied
along the Chilean Pacific coast (Gutiérrez and Otsú 1975; Zamora and Stotz 1992),
in the Magellanic region (Bay-Schmith et al. 1981; Oyarzún et al. 1999), in the
Beagle Channel (Orler 1992; Pérez et al. 2008, 2010, 2009) and at the Malvinas
Islands (Schuhbauer et al. 2010). The BC population of L. albus represents the
southernmost limit of distribution. There it is exposed to strong variations in
photoperiod, temperature, and marked seasonal fluctuations in primary produc-
tivity (Hernando 2006). In BC, a major peak of gonadal index occurred in winter,
followed by spawning in spring. A minor gonadal index peak occurred in autumn,
explained by an increase in the nutritive phagocytes, a non-gametic component of

gonads (Pérez et al. 2010). A similar spawning period was reported for L. albus
populations of the Magellanic region (Bay-Schmith et al. 1981; Arana et al. 1996;
Oyarzún et al. 1999). However, populations of L. albus from other sites do not
seem to follow this pattern. In the Malvinas Islands, L. albus spawns during the
warmest time of the year (summer). This allows for greater growth of their pelagic
larvae and newly settled juveniles (Schuhbauer et al. 2010). Spawning is delayed
along the Chilean coast as latitude increases (Zamora and Stotz 1992; Vásquez
2007).
In the BC population of Loxechinus albus, the reproductive cycle may be
influenced by photoperiod fluctuation and primary production. In contrast, water
temperature was not correlated with gonadal maturation (Pérez 2009; Pérez et al.
2010). Pérez et al. (2008, 2010) made the first attempt to analyze the energetic
content of different organs of L. albus using calorimetric determinations. High
values of gonad energy content suggest the presence of lipids in the gonads. Both
gonad energy content and total gonad energy content were higher in females than
in males. The reproductive output of L. albus was higher in females than males,
thus indicating that spawning is energetically more expensive in females (Pérez
et al. 2010). The values of gonad energy content and total gonad energy content
showed minimum values in winter (ripe stage) and maximum values in spring
(spawned stage) (Pérez et al. 2008). Total gonad energy content reached higher
monthly average values (50–200 kJ) than total gut energetic content (20–40 kJ).
These authors suggested this difference indicates that the gonads constitute the
most important storage organs in L. albus. Organic stores are built up in the gonads
after spawning and then used in gamete production.
The gonads of this sea urchin are a highly valued seafood product and are
considered as delicacies in many parts of the world. Pérez et al. (2010) suggested
that autumn is the best period to harvest this species since the gonads are in early
stages, having the quality (color, texture, and flavor) desired by the market.
Malanga et al. (2007) studied the addition of carotenoid in diet in order to improve
the organoleptic properties and protection against oxidative damage of gonads.
They fed L. albus with carotenoid during the pre-reproductive period and during
spawning. b-carotenoids, not only provided a desirable color to the product, but
also protect their lipid constituents against oxidation. The influence of seasonal
factors on oxidative metabolism in the L. albus gonads was tested by Malanga
et al. (2009). They found that lipid radical content showed non-significant changes
with seasons while the content of ascorbyl radicals and -tocopherol were lower in
spring and summer as compared to winter. The authors suggested the existence of
different profiles of response against oxidative stress of the lipophilic and hydro-
philic milieus in L. albus gonads.
The sand dollar Encope emarginata inhabits the Atlantic Ocean from Colombia
to the northern coasts of Argentina. Brogger and Penchaszadeh (2003) described
some aspects of population ecology at MDP at 15–25 m depth. Length frequency
analysis indicated that E. emarginata is characterized by slow growth rates,
reaching a smaller size than populations from Brazil. They reported a very low
frequency of occurrence of the commensal crab Dissodactylus crinitichelis
between oral surface spines.
Tripylaster philippii is a spatangoid echinoid known from the region. This
species is found in greater depths with a maximum at 135 m. It is associated with
substrates having high percentages of silt–clay fraction and very fine sands with a
clear predominance of the mud fraction. Despite the existence of suitable sub-
strates in large areas at SJG, this species is absent from those areas, probably
because the fine sediments of SJG are not stable enough to allow its establishment,
or perhaps summer temperatures are too high (Zaixso and Lizarralde 2000).
Tripylaster philippii is also found at great depths of NG (Carriquiriborde et al.
1983). At SJOG it is part of an important benthic association with the colonial
coelenterate Renilla sp. (Roux 2000).
The heart urchin Abatus cavernosus (Fig. 11.4a) is another spatangoid that
occurs off RP to TF. This species also occurs in the Subantarctic and Antarctic
Islands (Bernasconi 1953, 1966b; Schatt and Féral 1991; López Gappa and Sueiro
2007). It inhabits unconsolidated, muddy and sandy sediments and has a wide
bathymetric range, from the intertidal to 700 m (Bernasconi 1966b). In RP, this
species has a mean density of 63.9 ind m-2 and can be found buried in sand,
muddy sand, and pebbly sand substrata, from midlittoral to the shallow subtidal,
and within soft-bottom tidepools (Dolcemascolo and Zaixso 2000; Gil et al. 2009).
According to Gil et al. (2009), A. cavernosus is a deposit-feeder that broods its
young inside four dorsal ambulacral pouches. Sex ratio was not homogeneous
across size classes and females were found to be larger than males. The temporal
pattern in the size-adjusted gonad dry weight demonstrated that reproduction in the
littoral population of A. cavernosus is seasonal and synchronous. Spawning occurs
from May to July, and females breed within a 9-month period from May to
February, followed by a 1–2 month lag between gonad development and
decreasing seawater temperature. Spawning occurred when the seawater temper-
ature decreased and may be influenced by spring and summer periods of sediment
enrichment. Juveniles develop directly without formation of larval structures, and
are released during the austral summer. The heart urchin exhibited some repro-
ductive characteristics that differ from those of other Subantarctic and Antarctic
spatangoids, including relatively large egg size and low fecundity. Tolosano et al.
(2006) found that gonopore diameter may be used to sex immature heart urchins,
using a breakpoint regression model. The epibiont bivalve Waldo parasiticus is
usually found among the spines of A. cavernosus (Zelaya and Ituarte 2002).
According to Zaixso et al. (2003) the number of bivalves per host is not influenced
by sea urchin sex, but it is positively related to size. Epibionts were mainly found
on the ventral ambulacral areas but can also occupy other areas on the ventral and
dorsal side. Maximum abundance of epibionts per host was found during summer
and lowest values in winter. The relation between these two species should be
tentatively classified as inquiline or commensal.
Holothuroidea
Some holothurian specialists that worked with species from Argentina were Risso
(1826), Lesson (1830), Müller (1850), Phillipi (1857), Lampert (1889), Perrier
(1904, 1905) and Cherbonnier (1941). More recently Pawson (1964, 1969, 1970)
studied some species distributed from Chile to Argentina. Massin (1994),
O’Loughlin (2002, 2009), Bohn (2007), O’Loughlin and Ahearn (2008) and
O’Loughlin (2009) studied the distribution and taxonomy of many species from
Antarctica and near waters with distribution in Argentina. Important advances in
taxonomy of holothurians from Argentine waters have been made by Tommasi,
through multiple expeditions made with the Brazilian vessel N/Oc. Almirante
Saldanha (Tommasi 1969a, 1971, 1974; Tommasi et al. 1988a, b).
Bernasconi described some species from the collection of the MACN (1934,
1941b). Hernández (1981) described the holothurians of Puerto Deseado and three
members of the genus Trachythyone from the southwestern Atlantic, Trachythyone
peruana, T. parva and T. lechleri (Hernández 1982); he also described Trachy-
thyone baja (Hernández 1987). According to O0 Loughlin (2009) this species is a
synonym of T. bouvetensis. Hernández (1985) studied several aspects of different
holothurian species, including a taxonomic description of all species inhabiting the
Argentine Sea. He also analyzed the phenetic relationship among the holothurian
species found, and the phylogenetic relationships of five genera of the family
Cucumariidae, studied the biology and ecology of some intertidal species, pub-
lishing data on their distribution. Besides this, he studied the sediment feeding
behavior and evisceration as a defense mechanism of Chiridota pisanii (Hernández
1985). Martinez et al. (2009) reported the total number of holothurians from
Argentina, and developed tools for species recognition in the families Cucum-
ariidae (Fig. 11.4d) and Psolidae (Fig. 11.4c).
Different authors have studied multiple aspects of the reproduction of the
brooding dendrochirotid Psolus patagonicus (Martinez 2008; Giménez and
Penchaszadeh 2010; Martinez et al. 2011). Martinez (2008) studied the ecology
and reproduction of a P. patagonicus population that inhabits the shelf break at
38°S (*100 m depth). This species exhibited an annual reproductive cycle with
one reproductive event in late summer/autumn (February–March). The maximum
oocyte diameter registered in P. patagonicus was 900 lm with more or less
continuous development and growth of oocytes all through the year. Giménez and
Penchaszadeh (2010) estimated the brooding season from February to October and
the beginning of the brooding period is consistent with a decrease in the gonadal
index of females, larger oocytes and mature spermatozoa in males (Martinez 2008;
Martinez et al. 2011). Schejter et al. (2002) reported a seasonal food input in the
associated Patagonian scallop community, which is correlated with the repro-
ductive timing of P. patagonicus, thus reinforcing the contention that this factor
influences the timing of reproduction (Martinez 2008; Martinez et al. 2011).
11.3.2 Argentine Continental Shelf Waters
The study of benthic communities in ACS started four decades ago (Bastida et al.
2007). In some of these studies, echinoderm species showed high abundance and
diversity and were used to define different ecological areas along the ACS (Roux
et al. 1988, 1993; Bastida et al. 1992).
The ecological information on echinoderms from shelf waters comes from
monitoring studies aimed at determining the invertebrate by-catch associated with
commercial fisheries. The first faunistic inventory of the Patagonian shrimp
(Pleoticus muelleri) fishing grounds of SJOG was developed by Roux et al. (1995).
The benthic community between 27 and 98 m showed a total of 22 species of
echinoderms with a higher abundance in soft than in hard bottoms. Pseudechinus
magellanicus, together with four species of mollusks and one crustacean were
dominant in the benthic assemblage. Other species commonly collected in the gulf
were Henricia obesa, Cosmasterias lurida, Ophiomyxa vivipara and Ophiactis
asperula.
In external shelf areas the assemblage of macroinvertebrates dominated by the
Patagonian scallop Zygochlamys patagonica has been broadly studied. This
community is located along the shelf-break front, following the 100 m isobath,
between 37° and 44°S. Echinoderms comprise one of the most important taxa, in
terms of biomass and numbers. The northern sector (37–39°S) is characterized by
high density of Ctenodiscus australis, Diplasterias brandti, Labidiaster radiosus
(Asteroidea), Gorgonocephalus chilensis, O. asperula (Ophiuroidea) and the
echinoid Austrocidaris canaliculata. Sterechinus agassizii (Echinoidea), Ophia-
cantha vivipara and O. asperula (Ophiuroidea) and the sea star D. brandti stand
out in the southern sector (Bremec and Lasta 2002; Bremec et al. 2006; Schejter
et al. 2008).
Echinoderms, mainly sea stars, have a potentially important ecological role in
this community. The sea star Calyptraster sp. (possibly Diplopteraster clarki,
according to Escolar and Bremec 2009) appears as the top predator, preying on the
sea star C. australis, and the gastropods Odontocymbiola magellanica, Adelomelon
ancilla and Fusitriton magellanicus. These gastropod species are the main pre-
dators of the Patagonian scallop. Other echinoderm species such as L. radiosus,
C. australis and S. agassizii are intermediate predators, preying on scallops and
brittle stars, scallops being the principal item in the diet of L. radiosus (Botto et al.
2006).
Another conspicuous sea star in ACS is Astropecten brasiliensis. This is an
abundant species in the Buenos Aires Province coastal and shelf areas (Roux
2004). It inhabits soft bottoms (sand-mud) between 7 and 60 m. This species feeds
on banks of the mussel M. edulis platensis, commercially exploited until 1988
(Penchaszadeh 1980; Ciocco et al. 1998). This sea star is one of the most important
predators of mollusks, mostly young gastropods and bivalves (Penchaszadeh
1973). Astropecten brasiliensis shows a selective diet that varies with depth and
bottom type, limited only by the size of prey. This sea star ingests prey whole; the
maximum mussel size recorded in a stomach was 24 mm long. The diet compo-
sition of this sea star at M. edulis platensis banks varied according to the seasonal
recruitment and growth of mussels. When young mussels were absent (late fall and
winter), the contribution of cumaceans, other pelecypods and gastropods increased
in the gut contents of A. brasiliensis.
With respect to echinoderms from deeper areas of the ACS, there are only the
faunal inventories of the H.M.S Challenger, Vema and Walther Herwig expedi-
tions (Sladen 1889; Bernasconi 1966b, 1972, 1973b; Bartsch 1982). Recently,
samples taken from a submarine canyon at 365 m depth, around 43°350 S and
59°330 W, show that echinoderms are diverse and constitute an important part of
the benthic community (18 species, 14 % wet weight). The same species are found
in scallop fishing grounds, with the exception of the brittle star Astrotoma agassizii
(Bremec and Schejter 2010).
11.3.3 Chemical Studies
Secondary metabolites from Echinodermata have been thoroughly investigated for

the past four decades. Members of this phylum contain steroidal and triterper-
noidal oligoglycosides and saponins, which are widely diffused in terrestrial plants
but apparently not so common in marine organisms (Burnell and Apsimon 1983;
Minale et al. 1986, 1993, 1995; Stonik and Elyakov 1988; Kalinin et al. 1992,
1996, 2005; D’Auria et al. 1993; Verbist 1993; Stonik et al. 1999; Iorizzi and
Riccio 1999; Kalinin 2000; Iorizzi et al. 2001; Chludil et al. 2002b, 2003a, b;
Maier and Murray 2006; Maier 2008).
Asteroidea
Asterosaponins are steroidal glycosides which share a 3b-O-sulfated,6a-O-gly-

cosylated D9(11) steroidal skeleton- only one reported structure shows a 12a-
hydroxy functionality, with different side chains (Yasumoto and Hashimoto 1965).
Twelve asterosaponins have been isolated from four species of sea stars from the
Patagonian coast of Argentina: Luidia ludwigi scotti from SMG, Cosmasterias
lurida and Allostichaster capensis from NG, and Anasterias minuta from SJOG.
Analysis of their chemical structures showed six novel asterosaponins: Cosmas-
terosides A, B, C and D (from C. lurida) (Roccatagliata et al. 1994) and Anas-
terosides A (from A. minuta and A. capensis) and B (from A. minuta) (Díaz de
Vivar 2002; Maier et al. 2002) (Fig. 11.5a). The hexaglycoside Anasteroside
A showed antifungal activity against the plant pathogenic fungus Cladosporium
cucumericum. Two additional known saponins were isolated from C. lurida:
Ophidianoside F, previously isolated from Ophidiaster ophidianus (Riccio et al.
1985), as the main component and Forbeside H, previously isolated from Asterias
forbesi (Findlay et al. 1990). Acanthaglycosides B and C, previously isolated from
b Fig. 11.5 a Cosmasterosides a, b, c and d, and Anasterosides a and b. b Sulfated glycosides

from Cosmasterias lurida and Anasterias minuta. c New polyhydroxysteroid from Luidia ludwigi
scotti. d Patagonicoside A and its desulfated analogue; e Novel sulfated polyhydroxysteroid from
Ophioplocus januarii
the sea star Acanthaster planci (Itakura and Komori 1986), were purified from the
sea star L. ludwigi scotti (Roccatagliata 1996). Versicoside A, an asterosaponin
from Asterias amurensis versicolor (Itakura et al. 1983), was also purified with
Anasterosides A and B from A. minuta (Maier et al. 2002).
Polyhydroxysteroids are also extracted from echinoderms. These compounds
show a polyhydroxylated steroidal aglycone and one or two monosaccharide
moieties with variability in the hydroxylation pattern, the side chain functionali-
zation, the localization of the glycosidic link and the presence of the sulfate
moiety. Two novel steroidal xyloside sulfates (Luridosides A and B) and the
known Pycnopodioside C, previously isolated from Pycnopodia helianthoides
(Bruno et al. 1989), were isolated from the sea star C. lurida (Maier et al. 1993).
Anasterias minuta provided two new sulfated polyhydroxylated steroidal xylo-
sides: Minutosides A and B, together with the known Pycnopodioside B, previ-
ously isolated from P. helianthoides (Bruno et al. 1989). Minutoside B is the first
example of a polyhydroxylated steroidal xyloside containing an amide function in
the aglycone side chain (Fig. 11.5b). The three xylosides from A. minuta exhibited
antifungal activity against Cladosporium cucumerinum and Aspergillus flavus
(Chludil and Maier 2005). Helianthoside, previously isolated from Heliaster he-
lianthus (Vázquez et al. 1993) was obtained from the ethanolic extract of the body
walls of A. capensis (Díaz de Vivar 2002).
Sulfated polyhydroxysteroids were also extracted from sea stars. Roccatagliata
et al. (1995) reported a novel sulfated polyhydroxysteroid from L. ludwigi scotti
(collected at SMG) together with two other known compounds previously isolated
from Myxoderma platyacanthum (Finamore et al. 1991) and Luidia maculata
(Minale et al. 1984) (Fig. 11.5c).
The first study on sterols from sea stars of the coast of Argentina was by Seldes
and Gros (1985) on C. lurida, and later, a mixture of sterols was obtained from
L. ludwigi scotti by Roccatagliata et al. (1996b). These studies show that
24-methyl-D7-sterols account for *40 % of the sterol mixture in both sea stars,
followed by choles-7-en-3b-ol (more than 20 %).
Glycosphingolipids consist of a sphingoid or a ceramide linked at C-1 by a
b-glycosidic bond to a hydrophilic carbohydrate moiety. Glycosylceramides are
the simplest glycosphingolipids and consist of a mono- or a disaccharide chain
linked to the ceramide (http://www.chem.qmul.ac.uk/iupac/misc/glylp.html).
Glucosylceramides (monoglycosylceramides) have been isolated from C. lurida
(Maier et al. 1998), A. capensis (Díaz de Vivar 2002; Díaz de Vivar et al. 2002)
and A. minuta (Chludil et al. 2003a) (Fig. 11.6b). Two new glucosylceramides
were obtained from C. lurida (compounds 3 and 4), together with a mixture of
related cerebrosides obtained as molecular species and Ophidiacerebrosides C and
D (Maier et al. 1998), and first reported from Ophidiaster ophidianus (Jin et al.
1994). Anasterias minuta produced a new glucosylceramide: Anasterocerebroside
A, together with five other previously reported pure cerebrosides. Isolated as a
pure compound, Glucosylceramide 7, previously obtained as molecular species
from C. lurida and A. capensis, was characterized for the first time (Chludil et al.
2003a). The novel Inaequalocerebroside 1 was obtained from body walls and
gonads of A. capensis. The body walls also contained two other new glucosyl-
ceramides as nearly homogeneous mixtures and five known cerebrosides:
Phalluside 1, Ophidiacerebrosides C and D (Jin et al. 1994) and compounds 3 and
4, firstly isolated from C. lurida. Gonads had two new glucosylceramides (Ina-
equalocerebrosides 2 and 3), containing the new LCB (2S,3R,4E,15Z)-2-amino-
4,15-docosadien-1,3-diol and Phalluside 1 as major components. A comparative
analysis of the composition of the cerebroside mixtures obtained from body walls
and gonads of A. capensis showed differences in the fatty acid and LCB moieties:
the 9-methyl, D4,8,10 triene-LCB and saturated or unsaturated (D10 or D15)-24-C
fatty acids were the most abundant in the glucosylceramides obtained from body
walls, while the octadecatriene and the cis-D15-22-C LCBs and 2-hydroxyhepta-
decanoic acid were predominant moieties in cerebrosides from gonads. All fatty
acids isolated from body walls had normal chains whereas gonads had fatty acids
with iso chains as well (Díaz de Vivar et al. 2002).
Ophiuroidea
Secondary metabolites from three species collected at SMG: Ophioplocus januarii,

Ophiactis asperula and Ophiacantha vivipara were studied. Ophioplocus januarii
afforded four sulfated polyhydroxysteroids, one of them is new (Fig. 11.5d). The
antiviral activity of the four compounds was tested against herpes simplex virus
type 1, Junín virus, respiratory syncytial virus and polio virus. These studies showed
the importance of the unsaturation at C-22 for the inhibitory action towards
respiratory syncytial virus and at C24-28 for replication of Junín virus (Roccata-
gliata et al. 1996a). Ophiactis asperula and O. vivipara did not have sulfated
polyhydroxylated steroids as the major secondary metabolites, but showed
nucleosides and nucleobases in their ethanolic extracts instead. From O. asperula,
inosine (19.0 %), 20 -deoxyuridine (10.6 %), thymidine (14.9 %), and the bases
uracyil (17.4 %), cytosine (15.9 %) and thymine (22.2 %) were obtained. Ophia-
cantha vivipara afforded inosine (7.1 %), 20 -deoxyinosine (13.4 %), uridine
(3.8 %), 20 -deoxyuridine (7.1 %), 20 -deoxycytidine (1.4 %), and thymidine (4.6 %)
with the bases uracyl (8.1 %), cytosine (4.8 %), thymine (31.9 %) and hypoxan-
thine (17.8 %) (Murray et al. 2002). Considering echinoderms, 20 -deoxyribonu-
cleosides had only been previously isolated from the sea stars A. planci and
L. maculata (Komori et al. 1978, 1980). Their importance as systematic markers
was suggested by Dematté et al. (1986), since they had only been isolated from
tunicates and echinoderms. This is currently under discussion as they have also been
purified from two marine sponges (Kondo et al. 1992; Searle and Molinski 1994).
Holothuroidea
Secondary metabolites from holothurians are characterized by the presence of

triterpenoidal saponins (holothurins) and can be considered characteristic of the
class. Their toxicity (acting as membranotropic towards cell membranes) can be
associated with a defensive response of the organism and is the cause of their
biological activity (Maier 2008). They have also been used as chemotaxonomic
markers since some holothurins are specific for different taxa (see Maier and
Murray 2006; Antonov et al. 2008).
Two species of holothurians from the Patagonian coast of Argentina have been
studied: Hemioedema spectabilis (collected at SJOG) and Psolus patagonicus
(collected off Bahía Ensenada; 54°500 S, 68°290 W). Patagonicoside A, the major
component of the ethanolic extract of P. patagonicus was isolated and its structure
was determined by Murray et al. (2001) (Fig. 11.5d). Patagonicoside A has
revealed a broad spectrum of biological activities: the glycoside and its semi-
synthetic desulfated analogue were tested for their antifungal activity against the
phytopathogenic fungi Cladosporium cucumerinum (Murray et al. 2001), C. ful-
vum, Fusarium oxysporum and Monilia sp. (Muniain et al. 2008). The intact
compound is more effective than the desulfated analogue. It showed an antifungal
activity comparable with a commercial fungicide towards Monilia sp. Bioactivity
assays showed that the presence of the sulphate groups in the oligosaccharide
chain plays an important role in the antifungal activity. Patagonicoside A and the
desulfated derivative were also tested for antiproliferative, cytotoxic and haemo-
lytic activities and their effect on NF-jB activation (Careaga et al. 2009). Both
glycosides showed low cytotoxic activity in A549 tumour cells, which the authors
attribute to the uncommon presence of 12a,17a-hydroxyl groups in the aglycone
moiety. Both compounds exert an antiproliferative effect although their mecha-
nisms of action do not involve inhibition of NF-jB.
Chludil et al. (2002b) isolated and described two new sulfated triterpene gly-
cosides, Hemoiedemosides A and B from the Patagonian holothurian H. specta-
bilis (Fig. 11.6a). Hemoiedemoside B is a new example of a small number of
trisulfated triterpene glycosides from holothurians belonging to the family Cu-
cumariidae. Both glycosides exhibit considerable antifungal activity against the
phytopathogenic fungus C. cucumerinum, while the semisynthetic desulfated
derivative of Hemoiedemoside A is less active.
Although fisheries of the sea urchin Loxechinus albus has been extensive in Chile,
in Argentina it has been captured only sporadically by artisanal fisheries in the
Beagle Channel near Ushuaia Bay, especially for the local market, to serve tourists
from the Orient, and occasionally exported to Chile.
Fig. 11.6 a Triterpene glycosides from Hemioedema spectabilis. b Glucosylceramides from sea
stars of the Patagonian coast of Argentina
The trawl fishery is considered one of the major factors disturbing the seabed
(Kaiser et al. 2006). Otter trawls collect and kill huge amounts of non-target
species, physically disturb the seabed, and alter the structure of benthic
communities. There are several bottom trawling fisheries in Argentina. Currently,

three fisheries have by-catch monitoring programs in order to assess the condition
of the disturbed bottoms; these are the red shrimp fishery and the Patagonian and
Tehuelche scallop fisheries.
The red shrimp (Pleoticus muelleri) fishery has operated in SJOG since 1980.
Only 20 % of the species found in the benthic community are retained as incidental
catch during the fishery trawls. Echinoderms, together with crustaceans and mollusks,
are the main groups of by-catch. Diplasterias brandti, Henricia obesa, Pseudechinus
magellanicus, Arbacia dufresnii, Austrocidaris canaliculata, Trypilaster philippi,
Hemioedema spectabilis and Gorgonocephalus chilensis are the echinoderm species
caught by the shrimp trawl (Roux et al. 2007); Diplasterias brandti being the most
frequent. The benthic communities subjected to trawling partially maintain their
structure, species richness and spatial distribution (Roux 2008).
The most productive Patagonian scallop (Zygochlamys patagonica) fishing
grounds in Argentina are associated with the shelf-break frontal system and have
been exploited since 1996. Several studies on incidental catches show that brittle
stars, sea stars and sea urchins are the dominant taxa in most of the scallop beds;
reaching 86 % of the total catch in some of them (Bremec et al. 2006, Escolar et al.
2007; Schejter et al. 2008). The biomass of echinoderms as bycatch increased from
1997 to 2002 in sites with high fishing activity (Escolar et al. 2009). Because many
echinoderms are predators, they could be favored by fishery discards. In addition,
an increase in biomass of Ophiactis asperula (detritivore) was recorded in
intensively trawled sites, while Ophiacantha vivipara and Sterechinus agassizii
decreased (Schejter et al. 2008). Although relative abundances of some taxa varied
over time, no loss of species has been recorded in the Patagonian scallop com-
munity since the start of the fishery (Bremec et al. 2008).
The Tehuelche scallop (Aequipecten tehuelchus) fishery in SMG is the main
coastal bivalve fishery in Argentina. This fishery started in 1969 and was closed for
ten years from 1973 to 1983 (Ciocco et al. 2006). Impact studies on biodiversity
reported five species of echinoderms (Arbacia dufresnii, Pseudechinus magell-
anicus, Tripylaster sp., Ophioplocus januarii and Cosmasterias lurida) as part of
incidental catches. The total density of echinoderms remained stable in sites
without fishing activity and increased in sites that were intensively exploited until
2000; the dominant species were A. dufresnii, P. magellanicus and O. januarii.
In contrast, echinoderm density had been reduced at sites subjected to intensive
and continuous fishing activity. Patches of A. dufresnii and P. magellanicus were
depleted in these fishing grounds (Morsán 2009).
11.6 Concluding Remarks
Until recently studies on echinoderms in Argentina were very limited. During the
last century Bernasconi’s publications were the principal references for this
phylum. In the last ten years there was an increase of studies on the echinoderm
fauna of Argentina. This is reflected in recent Doctoral theses (Díaz de Vivar 2002;
Pérez 2009; Brogger 2010; Escolar 2010; Rubilar 2010, Epherra in prep., Gil in
prep., Martinez in prep., Meretta in prep.) and Licenciatura theses (Gil 2002;
Rubilar 2004; Brogger 2005; Marzinelli 2005; Martinez 2008; Villares 2008;
Epherra 2009; Rodríguez 2009; Di Giorgio 2011; Pérez-Gallo in prep; Reartes in
prep.). These studies are mainly focused on biological and ecological aspects.
However, some have focused on chemical, taxonomical, physiological or ana-
tomical aspects. We expect that these will be the beginning of new echinoderm
researches in Argentina.
Acknowledgments The present chapter was greatly improved with the valuable comments on
the manuscript of five reviewers: J Lawrence, D Pawson, P Lambert, J Pearse, and J Cortés. We
would like to thank them for kindly reviewing the manuscript and for their useful suggestions.
We are very grateful to all Argentinean researchers who contributed to increase the knowledge
about this phylum in the region. A special recognition must be done to the Institutions and the
directors for helping us conduct research on echinoderms. Thank you very much to P Penchas-
zadeh, H Zaixso, C Pastor de Ward, C Bremec, J Calvo, and to the late A Seldes and I
Bernasconi.
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Chapter 12
Echinoderms of the Cuban Archipelago
Rosa del Valle-García, Mercedes Abreu-Pérez, Francisco Alonso

Solís-Marín and Alfredo Laguarda-Figueras
12.1 Introduction
The Cuban Archipelago is found between 198500 –238170 N and 748080 –848570 W. It
is comprised of the island of Cuba (105,007 km2), Isla de Pinos (2,200 km2), and
about 4,195 cays and islets (Núñez-Jiménez 1982). The southern coasts border the
Caribbean Sea and the northern the Atlantic Ocean and the Gulf of Mexico.
The island of Cuba is the largest of the group comprising ‘‘the Greater Antil-
les’’. Its coast line is 3,209 km on the north side and 2,537 km on the south side.
The archipelago’s insular platform is shallow (\30 m in depth) and has a sub-
merged plain relief with an emerged surface that, including the cays and islets
(with exception of the island of Cuba), covers an area of 67,831 km2 (Núñez-
Jiménez 1982). According to Ionin et al. (1977), the insular platform may be
considered a unique area from geological, tectonic and geomorphological points of
view. It has four independent shallow regions surrounded by the islands, cays, little
cays and reefs forming archipelagos (Fig. 12.1).
The insular slope is found nearby the exterior edge of the cays. The reefs begin
to decline between 20 and 50 m depth (usually at depths between 25 and 30 m).
In most of the periphery of the platform it drops nearly vertically down to a depth
between 100 and 150 m. This is followed by a zone of smaller slope (60–808)
R. d. Valle-García (&) M. Abreu-Pérez

Instituto de Oceanología, Ministerio de Ciencias
Tecnología y Medio Ambiente (CITMA), La Habana, Cuba
e-mail: rosa.denisse1@hotmail.cominformatica@giron.sld.cu
F. A. Solís-Marín A. Laguarda-Figueras
Laboratorio de Sistemática y Ecología de Equinodermos,
Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma
de México (UNAM), Apdo. Post. 70-305, 04510, Mexico, D. F, Mexico
e-mail: fasolis@icmyl.unam.mx

404 R. d. Valle-García et al.
Fig. 12.1 Satellite image of the Cuban Archipelago (Jeff Schmaltz, MODIS, Rapid Response
Team, NASA/GSFC), and division of the Cuban marine platform into nine ecoregions (taken
from Areces 2002). 1 Southeastern Ecozone (Costa Sur de Oriente), 2 Southcentral Ecozone
(Jardines de la Reina Archipelago), 3 Southcentral Ecozone (Coast to the South of macizo de
Guamuhaya), 4 Southwestern Ecozone (Archipelago of Los Canarreos), 5 Southwestern Ecozone
(Península de Guanhancabibes), 6 Northwestern Ecozone (Archipelago of Los Colorados), 7
Northern Ecozone (Habana-Matanzas), 8 Northcentral Ecozone (Archipelago Sabana-Cama-
güey), 9 Northeastern Ecozone (Costa Norte de Oriente)
which continues with slight changes far beyond a depth of 600 m. These deep
mark the geographic limits of the Cuban Archipelago (Claro 2007).
The salinity and nutrient values of the Cuban platform water show compara-
tively wide seasonal and spatial variation since there is relative isolation of these
oceanic waters, the mean depth is small, and there is a high level of river dam-
ming. During the rainy season salinity may drop to an average of 32 % in some
regions and under certain conditions even more. On the other hand, in other sites
of limited exchange with the ocean, salinity may reach more than 50 % during the
dry season (Lluis-Riera 1972, 1977, 1981a, b, 1983, 1984; Alcolado et al. 1998;
Claro 2007).
The water circulation over the Cuban platform seems to be determined mainly
by the wind, the tide, and the influence of sea currents of the adjacent open sea
(Emilsson and Tápanes 1971) whereas the currents system in the oceanic water
that surrounds the Cuban Archipelago is extraordinarily complex and variable
(Sukhovey et al. 1980; García-Díaz et al. 1991; Victoria et al. 1997; Gallegos et al.
1998).
The horizontal distribution of the primary production in the platform is affected
by the hydrologic regime which is highly influenced by the oceanic currents and
winds and by the input of inland water. Changes on any of these factors or other
factors linked to them (e. g. local variation on the biogenic elements content), may
produce local conditions of variable duration resulting in different environmental
conditions for phytoplankton (Claro 2007).
12 Echinoderms of the Cuban Archipelago 405
12.1.1 Seagrasses
Seagrasses, known locally as ‘‘seibadales’’, are marine angiosperms on uncon-

solidated sediment bottoms. Six species of angiosperms have been recorded in
Cuba: Halophila decipiens, Halophila engelmanni, Thalassia testudinum, Halod-
ule wrightii, Syringodium filiforme, and Ruppia maritima. The greatest amount of
primary production on the Cuban platform is due to marine phanerogams, espe-
cially T. testudinum which generally constitutes more than 70 % of the macro
phytobenthos wet biomass on the Cuban platform.
Seagrasses are the main energy entrance pathway for biological and fishery
productivity on the Cuban platform. It constitutes a major ecological reserve of
matter and energy in biomass. Part of this is exported to the reefs and the ocean
and increases productivity in these biotopes. They also stabilize the substrate,
preventing the erosion on reefs and nearby beaches. They regulate oxygen con-
centrations on the sea water and strongly affect local biogeochemical processes.
Development of calcareous algae, mainly Halimeda, in the seagrass beds produce
most of the sand of beaches (Alcolado 2007).
12.1.2 Coral Reefs
Cuban coral reefs appear in the shape of ridges (‘‘restingas’’), coral barriers,
promontories (‘‘cabezos’’ or patch reefs), alternating bars with sand canals (bot-
toms of ‘‘camellones’’ (ridges), or ‘‘macizos’’ and canals), and carpet boulders and
rocky terraces, amongst other varied structures (Ionin et al. 1977). More than 98 %
of the Cuban marine platform edge is surrounded by frontal reefs, ridges, or
barriers (Alcolado 2007).
The seagrass-reef complex is the source of approximately 60 % of the com-
mercial catches of fish of high commercial value. Reefs contain a great diversity of
microorganisms, plants and invertebrates that have biologically active substances.
These are used or constitute potential resources as drugs or biochemical and
experimental reagents of interest. Its barriers or ridges are an effective protection
for coastal lines against wave erosion. They also protect coastal towns and
buildings. They are extraordinarily attractive for diving tourism, i.e. the reefs of
Archipiélago de los Jardines de la Reina (Alcolado 2007) (Fig. 12.1). Recently,
Figueredo-Martín et al. (2010) evaluated the environmental quality of coral reefs
at Jardines de la Reina and concluded the area was in optimal conservation
condition.
Isle of Youth, National Park of the National System of Protected Areas of Cuba
(Fig. 12.1), has a marine area of 5.15 km2 with coral reefs of major species and
biotope diversity. Forty two species of algae, three phanerogams, 29 sponges, 21
gorgonians, 35 corals, and 79 fishes have been identified in this area (De la
Guardia et al. 2004). The most representative habitats between the coastline and a
depth of 25 m are sand bottoms with or without seagrasses, reefs in shallow

patches, Acropora ridges, rocky esplanades with isolated corals and gorgonians
ending in an escarpment or contour. Reefs in deep patches and spurs and groves
very close to the wall. In general, the Isle of Youth possesses healthy reefs,
although some indicators, such as the damaging effect of a dense growth of algae
and cyanophytes over sessile benthic organisms and the scarcity of large herbiv-
orous or carnivorous fish indicate the existence of potential disruptions in the reef
(De la Guardia et al. 2004).
12.1.3 Mangrove Swamps
Most of the Cuban Archipelago coasts are lined with mangroves, as well as
swampy areas and thousands of coastal lagoons and estuaries. This biotope in
Cuba is integrated by four arboreal species: mangle rojo (red mangrove) (Rhizo-
phora mangle), mangle prieto (black mangrove) (Avicennia germinans), and
patabán (Laguncularia racemosa), considered true mangroves, and the yana
(Conocarpus erectus), a false or pseudomangrove. The mangrove swamps occupy
approximately 4.8 % of the national territory, 26 % of the forest surface of the
country (Alcolado 2007).
Approximately 30 % of Cuban mangroves are being affected by an increase in
salinity and nutrient depletion, the result of damming, pollution and illegal
deforestation. There has also been an increase in contamination from sugar mills,
mining, oil refineries, paper and alcohol industries and cattle farms. Hydrologic
changes on the coasts and at sea have been produced by building dams and
viaducts over the sea. The effect on coasts that have lost the frontline protection of
red mangrove produces a sudden accumulation of sand (Alcolado 2007).
The earliest researches on echinoderms made in Cuban seas were by the vessels
‘‘Blake’’ (1877–1879), ‘‘Fish Hawk’’ (1880–1881), ‘‘Albatross’’ (1883–1884),
‘‘Vanderbilt’’ (1921–1928), ‘‘Eagle’’ and ‘‘Ara’’ (1921–1930), and the ‘‘Atlantis’’
(1938–1939). Leading specialists from Cuba and abroad participated in these
voyages and made valuable contributions to echinoderm knowledge, especially for
those species inhabiting deep waters off Cuba. Most of the results obtained from
those investigations are in the works of Théel (1886), Agassiz (1869, 1878, 1880,
1888), Verrill (1899), Clark (1917, 1931, 1940, 1947, 1950), Clark (1919, 1933,
1941), Deichmann (1930, 1940, 1954, 1963), and Thomas (1963).
Other foreign specialists have made major contributions to the echinoderm
knowledge in Cuba, including Boone (1933, 1935), Mortensen, with his master-
piece ‘‘Monograph of Echinoidea’’ (1928, 1935, 1940, 1943a, b, 1948a, b, 1950,
1951), Tortonese (1956), Tommasi (1958), Pawson and Fell (1965), Downey
(1973), Meyer et al. (1978), Serafy (1979), Pawson and Caycedo (1980), Miller
and Pawson (1984), Clark and Downey (1992), and Hendler et al. (1995).
The first publication by a Cuban scientist was by Arango (1877) ‘‘Los Radiados
de la Isla de Cuba’’. He gave a list of marine invertebrates including the echi-
noderms, the first record for the phylum in the archipelago. Later, Delgado-Núñez
(1917) did his doctoral thesis in Natural Sciences at Universidad de la Habana,
with a contribution on the study of the nervous system of the sea urchin Centre-
chinus antillarum (= Diadema antillarum Philippi, 1845).
The first identification key for echinoderms of Cuba made by Aguayo (1945) is
still of great practical use for scholars. That same year, Sirven-Villuenda (1945)
did his doctoral thesis on the subject ‘‘Equinoideos litorales de Cuba’’ (‘‘Littoral
Echinoids of Cuba’’), and in 1959, Pérez-Farfante (1959) published his doctoral
thesis on irregular echinoids of Cuba, presenting dichotomous keys and descrip-
tions for 24 species of shallow and deep waters.
Subsequently there were no publications related to echinoderms until 1965, when
the Instituto de Oceanología de la Habana (Oceanology Institute of la Habana) was
created and study of echinoderms was resumed. The mission of this institute was to
establish the scientific basis to guarantee preservation and sustainable use of coastal
and marine resources of Cuban ecosystems and adjacent waters through multidis-
ciplinary research. Since then, major development of marine sciences began in Cuba,
under the direction of Dr. Darío Guitar, an internationally renowned and professor at
Universidad de la Habana. Consequently, new talents have emerged who acquired
different specialties and research cruises have been undertaken especially from the
former Soviet Union, Bulgaria and Romania.
During the second half of the twentieth century, joint expeditions were made by
Cuban and European institutions and specialists, like the ‘‘I y II Expediciones
Conjuntas cubano-soviéticas’’ (‘‘1st and 2nd Cuban-Soviet Joint Expeditions’’).
Both of these expeditions studied different aspects of echinoderms, including the
extraction of bioactive substances to determine their potential use in medicine,
especially as anticarcinogens (Archivo Científico, Instituto de Oceanología). To
accomplish this, work was done in close relationship with specialists from the
Oncology and Radiology Institute at la Habana, Cuba.
As a result of the collections made by these cruises and their identification of
the specimens deposited in the collection at Instituto de Oceanología, Levin and
Gómez (1975) published a work on the holothurians of shallow waters of Cuba.
They described the external morphology and spicules of 17 species of sea
cucumbers and also provided data on their population density, habitat and ecology.
Nine of these species were recorded for the first time for Cuba.
After this, Suárez (1974) published his work ‘‘Lista de los equinodermos
cubanos recientes’’ (‘‘List of the recent Cuban echinoderms’’), which provided the
scientific name and synonymies of all species recorded for Cuba until that date.
It included all five classes of echinoderms and gave new records for the archi-
pelago. She listed 308 species of echinoderms (Crinoidea: 33 species, Asteroidea:
83 species, Ophiuroidea: 116 species, Echinoidea: 67 species, and Holothuroidea:
9 species). It also includes bibliographic references to these species for the Cuban
Archipelago and/or the region. This dissertation is considered the first scientific
work in which all the taxonomic information existing in Cuba on the Phylum
Echinodermata is integrated. It laid the groundwork for the beginning of the dif-
ferent studies that have been made until today.
It was not until the 1980s when the study of echinoderms began in a deeper
way. The pioneers in this enterprise where Ángela Corvea and Mercedes Abreu-
Pérez, under the guidance of Dr. Osvaldo Gómez, an outstanding pedagogue
responsible for the formation of numerous generations of biologists at the
Universidad de la Habana.
Dr. Pedro M. Alcolado, a specialist from the Instituto de Oceanología, also
made major contributions to the development of this special field in Cuba. In the
early 1990s, Dr. José Espinosa began study of echinoderms, although for a short
period of time. Four years later M. Sc. Rosa del Valle assumed this position.
Among the main research projects for systematics and/or ecology developed by
echinoderm specialists in the benthos department from the Instituto de Oceano-
logía were ‘‘Comunidades del Bentos de Punta del Este’’ (‘‘Benthos communities
from Punta del Este’’); ‘‘El Bentos de la Macrolaguna del Golfo de Batabanó’’
(‘‘Benthos from the macro-lagoon of Golfo de Batabanó’’); ‘‘Comunidades mac-
robentónicas asociadas a tres arrecifes coralinos del Golfo de Batabanó’’ (‘‘Macro-
benthic communities associated with three coral reefs from Golfo de Batabanó’’);
‘‘Caracterización del bentos de las Bahías de Jururú y Vita’’ (‘‘Characterization of
the benthos from Bahías de Jururú y Vita’’); ‘‘Protección de la biodiversidad y
desarrollo sostenible en el ecosistema Sabana-Camaguey’’ (‘‘Biodiversity protec-
tion and sustainable development in the Sabana-Camagüey ecosystem’’); and
‘‘Ecología y sistemática de las comunidades de equinodermos de las costas del
Caribe Mexicano y Cuba’’ (‘‘Ecology and systematics of echinoderm communities
from the Mexican Caribbean coasts and Cuba’’) under the framework of the Joint
Collaboration Agreement with the Instituto de Ciencias del Mar y Limnología,
from the UNAM, Mexico.
Other projects in which echinoderms have been studied are ‘‘La Biodiversidad
Marina de Cuba’’ (‘‘Cuban Marine Biodiversity’’); ‘‘Evaluación del deterioro
ocasionado por los vertimientos del emisario submarino de Playa el Chivo,
mediante el análisis de las comunidades bentónicas’’ (‘‘Evaluation of the damage
made by the dumping of the submarine emissary of Playa el Chivo, by means of
benthic communities analysis’’); ‘‘Evaluación general del estado ecológico de los
arrecifes de Cuba y monitoreo de la estación regional cubana de CARICOMP’’
(‘‘General evaluation of the ecological status of reefs in Cuba and monitoring of
the regional Cuban station of CARICOMP’’); ‘‘Hacia el uso sostenible del Golfo
de Batabanó’’ (‘‘Towards the sustainable use of the Golfo de Batabanó’’);
‘‘Análisis de sistema y modelación de escenarios’’ (‘‘System analysis and scenario
modeling’’); ‘‘Evaluación del Archipiélago de los Canarreos con vista a su cate-
gorización dentro del Sistema Nacional de Áreas Protegidas’’ (‘‘Evaluation of the
Archipiélago de los Canarreos with a view to its categorization within the Sistema
Nacional de Áreas Protegidas’’). As a result of participation in these projects,
numerous scientific papers, popular science works, scientific-technical reports and

many book chapters have been published.
Among the most relevant results obtained by the echinoderm specialists from the
benthos department are the works by Corvea (1986, 1990) that reports three new
records of holothuroids for Cuba: Euthyonidiella destichada (as Neophyllophorus
destichaudus), Synaptula hydriformis, and Holothuria thomasi. Corvea et al. (1990)
worked with the echinoderm communities from the Golfo de Batabanó macro-
lagoon and presented taxonomic and ecological results for echinoderms from the
gulf.
Abreu-Pérez (1983, 1987) published early works on new records of brittlestars
for the Cuban waters, including 14 species of shallow waters. Later, Abreu-Pérez
(1990) presented an updated list of Cuban ophiuroids that reported 158 species of
brittlestars, including seven new records. Espinosa et al. (1997) added new records
of three species of echinoderms: Davidaster rubiginosus, Astropyga magnifica,
Lytechinus variegatus carolinus. They also give data of bathymetric distribution
and exceptional sizes of four other species previously reported for Cuba: Eucidaris
tribuloides, Podocidaris sculpta, Moira atropos and Holothuria thomasi.
Abreu-Pérez (1997) published ‘‘Los asteroideos (Echinodermata) del Archi-
piélago Cubano’’ (‘‘The seastars (Echinodermata) of the Cuban Archipelago’’) that
updated the records of the seastar species of the coasts of Cuba. She reported 20
species, two of them being new records: Asterina harmeyeri and Ophidiaster
guildingi (= Linckia guildingi). In addition, she referred to previous works citing
the existence of the seastar species and their bathymetric ranges.
Abreu-Pérez and Del Valle (1998) recorded the sea urchin Lytechinus williamsi
for Cuba. That same year, Del Valle and Abreu-Pérez (1998) participated in the
National Study of Biological Diversity with a chapter related to the Phylum
Echinodermata that provided a list of all echinoderms recorded for Cuba and gave
the main characteristics of the group and ecological data of interest.
Ibarzábal et al. (1999) published a list of 26 species of echinoderms inhabiting
the Jardines de la Reina Archipelago. The class Ophiuroidea was the most diverse
with 14 species.
Abreu-Pérez et al. (2000) published an updated list of the sea urchins of the
Cuban Archipelago. Del Valle et al. (2000) recorded for the first time the family
Chiridotidae for Cuba with the species Chiridota rotifera. Fernández (2001) made
a new contribution for the class Asteroidea with the species Copidaster lymani.
Del Valle et al. (2003) presented the Final Report of the ‘‘Inventory of marine
flora and fauna from the Sabana-Camagüey Archipelago’’ that included the list of
echinoderm species present in the northern part of the providences of Matanzas,
Sancti Spiritus, Villa Clara, Ciego de Ávila, and Camagüey (Archivo Científico
del Instituto de Oceanología). That same year, Abreu-Pérez (2003) published five
new records of brittlestars for Cuba: Ophiolepis gemmata, Ophiopsila hartmeyeri,
Ophionereis vittata, Ophioderma ensiferum, and Ophiactis quinquerradia,
extending the bathymetric range for some of them and increasing the number of
known species of brittlestars for Cuba. The next year, Del Valle and Barreras-
Rodríguez (2006) reported the sea cucumber Holothuria princeps for Cuba.
In the book ‘‘Corales Pétreos: Jardines Sumergidos de Cuba’’ (‘‘Stony corals:

underwater gardens of Cuba’’) Del Valle and Abreu-Pérez (2004) give a general
view of the Phylum Echinodermata for Cuban waters, listing the main species of
echinoderms associated with stony corals in Cuba. A year later, Abreu-Pérez et al.
(2005) and Del Valle et al. (2005) published the catalogue of neritic seastars and
brittlestars and a catalogue of neritic sea lilies, sea urchins, and sea cucumbers of
the Cuban Archipelago, respectively. In both, they listed 125 species for the neritic
zone (down to a depth of 30 m), the bibliographic references that cite them for the
country, habitat, depth, distribution in the Western Atlantic, and the collection
localities in Cuba. The catalogue of Del Valle et al. (2005) reported two new
records of holothuroids for Cuban waters: Ocnus suspectus and Phyllophorus
(Urodemella) occidentalis.
The next year, Del Valle and Barreras-Rodríguez (2006) recorded the holo-
thurian Holothuria princeps for Cuba. Soon after, Rodríguez-Barrera et al. (2007)
contributed a new record of an echinoid for Cuba, Paraster doederleini, increasing
the number of species and extending the knowledge on the geographic distribution
for this group.
In the book ‘‘ Marine Biodiversity of Cuba’’, Del Valle and Abreu-Pérez (2007)
present a taxonomic list of all echinoderm species recorded for Cuba, where the
distribution of each one is given by ecozones (Areces 2002), as well as the relative
abundance of them. For each class, the most remarkable species for each ecosystem
present in Cuba are highlighted. In this work they analyzed the status of exploitation
of the sea cucumber Isostichopus badionotus in the Cuban Archipielago.
In the book ‘‘Sabana-Camagüey Ecosystem: current state, progresses and
challenges on the protection and sustainable use of Biodiversity’’, Del Valle et al.
(2008) listed 1,370 marine species inventoried in the archipelago. Of these, 1,064
are invertebrates with 71 species of echinoderms. In addition, the areas with the
upper and lower species richness values are pointed out. Pertinent recommenda-
tions for protection and better knowledge of the marine fauna of the Sabana-
Camagüey Archipelago are given. This study found that in the outer zone of the
macro-lagoon, where human impact is minor, the seagrasses develop better, the
echinoderm communities have a greater number of species and moderated popu-
lation densities. This was associated with a reduction in percentage of suspension-
feeders. In contrast, towards the inner side of the macro-lagoon, where there is
little exchange with oceanic waters, seagrasses are rare or absent and the organic
matter and silt values are high. Fewer echinoderm species occur and a greater
percentage of invertebrate filter-feeders are present.
12.2.1 Other Studies
Echinoderms have been used by other specialists, in different fields of study, for
research. Elyakov et al. (1975) made a comparative and chemical composition
study of holothurins (glycosides) isolated from nine species of sea cucumbers:
Astichopus multifidus, Isostichopus badionotus, Actinopyga agassizii, Holothuria

mexicana, H. cubana, H. arenicola, H. grisea, H. surinamensis, and Euapta lappa.
Gómez et al. (1980) evaluated the macrobenthos of the Golfo de Batabanó.
They provide ecological data of interest related to echinoderms. Berovides-
Alvarez and Ortíz-Touzet (1981) studied color polymorphism in the seastar
Oreaster reticulatus, finding four patterns of coloration: olive green, red, reddish
brown and creamy yellow. They concluded that the most frequent pattern is the
reddish brown with 49.2 % of presence and the olive green pattern was frequent in
juveniles. Herrera-Moreno et al. (1981) used random sampling to study popula-
tions of the sea urchin Diadema antillarum in front of the rocky zone of the
Instituto de Oceanología. Espinosa (1982) analyzed the stomach contents of the
predatory seastar Astropecten articulatus and A. duplicatus. Martínez-Estalella
(1984) contributed a list of organisms associated with coral reefs. Beltrán et al.
(1988) used the sea urchin Echinometra lucunter as an indicator of oil pollution in
the rocky littoral adjacent to Bahía de la Habana. Ablanedo et al. (1990) used the
same species as an indicator of heavy metals (iron, zinc, manganese, and mercury)
pollution. They measured the concentration of these metals in sea urchins gonads
and skeletal parts (test, Aristotle’s lantern). The results obtained showed that heavy
metals accumulate differentially in different body tissues: iron and mercury in
gonads, zinc and manganese in skeletal parts. They also observed that accumu-
lation of heavy metal showed seasonal variation.
Ortea et al. (1991) pointed out the 30 most conspicuous species of echinoderms
inhabiting the Cuban Archipelago. These included Luidia senegalensis, A. duplic-
atus, Linckia guildingii, O. reticulatus, Astrophyton muricatum, Ophiocoma riisei,
Ophiothrix suensonii, D. antillarum, E. tribuloides, E. lucunter, E. viridis, Tripn-
eustes ventricosus, L. variegatus, Clypeaster rosaceus, Leodia sexiesperforata,
Meoma ventricosa ventricosa, Holothuria cubana, H. arenicola, I. badionotus and
Stichopus regalis.
Santana et al. (2001) documented the ingestion of a specimen of D. antillarum
(30 mm test diameter) by the sea anemone Condylactis gigantea in an aquarium.
Six stages were involved in this process during a period of 24 h. All of the skeletal
material of the sea urchin was discarded by the anemone at the end of digestion.
Castellanos-Iglesias et al. (2004) performed the ‘‘Reef Check Monitoring’’ at
‘‘Rincón de Guanabo’’ reef in la Habana. As part of the results, they showed that
the population density of D. antillarum was extremely low in this reef.
Nodarse-Konnorov (2001) analyzed the ecological structure of E. lucunter
populations in a coastal reef at Ciudad de la Habana, using a stratified random
sampling. The goal was to detect possible natural regularities in abundance and
size of the E. lucunter over space and time. The authors found that anfractuosity
was larger in the middle strata (5–15 m away from the shore), therefore containing
the major refuge availability, and with greater densities per month. Depth, an
indicator of the hydro-physical stress, increased farther from shore. Smaller
individuals were collected in the deeper strata during September and November.
For this reason, it was concluded distribution and size of individuals were related.
Ortiz (2001) provided a list of marine, estuarine and semi-terrestrial inverte-

brates of Playa Cojimar, located in the northern coast of the Province Ciudad de la
Habana. It showed that several species, Oreaster reticulatus and Arbacia punc-
tulata among them, were starting to disappear during a period of 40 years of
sampling in that locality.
Later on, De la Guardia et al. (2003) published a work on marine biodiversity in
the coastal lagoon ‘‘El Guanal’’, Cayo Largo, Cuba. They made an inventory of the
flora and fauna of this site. Four echinoderm species were identified: O. reticulatus,
D. antillarum, Tripneustes ventricosus, and Lytechinus variegatus. The authors
concluded that none of the species identified during this study is exclusive to this
zone, nor in danger of extinction or overexploitation. Despite the intense nautical
activity and dredging work prior to this study, the system’s characteristics have
allowed the maintenance of a phase, undistinguishable from natural situations.
Due to the importance of the results obtained by the joint expeditions from
Harvard and La Habana Universities, on board the B/I ‘‘Atlantis’’ during 1938 and
1939; as well as for the difficult access to the original literature on the subject,
Lalana et al. (2004) published a taxonomic list (with bathymetric distribution) of
371 species and 5 subspecies of mollusks, crustaceans and echinoderms from that
expedition (31 species and one subspecies were echinoderms).
Recently, Martín-Blanco et al. (2011) determined the abundance, distribution,
and size structure of Diadema antillarum and the influence of their populations on
the community structure of algae in Jardines de la Reina Archipelago.
Three hundred and fifty seven species of echinoderms have been recorded for
Cuba. The Class Crinoidea is represented by 34 species (four orders, 19 families,
and 21 genera), the Class Asteroidea by 76 species (six orders, 23 families, and 50
genera), the Class Ophiuroidea by 163 species (two orders, 14 families, and 61
genera), the Class Echinoidea by 63 species (13 orders, 23 families, and 35 gen-
era), and the Class Holothuroidea by 51 species (five orders, 12 families, and 24
genera).
The 34 crinoid species recorded for Cuba are distributed from the littoral zone
down to the great depths, as in the species Coccometra hagenii which was col-
lected at 1,046 m depth. Only eight species of crinoids have been recorded at
depths below 30 m. Conspicuous by their apparent frequency in coral reefs are the
species Davidaster rubiginosus and N. discoideus.
The Class Asteroidea is represented in Cuban waters by 76 species; of these,
only 20 (26 %) are distributed down to a depth of 30 m. Conspicuous by their
apparent frequency in coral reefs are Linckia guildingii and Ophidiaster guildingi.
Whereas, in sandy substrates with macro-vegetation, the most frequent and
abundant seastar is O. reticulatus. In bays and coastal lagoons, where sand-mud
substrates predominate mostly without vegetation, the nine-arm seastar Luidia
senegalensis occurs frequently. In beaches or sandy plains Astropecten duplicatus

can be observed, with relative frequency. The 56 species left are distributed in the
deeper zones. Conspicuous amongst them are Benthopecten spinosus and Pteraster
aciculata, the first recorded at depths between 1,300 and 3,700 m and the second
at depths between 260 and 3,712 m.
The Class Ophiuroidea is the best represented in all coastal ecosystems of the
Cuban Archipelago, both in species number (163) and appearance frequency and
abundance. Of the total of recorded species, 55 have been observed at depths
shallower than 30 m. Conspicuous in coral reefs are the genera Ophiomyxa,
Astrophyton, Ophiocoma, Ophioderma, and Ophiothrix. In particulate substrates
with macro-vegetation, stand out, by its appearance frequency, the species Ophi-
olepis paucispina and Ophionereis reticulata. In particulate substrates without
vegetation in bays, coastal lagoons, beaches or sandy plains, Amphiodia trychna
and Amphipholis gracillima are the most frequent.
In semi-enclosed areas with limited water circulation and with major input of
organic matter, as is the case for Bahía de Santa Clara (northern-central Cuba),
very high densities of the species Ophiothrix suensonii (up to 8.84 ind m-2) and
Ophiothrix angulata (up to 7.57 ind m-2) have been recorded. Both are associated
with macroalgae of the genera Laurencia. In the mangrove swamps of Cuba, the
brittle stars species Ophiopsila riisei and Ophionereis olivacea are recorded with
greater frequency It is interesting that the only species of brittle star that has been
recorded in the rocky littoral is Ophioderma brevicadum and the two species
recorded at greatest depths are Ophiomusium lymani and Homolophiura abysso-
rum (584 and 983 m, respectively).
Of the 63 echinoids recorded for Cuba, the best represented orders are
Cidaroida, with nine species, Clypeasteroida, also with nine species, and Echi-
noidea with seven. Twenty two species have been observed at depths less than
30 m. Conspicuous, both by its appearance frequency and abundance (in density
and biomass), are E. lucunter, E. viridis, E. tribuloides, T. ventricosus and
D. antillarum.
In particulate sand-mud substrates with macro-vegetation, Meoma ventriciosa
and Clypeaster rosaceus have been recorded in high densities. The latter is also
abundant in sandy bottoms. In muddy substrates without macro-vegetation in bays
and coastal lagoons, the irregular echinoid Moira atropos is the best represented.
On beaches and/or protected sandy plains Leodia sexiesperforata is the most
frequent and abundant. It must be emphasized that prior to the 1980s, D. antil-
larum was very abundant in the rocky littoral of Cuba, where it occurred along
with E. lucunter. However, after the mass mortality of this sea urchin, E. lucunter
became the most frequent and abundant species of this ecosystem. The deepest
records for the Phylum Echinodermata are for Pourtalesia miranda (450–5,850 m)
and Aceste bellidifera (550–5,220 m).
Among the 51 species of holothurians recorded for Cuba, 26 inhabit depths
greater than 30 m. The other 25 species are distributed in the different ecosystems
present on the island platform. In coral reefs, 10 species are recorded, of which
Euapta lappa, Holothuria impatiens, and H. thomasi inhabit exclusively in this
ecosystem. Others, like the sea cucumber Actinopyga agassizi, have been found
with relative frequency in coral reefs. It can also be collected with some frequency
in seagrasses associated with rocks and stones. Holothuria thomasi has been
observed with less frequency. In seagrasses, the most frequent and abundant
species are H. mexicana and H. floridana. In bays and coastal lagoons I. bad-
ionotus can be found at times with very high densities. Holothuria arenicola is
common on beaches and/or sandy plains.
12.4 Ecology
The Workshop of Eco-regionalization and classification of marine habitats in the

Cuban platform, celebrated in 2002, brought together Cuban and foreigner spe-
cialists in different fields of study. They undertook the task of analyzing the
geographic, geomorphologic, and ecological characteristics of the Cuban platform,
as well as the distribution of its biotopes. The main objectives of this workshop
were to identify the regions that form the marine platform of the Cuban Archi-
pelago and to determine the main biotopes of the Cuban marine ecosystem. As
result of the analysis, nine ‘‘ecoregions’’ were established (Areces 2002)
(Fig. 12.1), named later by Claro (2007) as ‘‘ecozones’’. It was stated that the main
biotopes of the marine ecosystem in Cuba are coral reefs and hard non-reef bot-
toms (from interior waters); non-consolidated sediments (sand, mud); submerged
vegetation (seagrasses and macroalgae, mangrove swamps, coastal lagoons and
estuaries, low rocky coasts or with cliffs, and beaches).
Among the ecological studies performed in the island platform of Cuba are the
major ones made in Punta del Este (at La Isla de la Juventud, ecozone 4), and the
Golfo de Batabanó macro-lagoon (ecozone 4), the echinoderm communities of
Sabana-Camagüey Archipelago (ecozone 8), of Jururú and Vita bays (ecozone 9),
of Jardines de la Reina Archipelago (ecozone 2) and the northern coast of Habana-
Matanzas (ecozone 7) (Fig. 12.1).
An integrative analysis of the ecological results for the Golfo de Batabanó
macro-lagoon, Sabana-Camagüey Archipelago (ecozones 4 and 8, respectively)
(Fig. 12.1), and the Jururú and de Vita bays (ecozone 9) showed all were domi-
nated by unconsolidated sediments (sand and mud) and submerged vegetation
(seagrasses and macroalgae). Both biotopes are present in different proportions
(sand-mud, seagrasses-macroalgae), according to the oceanographic and geo-
morphologic characteristics of each locality.
At Golfo de Batabanó (ecozone 4) (Fig. 12.1), were recorded 50 species of
echinoderms (33 brittle stars, four seastars, seven echinoids, and six holothurians).
The most important species of brittle stars, due to their wide distribution within
that macro-lagoon were Ophioderma olivaceum, Ophiolepis elegans, and Ophio-
stigma isacanthum (Corvea et al. 1990). The richest and most diverse sites cor-
responded to those zones where appropriate geomorphologic and oceanographic
conditions exist for the development of macro-vegetation and the establishment of
diverse and abundant echinoderm communities. Whereas, in some interior zones,

where muddy bottoms without vegetation predominate, very few or no species
were recorded. The dry biomass values varied between 0 and 4.72 g m-2. The
highest ones corresponded to ‘‘Cayo Largo del Sur’’ zone, part of the Northwestern
zone of ‘‘Isla de la Juventud’’ (ecozone 4) (Fig. 12.1). The poorest zones in bio-
mass were the Southeast part of Golfo de Batabanó, ‘‘La Broa’’ inlet (ecozone 1)
(Fig. 12.1), which suffers from strong erosion in the northern muddy coasts, due to
disappearance of the protective red mangrove. This produced a marked and per-
manent water turbidity that has eliminated the seagrasses throughout the west part
of this inlet where some benthic organisms were once abundant (Alcolado 2007).
All over the macro-lagoon, the dominant species in biomass were the sea urchin
L. variegatus, the seastar O. reticulatus, and the irregular echinoid C. rosaceus.
This study showed L. variegatus plays an important role in seagrasses from Golfo
de Batabanó, because is an active and abundant micro phytobenthos consumer and
represents the main biological energy mobilizing mesh in the ecosystem (Alcolado
1990).
The mega-benthic communities of the Sabana-Camagüey Archipelago (ASC,
ecozone 8) were analyzed using a 109 stations network and numerous oceano-
graphic parameters including temperature, current, salinity, particulate organic
matter and silt content in sediments (Espinosa and Abreu-Pérez unpublished data;
Alcolado et al. 1998). Echinoderms represented by 53 species (29 brittle stars, 12
holothurians, seven echinoids, and five seastars) were recorded in 71 stations just
like the Golfo de Batabanó macro-lagoon, the dominant species in number in most
stations (18) and with the largest frequency of appearance (32 stations), was
L. variegatus. Other conspicuous species in terms of numbers were the brittle stars
were evident: O. angulata (seven stations), Ophiostigma isacanthum, Ophioderma
olivaceum, Ophiothrix orstedii, and O. angulata. These also were important for
being recorded in 20, 19, 18, and 12 stations, respectively.
It is interesting to note the high abundance in number in which the brittle star
O. angulata occurred. Over two thousand specimens were collected in 1 min of
sampling. This seston-feeder species was found on the red algae Acanthophora sp.
at a station located at the western of Bahía de Santa Clara, where the current
provoked by predominant winds (from east to west) accumulates a larger amount
of nutrients, producing the so called ‘‘pocket’’ effect which favors this kind of
organism (Alcolado 1990).
The greatest species number, as well as the highest Margalef’s Richness Index
(1951) (R1 = 3.9), was found at station 36. This station is located in an external
area where good exchange of water with the ocean exists and the bottom is covered
with seagrass. When the 109 stations are combined, taking into account the biotope
type, the greatest species number corresponded to seagrass growing on thick
aleurite. In addition, it was observed that the brittle stars I. isacanthum, O. angulata,
and O. oerstedii were the species appearing on a larger numbers of biotopes.
The effect of salinity was also analyzed. Of the 53 potential species existing in
the Sabana-Camagüey Archipelago at salinities of 36 %, the number is markedly
reduced as salinity rises, down to 10 species at 41 % and only one, the brittle star
Ophiolepis elegans, at a salinity of 42.2 %. This is the most salinity-tolerant

species reported in Cuban waters.
A major variation in particulate organic matter content was found within the
sediments of the interior parts of Sabana-Camagüey Archipelago. The greatest
species numbers (33–36) were found at stations with low-medium content of
organic matter (2.5–15.5 %). The species number was greatly reduced (11–5) in
places with high-medium percentages (15.6–35.0 %) (Espinosa and Abreu-Pérez
unpublished data). This observation is very similar to that found by Alcolado et al.
(1998) for all mega-zoobenthic organisms he studied.
The sites with greater species richness were those with good water circulation
and exchange of interior waters with the open sea, presence of good development
of macro-vegetation, salinity values lower than 38 %, and moderate particulate
organic matter and silt values. The highest species density values were recorded in
regions where the geographic configuration favors particle retention (pocket effect)
(seston and larvae from benthic colonizers), provided that stress factor values were
at the levels indicated above.
In a general sense, the results from studies at Bahías de Jururú and Vita
(ecozone 9), Cárdenas and Nuevitas (ecozone 8) (Fig. 12.1) showed the same
pattern observed in Sabana-Camagüey Archipelago.
Comparison of the echinoderm list of ASC with the Golfo de Batabanó macro-
lagoon and Bahías de Jururú and Vita, using Sorensen’s Similarity Index (1948),
shows that a high faunistic affinity exists. Integration of the echinoderm species
lists from all of the studied sites in Cuba indicates that the echinoderm commu-
nities present in soft bottoms total 91 species. Fifty six are brittle stars, 18 holo-
thurians, nine echinoids, and eight seastars.
Another relevant ecological study was performed on three coral reefs in Golfo
de Batabanó (Espinosa and Abreu-Pérez unpublished data) (Fig. 12.1). The main
characteristics of echinoderm communities present were analyzed in different
zones (reef lagoon (L), rear zone (ZT), plain (M), pounding zone (ZE), abrasive
rocky esplanade (ER) at 5 m depth, and external slope (PE) at 10, 15, and 20 m).
The ‘‘Juan García’’ reef, located in the western part of Golfo de Batabanó, is the
most exposed of the three reefs. It receives a strong influence from enriched water
with organic matter from Isle of Youth and the numerous adjacent cays and little
cays. The ‘‘Cantiles’’ reef is located south of Golfo de Batabanó, in a more
protected zone and is the one receiving the least land influence. The ‘‘Diego
Pérez’’ reef, located east of Golfo de Batabanó, is the most protected one and
receives a major land influence from Península de Zapata, a region rich in swamps
and mangroves. Major differences were found among the three reefs in the eco-
logical structure of echinoderm communities, as well as between the zones that
composed them (coastal lagoon, reef plain complex, abrasive rocky esplanade, and
external slope at 10, 15, and 20 m)
The number of species in ‘‘Juan García’’ reef was less than in the other two
reefs, except on the external slope. These results have an important relationship
with different environmental factors acting negatively on the ecological structure
of echinoderm communities. Among them are the greater degree of exposure to
Fig. 12.2 Echinoderm

species distribution in the
different zones at the coral
reefs (Juan García, Cantiles,
and Diego Pérez) from Golfo
de Batabanó. L reef lagoon,
ZT rear zone, M plain, ZE
pounding zone; ER rocky
esplanade at 5 m depth; and
PE external slope at depths of
10, 15, and 20 m
waves and winds present. This became evident with the high values recorded by
the Agitation Index (IA) by Alcolado (1990): rear zone (IA = 98), pounding zone
(IA = 100), and abrasive rocky esplanade (IA = 99). These harsh conditions,
along with the major input of organic matter received from Isle of Youth and the
numerous adjacent cays and little cays, create a constant and severe re-suspension
of sediment in these zones. This stress factor may turn out to be intolerable for
many echinoderm species. In this reef, brittle star species predominated. In some
stations, only this class was recorded, especially in the Rear Zone-Plain-Pounding
Zone complex (ZT-M-ZE), where the species Ophiocoma echinata and Ophio-
coma riisei dominated. However, on the external slope (at 15, 19, and 20 m depth),
where conditions are not so severe, a remarkable increase in species number was
observed. This also happened in Cantiles and Diego Pérez reefs (Fig. 12.2).
Valdivia et al. (2004) described representative biotopes of coral reefs from the
southern zone of Guanahacabibes Peninsula (western edge of Cuba) and a taxo-
nomic list of marine organisms identified in the zone during 2001. This zone is
characterized by the greatest concentration of reef-former organisms and fish in
three biotopes: rocky plain with coral and gorgonians (down to 2–4 m), ‘‘cabezos’’
(5–8 m), and ‘‘camellones’’ (ridges) (12–15 m). At a depth of 3 m, and forming
bands 20 m wide, sea urchins E. lucunter and E. viridis form a characteristic
biotope of this zone. In addition, 109 algal species, 23 sponges, and 86 fish species
typical of coral reefs were identified. They argued that excessive overfishing may
be one cause of large fish in the area.
Since 1999, the sea cucumber I. badionotus has been exploited in Cuba. This
activity is in the charge of the Ministerio de la Pesca (Fishing Ministry) (MIP).
During the first 2 years of fisheries (1999–2000), more than 3 million individuals
were captured. It seems that, as result of these large extractions, catches and
production decreased from 1,153 sea cucumbers boat-1 d-1 in 1999 to 350 sea
cucumbers boat-1 d-1 in 2002 (Alfonso et al. 2003).
Due to major decrease in resources in zones like Bahía de Banes, Banco
Chinchorro, and Cayos de Manzanillo, the MIP established several regulations.
These included a fishing ban in affected regions, fishing effort limits regarding to
previous evaluations by zone, minimal size of 220 mm total length, a fishing ban
during the peak months in reproduction (February–April), and a rotation system of
catch areas. As a result of these measures, a resource recovery was observed in
2005 in every fishing zone in the country, including the most affected ones
(Alfonso 2004; Alfonso et al. 2004, 2005, pers. comm. R. Castelo).
12.6 Threats
The main threats affecting echinoderms of the Cuban Archipelago are closely
related to habitat destruction, fundamentally caused by coastal development and/or
natural factors (e.g. hurricanes, cyclones, etc.).
The known effect in Cuba is related to the sea urchin D. antillarum populations.
The mass mortality of D. antillarum has been widely documented, not only in
Cuba but all along the Caribbean region (Hughes et al. 1999; Alcolado et al. 2004).
In many reefs from the Cuban Archipelago, an excessive growth of algae has been
observed (Habana littoral; Sabana-Camagüey Archipelago; Largo, Paraíso, and
Levisa Cays; and nearby Bahía Honda). This seems to be, in general terms, due to
the mass mortality of D. antillarum and also because of the obvious low numbers
of herbivore fishes.
Alcolado et al. (2004) studied population densities of D. antillarum in 199 sites
at different reef ridges from the Cuban Archipelago. They showed that, even
though D. antillarum existed in some sites in ecozones Jardines de la Reina
(0.925–21.1 ind m-2) and in Golfo de Batabanó (0.58 a 1.17 ind m-2) with
densities ranging from moderate to very high, the recovery of populations of this
species is happening slowly. Caballero et al. (2007) mentioned that D. antillarum
density in ‘‘María La Gorda’’ reef (Bahía de Corrientes) was very low
(3.5 ind 10 m-2).
Martín-Blanco et al. (2011) presented the results of a study about abundance,
distribution, and size structure of D. antillarum in 22 frontal reefs from Jardines de
la Reina Archipelago. The authors found that D. antillarum densities were large in
reef ridges (0.08–2.18 ind m-2) and decreased down to three orders of magnitude,
with a maximum density of 0.13 ind m-2 on the slope. These authors concluded
that the size frequency distribution of D. antillarum suggests that recruiting began
to be evident approximately 3 years prior to the sampling; indicating that their
populations were affected during the past and are now recovering.
Another species that seems to have been affected in some way is the sea urchin
A. punctulata, for it was highly abundant in the northern coast of Cuba, especially
in Bahía de Cárdenas (pers. comm. M. Ortiz). However, it has not recorded for
over 20 years, despite numerous exploratory samplings in that bay. Moreover, it
was never observed during the several samplings made in the Sabana-Camagüey
Archipelago. Nevertheless, some specimens have been observed recently at
Provincia de Villa Clara (ecozone 8). High densities have being observed of
A. punctulata in the north and south coasts of Península de Guanahacabibes
(ecozones 5 and 6, respectively) (pers. comm. R. Arias and M. Esquivel).
12.7 Conclusions and Recommendations
The Phylum Echinodermata is among the best studied faunistic groups in the
Cuban Archipelago with observations from the seventeenth century until the
present. However, knowledge of this group differs among classes. The least
studied is Class Crinoidea. There are remarkable differences in knowledge among
the platform’s nine ecozones. These results will contribute to the knowledge of
echinoderm diversity in Cuba, and will be of practical and teaching purposes for
scholars interested in the subject.
One recommendation is to increase the scientific formation of young men and
women specialized in this group with the aim of ensuring the continuity of studies
related to the Phylum Echinodermata. New projects and lines of investigation
(genetics and molecular amongst them) should be developed. It is necessary to
develop a database and an information system on Cuban marine biodiversity,
including echinoderms, and to regulate and promote its use. Existing laws pro-
hibiting echinoderm extraction for ornamental purposes without prior impact
studies should be effectively enforced. Environmental education, awareness, and
citizens’ involvement should be promoted so that the population, especially
children and young people learn about echinoderm species inhabiting our seas.
Acknowledgments We want to express our gratitude to Cynthia Ahearn and David Pawson
(United States National Museum of Natural History, Smithsonian Institution) and Gordon Hen-
dler (Los Angeles County Museum) for they support and for always enriching our minds with
their wise knowledge. We also thank the Department and Laboratories of the Oceanology
Institute of La Havana, Cuba (Oceanology Department and Laboratory), for their unconditional
support during all these years. Without them this work would have been impossible.
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Chapter 13
Recent Echinoderms from Hispaniola
Alejandro Herrera-Moreno and Liliana Betancourt
13.1 Introduction
The Island of Hispaniola is the second largest in the Archipelago of the Greater
Antilles. It is located between 17°360 1500 N and 19°570 0900 N, and 68°190 2200 W and
74°410 3300 W. Its geographical boundaries are the Atlantic Ocean to the north, the
Caribbean Sea to the south, the Mona Channel to the East, which separates it from
Puerto Rico, and in the West, the Windward Passage and the Jamaica Channel,
which separate it from Cuba and Jamaica, respectively. The 76,480 km2 of its
territory is divided politically into two countries: Dominican Republic to the east
and the Republic of Haiti in the west, separated by a land border of 360 km.
The Republic of Haiti has a total area of 27,750 km2, which includes the
mainland territory plus several islands and islets such as Gonave (743 km2),
Tortue (180 km2), Vache (52 km2), Cayemites (45 km2) and Navassa (5.2 km2).
The Dominican Republic has an area of 48,442 km2 of mainland in addition to
several islands and islets such as Saona (117 km2), Beata (27 km2) and Catalina
(9.6 km2). Two oceanic banks: Christmas and Silver Banks, with areas of 70 and
150 km2 respectively, are located to the north of Samana in the Atlantic Ocean.
The coastline of Hispaniola extends for about 3,059 km (1,288 km belong to 16
Dominican coastal provinces and 1,771 km to nine Haitian coastal provinces)
containing ecosystems such as sandy beaches, rocky shores, cliffs, estuaries,
coastal lagoons and mangrove forests. On the Haitian coast important mangroves
areas are reported along the north and northeast coast in the Bays of Fort Liberté,
l’Acul, Caracol, the Artibonite estuary and the Islands Les Cayes, Vache, Gonave
and Cayemite (Ehrlich et al. 1987). Along the Dominican coast the largest areas of
mangroves are found in the Bay of Samaná and Montecristi.
A. Herrera-Moreno (&) L. Betancourt

Programa EcoMar, Inc., Santo Domingo, Dominican Republic
e-mail: ongprogramaecomar@yahoo.com

426 A. Herrera-Moreno and L. Betancourt
The Dominican shelf has a surface area of 8,130 km2 extending down to 200 m
depth, with an average width of 7.5 km. The platform area of Haiti is estimated at
6,683 km2 and is on average narrower. Important coastal habitats include sandy or
muddy bottoms (related to the degree of terrigenous influence), commonly pop-
ulated with abundant macrophytobenthos, especially seagrasses, followed by coral
patches and fringing or barrier reefs. In Haiti coral reefs occur near Vache Island,
all around Gonave Island in the central Bay of Port-au-Prince, on the Rochelois
Bank, at Cayemites Islands, off the northern coast of the southern peninsula, and
from the border with the Dominican Republic in the east to Acul Bay just west of
Cape Haitien (Creary et al. 2008). Well preserved reefs are found at Navassa
Island (Causey et al. 2000).
In the Dominican Republic important reef areas include the Montecristi
National Park barrier reef in the north-west (where the shelf is widest), narrow
high-energy reefs in the central region, and the Bávaro-Macao-Punta Cana barrier
reef system at the eastern end. Samaná Bay receives many rivers and is the largest
estuary of the insular Caribbean. Coastal reefs in the vicinity are poorly developed,
but the Christmas and Silver Banks reef systems are about 100 km to the north. To
the south, on the Caribbean coast, are the well-studied reefs of National East Park
and the adjacent Saona Island. Uplifted carbonate terraces with reefs growing on
narrow platforms are present in the west from Catalina Island to beyond Santo
Domingo (e.g. Boca Chica and the Submarine National Park La Caleta). Condi-
tions are not good for reefs in the south-west, except on the shallow sheltered shelf
east of Cabo Beata at National Park Jaragua (Woodley et al. 2000).
13.2 Research
Echinoderm studies of the waters around Hispaniola Island have a long history.
The Haitian collection of David Friedrich Weinland from 1857 (NMNH 2002) and
the work on holothuroids by Emil Selenka (1867), with the record of Actinopyga
agassizi (Selenka, 1867) in Haiti and the description of the type specimen of
Holothuria grisea Selenka, 1867, are among the earliest contributions.
William More Gabb made collections in the Dominican Republic in 1878 and
deposited several specimens of Echinometra lucunter (Linnaeus, 1758) in the U.S.
National Museum of Natural History (Rathbun, 1886). In December 1878, during
the Blake Expedition, the echinoid Salenocidaris varispina (A. Agassiz, 1880) was
collected northwest of Haiti at 2,195 m depth (A. Agassiz, 1880). Dodërlein and
Hartmeyer (1910) and Verrill (1915) summarized previous records of asteroids
from Hispaniola. Hubert L. Clark (1919) provided the first taxonomic summary of
Haitian echinoderms listing more than 20 species of all classes, except crinoids.
In the 1930s, new contributions to the knowledge of echinoderms from
Hispaniola include Deichmann (1930) who summarized previous records for the
Dominican Republic and Haiti in her review of the holothurians of the western part
of the Atlantic Ocean. The handbook of the littoral echinoderms of Porto Rico and
13 Recent Echinoderms from Hispaniola 427
the other West Indian Islands (H.L. Clark 1933) also summarized historical
information about all classes of echinoderms for Hispaniola, except crinoids, with
some new records. Austin H. Clark (1939) listed 19 species of shallow-water
echinoderms (between 1 and 2 m depth) collected in two localities in the north of
Haiti (Cape Haitien and Tortue Island) during the Smithsonian-Hartford expedi-
tion of March 1937, aboard the RV Joseph Conrad.
New collections took place during the 1930s on five expeditions: Parish-
Smithsonian in 1930 to Haiti on the yacht Esperanza, Norcross-Bartlett in July 1931
to the southwest of Santo Domingo down to 200 m depth on the schooner Effie
Morrissey, Johnson–Smithsonian Deep-Sea in February 1933 on the Yacht Caroline
at depths from 26 to 40 m at twelve stations inside the Bay and around the Peninsula
of Samaná (Bartsch 1933), RV Stranger in February 1933 in Port-au-Prince and the
RV Atlantis in April 1939 dredging to 1,170 m deep at one station to the northeast of
Haiti (H.L. Clark 1941). These expeditions contributed new records for Hispaniola
including two deep-water asteroid species: Persephonaster patagiatus (Sladen,
1889) and Phormosoma placenta sigsbei (A. Agassiz, 1880). The expeditions of
1931 and 1933 collected the first specimens of crinoids: Poliometra prolixa (Sladen,
1881) and Nemaster rubiginosa (Pourtalés, 1869).
In the decade of 1960s two expeditions took place. The exploratory fishing
vessel Silver Bay in October 1963 made collections at six stations between 92 and
348 m deep in the north, northeast and east of the Dominican Republic. The
exploratory fishing vessel Oregon in June and May, 1965 and December 1969
collected between 11 and 59 m deep at four stations off the north Haitian coast and
the northeast Dominican coast. During this period, Parslow and Clark (1963)
summarized the zoogeographical distribution of shallow-water (less than 17 m
deep) ophiuroids known for the West Indies, including 11 species from Hispaniola,
updating the previous list of H.L. Clark (1919). Deichmann (1963) summarized
previous records of holothuroids for Haiti. Halpern (1969) mentioned the asteorid
Litonotaster intermedius (Perrier, 1884) found in the Windward Pass, between
Cuba and Hispaniola, at a depth between 1,958 and 3,294 m (Downey 1973).
From January to July 1970 and July 1971 the most prolific expeditions carried
out in Hispaniolan waters took place aboard the RV John Elliot Pillsbury.
Collections were made at seven stations in Haiti, between 31 and 2,545 m deep,
and 33 stations in Dominican Republic, between 9 and 3,109 m deep (Fig. 13.1)
(Staiger and Voss 1970). Meyer et al. (1978) included the RV John Elliot Pillsbury
collections in their zoogeographical study of western Atlantic Ocean crinoids,
offering the most complete summary of this group, with 19 species listed for
Dominican and Haitian waters. In her review of the Order Brisingida for
the Atlantic Ocean, Downey (1986) also incorporated Haitian specimens from the
Pillsbury expeditions. A.M. Clark (1987) described the type specimen of the
asteoridHenricia downeyae A.M. Clark, 1987 from a Haitian locality, registered in
the catalogue of types of the NMNH (Ahearn 1995). Two more records for Haiti
from this period were the holothuroid Psolus tuberculosus Théel, 1886 (Miller and
Pawson 1984) and the asteroid Ceramaster grenadensis grenadensis (Perrier,
1881) (Halpern 1970).
Fig. 13.1 Localities where collections of echinoderms have been made by several expeditions
and national and international projects on the shelf and oceanic waters of Hispaniola
All these expeditions increased the collections of the five classes of echino-
derms from Hispaniola in international museums with 541 specimens of 132
species located in the U.S. Museum of Natural History (NMNH 2002), Florida
Museum of Natural History (FLMNH 2010), Museum of Comparative Zoology at
Harvard University (MCZ 2010) and Museum of Natural Sciences of Berlin (ZMB
2010). The RV John Elliot Pillsbury expeditions had a decisive role in the
enrichment of collections, particularly with deep-sea species, that at the end of the
1970s, 90 % of the crinoid, asteroid, and echinoid species known from Hispaniola
were already described and deposited in different museums.
In the Dominican Republic, the creation of the Center of Marine Biology
Research (CIBIMA) in 1962 promoted coastal and marine biodiversity research.
Between the decades from 1970 to 1990, CIBIMA produced summarized list of
echinoids and asteroids (Cicero et al. 1976), ophiuroids (Rathe 1978) and holo-
thuroids (Briones 1983). Multiple reports were finally compiled into a preliminary
study on coastal and marine biodiversity of the Dominican Republic (CIBIMA
1992). From these investigations, 242 specimens of 58 species are deposited in the
National Museum of Natural History of Santo Domingo (MNHNSD 2008).
In May 1979, the Autonomous University of Santo Domingo, along with the
University of Puerto Rico, implemented an expedition on board the RV Crawford
to make inventories of the southeast Dominican reefs of La Caleta and Catalina
and Saona Islands (Williams et al. 1983). Later the IDECOOP expedition
(Fig. 13.1) made collections between 142 and 270 m deep with three new records
for the deep-sea Dominican echinoderms: the crinoid Cenocrinus asterius (Lin-
naeus, 1775); the ophiuroid Astronicda isidis Lyman, 1872 and the echinoid
Conolampas sigsbei (A. Agassiz, 1878) (Rivas 1983). Studies carried out by
researchers from East Carolina University in Montecristi reefs (Luczkovich 1991)
extended the reports of Dominican crinoids with the record of Tropiometra cari-
nata (Lamarck, 1816).
The information on Haitian echinoderms comes from old collections, that have
already been mentioned, and some more recent studies carried out by foreign
institutions. In June 1988, a detailed study at 13 stations on different ecological
zones (from the reef lagoon to the frontal reef) of the Arcadines coral reefs was
made by the World Wildlife Fund and the Conservation Foundation Wilcox
Associates, between 0.3 and 21 m depth (Wilcox et al. 1989). Twelve species of
common shallow-water echinoderms were reported. Hendler et al. (1995) sum-
marized the echinoderms from approximately 17 old Haitian records.
With the implementation of the Hispabiota Marina Project (Herrera-Moreno
and Betancourt 2012) by Programa EcoMar, Inc. in the Dominican Republic a
bibliographical and taxonomic review of echinoderm species was done for the first
time, with a historical and insular approach (Herrera-Moreno and Betancourt
2004). A database was created with the results of collections of echinoderms from
more than 150 localities of the coastal zone, the shelf and the deep zone of
Hispaniola (Fig. 13.1). This project added 60 species to the national inventory of
CIBIMA (1992) confirming 123 species for Dominican coastal and marine zone. It
also compiled for the first time 79 species for Haiti. A total of 156 species for
Hispaniola (Appendix) is discussed in the present report.
The numbers of echinoderms in different taxonomic categories for the Dominican

Republic, Haiti and Hispaniola are summarized in Table 13.1. There are 22 species
of crinoids (18 for the Dominican Republic and eight for Haiti), subdivided into 17
genera, nine families and four orders. About five species of crinoids are found in
coral reef environments at depths to 45 m deep including those which, according to
Hendler et al. (1995), are more common and accessible to conventional scuba
diving. The remaining species are distributed below 100 to 1,033 m deep.
The list of asteroids has 33 known species (18 for Haiti and 21 for the
Dominican Republic) subdivided into 24 genera, 11 families and seven orders.
Downey (1973) in her review of the asteroids in the Gulf of Mexico and the
Caribbean described 95 species and noted that the asteroid fauna from Hispaniola
was practically unknown. The bathymetric ranges of the asteroid species known
from Hispaniola varies from 0.3 to 3,493 m deep. About eight species are dis-
tributed from the surface to 50 m deep primarily from the mangroves to the reef
front, with the remaining species found below 50 m deep.
Table 13.1 Summary of the numbers of different taxonomic categories for echinofauna groups
of Dominican Republic (DO), Haiti (HA) and Hispaniola (HI)
Orders Families Genus Species
Crinoidea HA 3 4 7 8
DO 4 9 14 18
HI 4 9 17 22
Asteroidea HA 7 11 16 18
DO 4 7 14 22
HI 7 11 24 33
Ophiuroidea HA 2 9 12 21
DO 2 11 15 24
HI 2 12 20 30
Echinoidea HA 7 12 15 21
DO 10 16 31 41
HI 12 17 36 50
Holothuroidea HA 3 6 7 11
DO 3 5 7 18
HI 3 7 10 21
TOTAL HA 22 42 57 79
DO 23 48 81 123
HI 28 57 107 156
Thirty species of ophiuroids are known for Hispaniola (21 for Haiti and 24 for
the Dominican Republic), subdivided into 20 genera, 12 families and two orders.
Most species of ophiuroids are distributed in water less than 30 m deep found in
seagrass beds and coral reefs. Only four species were collected between 148 and
366 m deep.
Fifty species of echinoids are listed (21 for Haiti and 41 for the Dominican
Republic) subdivided into 36 genera (including one species identified only to the
genus level), 17 families and 12 orders. Although bathymetric data are not
available for all species, the collections from Hispaniola contain the more common
shallow-water species from mangrove habitats, seagrass beds and coral reefs. At
least about 30 species are distributed from 100 to 2,545 m depth.
The list of holothurians of Hispaniola includes 21 species (11 for Haiti and 18
for the Dominican Republic) with ten genera, seven families and three orders.
Most species of holothuroid are common in shallow sedimentary environments.
Three species are found between 243 and 1,400 m deep.
13.4 Ecology
In the Dominican Republic, echinoderm species are mentioned in the inventories

of invertebrates of seagrass beds and coral reefs from Manzanillo to Punta Rucia in
Montecristi (Luczkovich 1991; CIBIMA 1998), Puerto Plata (Herrera-Moreno and
Betancourt 2009), Samaná (Sang and Lysenko 1994; Sang 1996), Punta Cana,
Bávaro (CURPOB 2000; Brandt et al. 2003), National East Park (Vega et al. 1997)
and Isla Saona in La Altagracia, Catalinita in La Romana, La Caleta in Santo
Domingo (Williams et al. 1983), Haina in San Cristóbal (Herrera-Moreno et al.
2009), Las Calderas Bay in Peravia (Almonte 1976), Puerto Viejo in Azua
(González et al. 1978) and Jaragua National Park in Pedernales (Weil 2006).
In Haiti, inventories with echinoderm species include Lamentin (Beebe 1928), Les
Arcadins reefs (FoProBIM 1985; Wilcox et al. 1989) and Navassa (Miller 2003).
Due to the dramatic reduction of black sea urchin Diadema antillarum in the
Caribbean in the 1980s, the presence of this species is always highlighted in all
inventories and some abundance surveys have been done. In the Dominican reefs,
Chiappone (2001) estimated an average density of D. antillarum of 0.03 ind
50 m-2 for the coral reefs in the National East Park, between 9 and 17 m depth.
He also estimated 9.15 ind 50 m-2 in the Boca Chica reefs between 4 and 20 m
depth. Brandt et al. (2003) reported a maximum density of 5.4 ind 10 m-2 in the
reefs of Punta Cana and Bavaro with the highest average values in the deep fore-
reef (1.2 ind 10 m-2) in relation to the shallow areas of the back-reef (0.5 indi-
viduals/10 m2). For Caracol Bay in Haiti, Hay (1984) reported 20 ind m-2,
between 3 and 10 m depth. FoProBIM (FoProBIM 1985) found densities of D.
antillarum up to 14.5 ind 100 m-2 at 5 m depth in the area of Les Arcadines. More
recently, abundance values from 0 to 1.5 ind 10 m-2 have been reported in the
same area by Linton (2003).
However, ecological studies at the level of echinoderm populations or com-
munities seem to be scarce. In the Dominican Republic, Chiappone (2001) con-
ducted a study measuring density and size distribution for five species of echinoids
(D. antillarum, Echinometra viridis, Echinometra lucunter, Eucidaris tribuloides
and Tripneustes ventricosus) in the National East Park and Boca Chica. Tewfik
et al. (2005) studied the impact of anthropogenic enrichment on the seagrass food
web at two locations in southern Dominican Republic (Pedernales and Barahona),
involving several species of echinoderms, categorized by them as three consumer
functional groups: (a) generalists (regular urchins: Lytechinus variegatus, T.
ventricosus, D. antillarium); (b) subsurface deposit feeders (red heart urchin
Meoma ventricosa and sand dollars) and (c) surface deposit feeders (cushion sea
star Oreaster reticulatus and sea cucumbers). Hay (1984) compared the activity of
grazing fish and sea urchins in the Bay of Caracol in Haiti, on an overexploited
coral reef including density data of Diadema antillarum and its rate of con-
sumption of Thalassia testudinum.
Commercial fishing of sea cucumber takes place in the Dominican Republic.

According to the Dominican Council of Fisheries and Aquaculture CODOPESCA
in 2002 there was an export of 57.6 MT, equivalent to 9 % of seafood exports. The
figures for years 2007, 2008 and 2009 indicate much smaller export volumes with
1.27, 2.37 and 1.37 MT, respectively. The trading company for sea cucumbers in
the Dominican Republic is NETCO C x A which operates from Juan de Bolaño
Beach in Montecristi with CODOPESCA permissions for exportation to the United
States. There are no studies that scientifically support this exploitation. In fact,
there is no information on the species being harvested, minimum sizes, close
seasons or any other descriptive parameters of an organized fishery. The survival
of these echinoderm populations is seriously threatened. According to Toral-
Granda (2008), between 2002 and 2003, China imported 2,607 kg of dry weight of
sea cucumbers from Dominican Republic and 10,680 kg of dry weight from Haiti.
In Haiti, the Caribbean SeaFood Company announces online the sale of dried sea
cucumbers of the species Stichopus badionotus.
13.6 Threats
The threats to echinoderms are not very different from those faced by other marine
groups of Hispaniola. The destruction of ecosystems (mainly mangroves, seagrass
beds and coral reefs) due to coastal development (human settlements, agriculture,
industrial and tourist facilities and/or fishing) and tourism activities (especially
diving) is the major threat that affects all groups. In addition to impacts of habitat
destruction, some groups of echinoderms are exploited for commercial purposes.
These include some species of sea cucumbers used for human consumption and
some species of starfish and sea urchins used for sale as aquarium species
(SERCM 2004). Other echinoderm species, such as the sand dollar Mellita quin-
quiesperforata and the starfish Oreaster reticulatus are collected for handicrafts
that are sold in tourist establishments. Some fishermen also use this asteroid
species as bait in net traps. No ecological studies or fisheries biology research have
been done to support a sustainable exploitation of these resources.
13.7 Recommendations
It is essential to train a group of national specialists from Haiti and the Dominican
Republic in the systematic of all echinoderm classes that can work closely together
to better understand the echinofauna of Hispaniola. The National Museum of
Natural History in Santo Domingo should be involved in the development and
review of the marine collections. New investigations in structural ecology and
population dynamics of echinoderms must be initiated for those species that may
be subject to some kind of extractive use. For exploited species, it is also necessary
to conduct fishery biology investigations for the establishment of closed seasons,
protected areas from fishing and legal minimum sizes. Fishing regulations and
fishery statistics are crucial for a sustainable exploitation of these species. Marine
research should continue increasing the knowledge of the echinoderms in new

areas of the Dominican and Haitian platforms, including exploration of deeper
zones.
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and Jamaica. Rosentiel school of marine and atmospheric science. University of Miami,
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food web. Ecology 86:2726–2736
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A global review of fisheries and trade. FAO Fisheries and Aquaculture Technical Paper. No.
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evt3.senckenberg.de/page/index.htm
Chapter 14
Review of Echinoderm Research in Puerto
Rico, with the Focus on Biological
and Ecological Aspects
Stacey M. Williams, Milena Benavides-Serrato, Jose E. García-Ar-

rarás, Edwin A. Hernández-Delgado and Ruber Rodríguez-Barreras
14.1 Introduction
The island of Puerto Rico is located in the northeastern part of the Caribbean Sea
(18°130 N–66°280 W). Puerto Rico is considered an archipelago formed by the main
island and other small islands mainly off its eastern and southern coastlines
(Fig. 14.1). The eastern shelf is very shallow (typically less than 30 m in depth)
and delimited in the north by a chain of small islands and emergent monoliths
that extend from Cabezas de San Juan to Culebra island (18°180 N–65°160 W),
27 km off the main island. The island of Vieques [or: Vieques Island] (18°070 N–
65°250 W) lies south of the eastern shelf. Numerous mangrove keys and an offshore
island (6 km), Caja de Muertos (1.54 km2), characterize the southern coast of
S. M. Williams (&)
Universidad de Costa Rica, San Pedro, San José, CR 11501-2060, USA
e-mail: stcmwilliams@gmail.com
M. Benavides-Serrato
Echinoderm Curator Museo de Historia Natural Marina de Colombia,
Biodiversity Marine Ecosystems (BEM), Instituto de Investigaciones Marinas
y Costeras-INVEMAR, A.A 1016, Santa Marta, Colombia
J. E. García-Arrarás E. A. Hernández-Delgado R. Rodríguez-Barreras
Department of Biology, University of Puerto Rico,
PO Box 23360 Río Piedras, San Juan, PR 00931-3360, USA
e-mail: jegarcia@hpcf.upr.edu
E. A. Hernández-Delgado
e-mail: coral_giac@yahoo.com
R. Rodríguez-Barreras
e-mail: ruber.rodriguez@hotmail.com

438 S. M. Williams et al.
Fig. 14.1 Map of Puerto Rico with outlying islands. Coral reefs are marked in red and insular
shelf is displayed in light grey. Boxed in area identifies La Parguera, Puerto Rico
Puerto Rico. The western shelf is broad and shallow, especially in the south-
western portion. Off the western edge of the shelf in the Mona Passage is Isla de
Mona (18°040 N–67°520 W), located 68 km west from the main island with an area
of 55 km2 (Frank et al. 1998). The Mona Passage, which is about 120 km wide,
separates Puerto Rico from Hispaniola. The northern coastline is characterized by
a very narrow shelf with high energy and sandy beaches. There are other smaller
islets surrounding Puerto Rico: Monito and Desecheo on the western side, and
Culebrita and Palomino on the eastern side. The coastline of the main island is
580 km. However, if we add the adjacent islands of Vieques and Culebra, the total
shoreline is around 700 km.
14.1.1 Coastal Morphology
Sediment discharge from rivers is greater along the northern and western coasts,
than the eastern and southern coasts. This is due to the higher annual precipitation
levels in the northern and western parts of the island. Abundant sand dunes
characterize the northern and northwestern coast. Parts of these are submerged
while others have fossilized and formed rocky outcrops and beaches (García-Sais
et al. 2005). The eastern coast has sandy deposits with scattered rock formations.
Submarine canyons (Morelock et al. 1977) and mangrove islets characterize the
southern coast. In Culebra, mangroves grow along the southern while sand beaches
dominate the northern side. However, Vieques has an opposite pattern. This is the
14 Review of Echinoderm Research in Puerto Rico 439
result of differences in wave action. The northern coast is exposed to more wave
action resulting from northeast trade winds and North Atlantic winter storms
(Ruiz-Ramos et al. 2011).
14.1.2 Shelf Morphology
Puerto Rico is crossed by two important geological troughs, the Muertos Trough in
the south with a depth of 5,000 m and the Puerto Rico Trench, the deepest
submarine trough ([8,400 m) in the Atlantic Ocean, located approximately 55 km
off the north coast. The ocean depth, shelf, and slope morphology varies greatly in
Puerto Rico (Morelock et al. 1977). The shelf off the northern and northwestern
coasts is narrow (3 km wide, Fig. 14.1) and is subjected to strong wave action
during the winter season (December–March). Along the southern coast, the shelf-
edge drops off at a depth of about 200 m with an almost vertical and abrupt steep
slope in some areas (García-Sais et al. 2005). Also along the southern coast is a flat
platform (*10 km wide) at a depth of about 30 m located between Salinas and
Ponce municipalities. The eastern coast is broad compared to the rest of the island
and has a wider shelf (Fig. 14.1). This area is partially protected by a cordillera
aligned between Culebra and Fajardo, the Cordillera Reefs System (CRS). The
southern coast is usually characterized by lower wave energy. However tropical
storms and southern trade winds can affect this area. In the west, the Mona Passage
has an average depth of 400 m, but some areas are more than 1,000 m deep. Puerto
Rico has 3,370 km2 of fringing coral reefs surrounding the main island and also
Culebra, Vieques, Mona, Monito, and Desecheo (García-Sais et al. 2005).
14.1.3 Oceanographic Conditions
Coastal currents around Puerto Rico are mainly driven by tides and wind. The tide
along the north and western coast is semidiurnal and diurnal along the south coast
(Morelock et al. 1977). Wind driven currents are dominated by the easterly trade
winds, which push surface waters towards the west. Near bottom waters tend to
reverse the flow during times of weak easterly winds.
The main objective of this chapter is to describe the species diversity and
distribution of echinoderms around Puerto Rico. We also want to present a
comprehensive historical account of echinoderm research carried out in Puerto
Rico. Lastly, recommendations are given for future ecological echinoderm
research and management strategies that seem fit for the conservation of echino-
derms in Puerto Rico.
14.2.1 Taxonomic and Inventories Studies
The study of echinoderms in the Caribbean was initiated at the end of nineteenth
and beginning of twentieth centuries, as part of the expeditions HMS Challenger,
RV Blake and US Fish Commission RV Albatross. During these expeditions many
marine organisms many marine organisms from shallow and deep waters were
collected in the Caribbean, including Puerto Rico, Jamaica, Cuba, Bermuda,
Tobago, Cayman Island, Virgin Islands, Curaçao, Aruba, Bonaire, Venezuela, and
the coast of French Guyana. The main contributions have been the works of Clark
(1898, 1901a, b, 1919, 1933, 1941, 1942), (1939), Engel (1939), Cherbonier
(1959) and Engel and Aberson (1960). Not only taxonomic characteristics, but also
the morphological and ecological aspects of echinoderms were studied. Clark and
Downey (1992) was an important contribution to the study of sea stars in the
Caribbean Sea, including the Lesser and Major Antilles with the citation of 23
species. The most recent work is by Hendler et al. (1995), who produced an
identification guide and life history of 150 echinoderm species from the shallow
waters (0–33 m deep) of the Florida Keys, Bahamas and the Caribbean.
The first major report of marine biodiversity of Puerto Rico was done by Clark
(1901a), which was based primarily on the extensive collections made by the U.S.
Fish Commission steamer Fish Hawk in January and February 1899 and by Mr.
George R Gray 1900 collection near San Juan. It listed 87 species: three crinoid
species, 11 asteroid species, 49 ophiuroid species, 13 echinoid species and 11
holothurian species. This survey produced the first systematic list of marine
invertebrates and fish for the island.
The collections of echinoderms made by the New York Academy of Sciences in
1914 were reexamined in a new report (Clark 1921). The revised report contained
around 550 specimens. Eleven new species were identified in Puerto Rico. The list
was completed and reviewed three decades later by the same author (Clark 1933).
Several other studies on echinoderms have been done in Puerto Rico. For
example, Carrera (1974) studied the Class Ophiuroidea (particularly the Amphi-
uridae family) at La Parguera and found 17 new records for the island. Wolf
(1978) studied aspects of the biology of Astrophyton muricatum (Lamarck) at
Enrique, La Parguera; González-Liboy (1971) emphasized the echinoderms in his
study of the macro-invertebrates at the Maní beach, Mayagüez et al. (1976)
focused on the role played by the sea urchin Diadema antillarum (Philippi) on the
composition of corals and sea grass beds at Jobos Bay.
Recently, Benavides-Serrato (2006), based on a review of the literature,
museums revision and collections, made the most complete and current taxonomic
list of the shallow water (depth from 0 to 100 m) echinoderms from Puerto Rico.
One hundred and eight species were observed. Nine species, Davidaster discoidea
(Carpenter), Poraniella echinulata (Perrier), Ophioderma squamosissimum
(Lütken), Ophioderma phoenium (Clark), Ophionephthys limicola (Lütken), Pla-

giobrissus grandis (Gmelin), Actinopyga agassizi (Selenka), Holothuria (Ha-
lodeima) floridana (Pourtalés), and Synaptula hydriformis (Lesueur) were
considered as first records for Puerto Rico.
14.2.2 Marine Science Collection of Department of Marine

Sciences (UPR Mayagüez)
The echinoderm collection of marine invertebrates of the Marine Science

Department (MSC, Mayagüez Campus, University of Puerto Rico) is located at
Isla Magueyes (La Parguera), and is considered the most complete and important
one for Puerto Rico. The invertebrates collection has approximately 4,000 speci-
mens (Weil 2005) belonging to different taxonomic groups, including the Phyla
Porifera, Cnidaria, Mollusca, Nemertea, Echinodermata, Arthropoda and Annelida
(Class Polychaeta). Echinoderm collection is composed of approximately 74
species, belonging to all five classes.
14.2.3 Reproduction, Larval Biology, Settlement, and Recruitment
Free-spawning is a mechanism that organisms utilize to disperse their genes and

increase their population success. Egg fertilization of a free-spawning individual is
difficult because of the rapid diffusion and limited life span of gametes in a dynamic
ocean (Levitan 1995). The fertilization of the egg is critical for population survival
and there are many mechanisms that organisms use to increase fertilization success.
Some of these tactics are simultaneous spawning, aggregation, and large population
size (Levitan 1995). One way to measure fecundity is to measure the gonad index,
which is calculated as gonad weight divided by total weight times 100 (Giese and
Pearse 1974). The drop in gonad index can be interpreted as spawning events
(Moore et al. 1963). Measurements of fecundity can provide a first estimate of adult
contributions to larval populations (Cameron 1986).
Echinoids have a complex life-history, which involves external fertilization of
pelagic eggs, planktonic larval development and recruitment into a benthic habitat
where it reaches an adult stage and reproduces (Smith 1997). The distribution of
larvae is influenced by the dynamics of water motion in the coastal ocean in
relation to the location of the spawning sites (Williams and García Sais 2010).
Motility depends both on internal, physiological processes and on the physical
properties of the fluid environment. The physical properties of water (e.g. density
and dynamics viscosity) have a predominant influence on an organism’s motion
and are considered to be a major force in the evolution of their various modes of
locomotion and feeding (Podolsky and Emlet 1993). Questions about larval
transport focus largely on: (1) where they go and (2) where they came from (Levin
2006). Assessments of temporal and spatial abundance patterns of pelagic larval
stages are relevant for understanding dispersal strategies, spatial connectivity and
recruitment dynamics (Williams and García Sais 2010). Larval abundance mea-
surements is a challenging undertaking for echinoderms due to their small size,
relatively long pelagic larval life, potential for patchy distributions and the
dynamic environment in which they live in (Williams et al. 2009). Larval
community characterizations are typically incomplete and may underestimate
abundance, especially when using large mesh diameters such as C202 lm for
plankton tows (Williams and García Sais 2010). There are several studies of
coastal zooplankton communities of Caribbean systems (Reeve 1970; Youngbluth
1980; McWilliam et al. 1981; Yoshioka et al. 1985) however, these studies do not
report on echinoderm larvae from plankton sampling.
The colonization of benthic habitats by marine organisms with planktonic life-
stages involves three phases: planktonic development and dispersal, habitat
selection and settlement (Keough and Downes 1982). Settlement rate is defined as
the rate at which planktonic larvae of benthic organisms establish permanent
contact with the substrate. Trying to identify sources and/or sinks of larvae is
important to understanding supply to populations (Miller and Emlet 1997). Marine
invertebrate larvae may navigate to suitable settlement sites by responding to
gradients of environmental stimuli (Kingsford et al. 2002). Recruitment, which is
the fourth phase, includes the survival of the organism until observable by an
observer (Keough and Downes 1982). Recruitment is a key mechanism shaping
and structuring reefal communities (Cameron and Schroeter 1980). Recruitment
rather than reproductive output is the key concept for marine species demography
and is one of the main determinants of spatial and temporal variations in popu-
lation sizes (Cowen et al. 2000). Larval availability, substrate selection and
juvenile mortality are factors that influence recruitment to a population. There
have been only a few larval and recruitment studies of echinoderms in Puerto Rico.
Most of these studies focused on Echinoidea.
Studies on the Reproduction, Larval Biology, Settlement, and Recruitment

of Lytechinus variegates (Lamarck), Echinometra viridis (Agassiz),
Echinometra lucunter (Linnaeus), and Tripneustes ventricosus (Lamarck)
Cameron (1983, 1986) measured the reproduction, larval abundance, and size
distribution for Lytechinus variegatus, Echinometra viridis, E. lucunter and
Tripneustes ventricosus in La Parguera, Puerto Rico (Fig. 14.2). The mean gonad
index of L. variegatus ranged from 0.8 in October 1981 to 7.8 in October 1983.
The higher gonad indices occurred during the late summer and early fall months
for L. variegatus and Echinometra spp. This is when the sea surface temperatures
were at their highest. Spawning occurred eight times ranging over all seasons of
the year for L. variegatus. The gonad indices for Echinometra spp. reached a peak
in August 1983 (10.0). The degree of synchrony was much greater for
Fig. 14.2 Photographs of a Lytechinus variegatus. b Echinometra viridis. c Echinometra

lucunter. d Tripneustes ventricosus (Photos by Carlos Muñoz)
Echinometra spp. than for L. variegatus. The reproductive period of the white-
spined urchin, T. ventricosus in not very synchronous like the other three species
and also contrasts by a peak in the coolest time of the year. Tripneustes ventricosus
displayed gonad indices between 1.2 (July 1982 and June 1983) and 4.4 (May
1982) (Cameron 1986).
All four of these species displayed similar reproductive strategies; they all
broadcast gametes which develop into feeding pelagic larvae. Cameron (1986)
found that at 23–25 °C L. variegatus, E. lucunter and E. viridis achieve meta-
morphosis in about two weeks and T. ventricosus in about four weeks (Lewis
1958). The seasonal larval occurrence of L. variegatus and Echinometra spp.
coincides with their differences in synchrony. Lytechinus variegatus plutei were
present in all samples from the spring to fall 1982 and 1983. Plutei of Echinometra
spp. was only present during the fall reproductive peak. Larval abundances of
L. variegatus, E. viridis, and E. lucunter co-varied with the reproductive period.
Tripneustes ventricosus plutei were only found during August and September
1982. The larval period of T. ventricosus is twice as long as the other three species.
Summer currents may carry larvae of L. variegatus offshore followed by a later
return of mature larvae. Larvae of the other three species may be present in the
water column at a time when retention is favorable.
Cameron (1986) found L. variegatus juveniles (\15 mm in test diameter) at

Tres Marias in all four 1982 samples, April 1983 and June 1984. At Caracoles
juveniles were present during April and August 1982, April 1983 and June 1984.
Settlement of L. variegatus occurred throughout the year. The recruitment pattern
of this species follows the expected and observed pattern of reproduction and
larval occurrence. Juveniles (20 mm in diameter) of T. ventricosus found in
January or February may have settled the previous September. They most likely
settle on the reef face and crest. The larvae of T. ventricosus were observed during
August 1982, a time that coincides with the expected fall recruitment. Seasonal
recruitment was not apparent for E. lucunter at Papayo. There were E. viridis
recruits several times during the year.
Cameron et al. (1985) measured the embryonic response to temperature and
salinity by four species of Caribbean sea urchins. They wanted to test if embryos
of highly synchronous species of echinoderms are more sensitive to physical
factors. They cultured embryos of four species of sea urchins, L. variegatus,
E. lucunter, E. viridis, and T. ventricosus in a 3 9 3 factorial experiment com-
paring temperatures of 20, 25, and 30 °C and salinities of 31.5, 35, and 38.5 % at
La Parguera. The responses to different temperatures and salinities differed among
the embryos of the four species. The asynchronous species, L. variegatus and
T. ventricosus exhibit a more gradual response to temperature at cleavage than the
synchronous Echinometra spp. Lytechinus variegatus developed more rapidly at
higher salinities and T. ventricosus at lower salinities. Echinometra spp. are less
affected by salinity changes and have a linear response to an increase in temper-
ature. These results support the suggestion that species with a narrower repro-
ductive season have greater responses in development rate of extreme
environmental conditions. Salinity proved to be less harmful at all development
stages than temperature. Significant mortality occurred during the blastula and
early gastrula stages. The development rate of all four species is temperature
dependent. At 20 °C the hatching interval was the shortest for L. variegatus and
longest for T. ventricosus; Echinometra spp. had an intermediate response.
At 25 °C all four species hatched at about the same time, 7.5 h. Tripneustes
ventricosus had 100 % mortality at extreme temperatures. All four species reached
the pluteus stage (18–24 h) at optimum temperature. Development of L. variegatus
was fastest (18.3 h), while L. variegatus was slowest (28.6 h). Most synchronous
species (Echinometra spp.) were the most responsive to temperature changes.
14.2.4 Reproduction, Larval Biology, and Settlement

of Diadema antillarum
Lugo (2004) described the population structure and reproductive biology of

D. antillarum in La Parguera. The average gonad index per month was used to
characterize the annual reproductive cycle. The gonad index ranged from 2.9
Fig. 14.3 Early (4–8 day-

old) and late (*24 day-old)
larval stages of Diadema
antillarum
(June–July 2001) to 11.2 (December 2001). High gonad indices were observed
during the late fall (9.5) to winter during times when sea surface temperatures were
at its lowest values (26 °C). Lugo (2004) estimated that D. antillarum spawn
during early spring and early summer. He also found a significant negative rela-
tionship between monthly mean gonad indices and sea surface temperature. The
smallest sexually mature urchin was 3.8 cm in diameter. The male:female ratio
was 0.89:1 during his study.
Williams et al. (2009) measured the temporal variations of early larval stages in
neritic waters of La Parguera. They also measured the water current velocities and
directions of inshore areas of La Parguera. During April 2005, six drogues were
deployed. They also deployed an Acoustic Doppler Current Profiler (ADCP)
during April 2005. Only early larval stages (4–8 day-old, Fig. 14.3) of D. antil-
larum were collected in the study. However, Williams and García Sais (2010)
observed older larval stages (*24 day old) (Fig. 14.3) of D. antillarum at the
shelf-edge and oceanic stations.
Early larval stages were collected in 13 of the 16 monthly samples, which
indicates that D. antillarum spawn throughout the year. Mean larval abundances
(ind 100 m-3 ± SD) ranged from no individuals in September and October of
2005 and January 2006 to 1,239 ± 309 in April and 1,078 ± 648 in July. There
were peaks in larval abundances in April and July 2005. Echinoplutei were patchy
within sample replicates. The inconsistent peaks of abundance between years and
months may be a result of patchy distribution of these larvae. Williams et al.
(2009) did not find a significant correlation between monthly larval abundance and
SST. Drogue trajectories and the ADCP time series indicated that the circulation of
coastal waters at La Parguera is towards the west-northwest. The flow in La
Parguera seems to be influenced by the boundary layers of the coastline and coral
reefs. Early dispersal of D. antillarum larvae during the first two days is most
likely within the insular shelf of La Parguera. It is uncertain if the older larvae
continue offshore towards the Mona Passage or follow the shelf contour towards
the north along the westcoast of Puerto Rico. The recovering populations of
D. antillarum in La Parguera may be an important source of larval replenishment
for downstream islands.
Fig. 14.4 Aerial photograph of La Parguera, Puerto Rico. Major reef systems and Isla Magueyes
Marine Station have been identified
Larval settlement of D. antillarum was assessed in three separate surveys

(Miller et al. 2009; Williams et al. 2010, 2011) in La Parguera, Puerto Rico. Miller
et al. (2009) assessed the rates of D. antillarum larval settlement at two sites
(Enrique and The Buoy) in La Parguera (Fig. 14.4). Settlement was quantified
from August 2005 to July 2006 with experimental settlement plates modeled on
Bak’s (1985) study. During each month plates were examined in the field and all
D. antillarum settlers were recorded. Williams et al. (2010, 2011) determined the
settlement pattern of D. antillarum in La Parguera during two separate studies. In
2010, D. antillarum settlement was examined at three reefs: an inshore reef
(La Pelotas), a mid-shelf reef (Media Luna), and a shelf-edge reef (Old Buoy,
‘‘The Buoy’’) for one year. The study of Williams et al. (2011) included six reefs
in La Parguera: Pelotas and Enrique (inner shelf reefs), San Cristobal and Media
Luna (mid-shelf reefs), El Hoyo, and Old Buoy (shelf-edge reefs). Settlement was
measured for 6 months (Fig. 14.4). Artificial turf was used as the recruitment
plates for both studies (Williams et al. 2010, 2011). Plates were inspected for
D. antillarum recruits in the laboratory.
Mean test size of settlers (mean ± SD) collected in the studies of Williams
et al. (2010, 2011) was 0.90 ± 0.50 mm (Fig. 14.5). The maximum monthly
settlement of D. antillarum in all studies was *16 ind m-2 (Miller et al. 2009),
1,064 ind m-2 (Williams et al. 2010), and 265 ind m-2 (Williams et al. 2011). The
temporal distribution of settlement by D. antillarum in the 2011 study was similar
to previous observations made by Williams et al. (2010). Diadema antillarum
Fig. 14.5 Diadema

antillarum settler
settlement mainly occurred during the summer (July–September). In 2010, settlers

of D. antillarum were collected during 12 of the 13 sampled months, confirming
year-round reproduction in La Parguera (Lugo 2004; Williams et al. 2009, 2010).
Spatial variations of D. antillarum settlement were evident. There were signifi-
cantly more recruits collected at the shelf-edge than at the inshore reefs (Miller
et al. 2009; Williams et al. 2010, 2011). Williams et al. (2010) suggested three
non-mutually exclusive explanations for the low settlement at the inshore reefs: (1)
stochastic variation, (2) a shelf-edge reef larval ‘filtering effect’ (Gaines et al.
1985), and (3) high abundance of fleshy macroalgae colonizing the artificial turf
plates precluded settlement at inshore reefs. However, it was proven that fouling of
the substrate was not a major factor precluding the settlement of D. antillarum at
the inshore sites in La Parguera (Williams et al. 2011). Settler mortality and/or low
larval supply were two other possible explanations for the low settlement at
inshore sites in La Parguera. Settlement in all studies was concentrated at the shelf-
edge where adult D. antillarum are rare. Therefore, D. antillarum recruitment in
La Parguera may be independent of local adult density (Lessios 1988). Post-
settlement mortality on D. antillarum at the shelf-edge may be high, which
explains the absence of adults in this area.
14.2.5 Bioerosion
Sea urchins are considered a major bioeroder on coral reef communities. Griffin
et al. (2003) studied the bioerosion of Echinometra viridis on coral reef commu-
nities in La Parguera, Puerto Rico. The mean size and densities of E. viridis were
determined at two patch reefs: Palmas and Mario. The daily rate of bioerosion
Fig. 14.6 Holothuria

glaberrima specimen
(amount of CaCO3 removed) was estimated and measured in fecal pellets pro-
duced over 24 h by urchins from depths of 1–3 m. The size ranges of the urchins
varied between 2 and 2.5 cm in diameter. They repeated four different treatments
11 times during March and April 1998 with different sea urchins. Substrate (pieces
of dead coral) was the same for each treatment. In the different treatments they
estimated (1) endolithic and external CaCO3 sediment production by urchins, (2)
endolithic bioerosion only, (3) the amount of particulate CaCO3 the sea urchins
had in their gut, and (4) control for external sedimentation. Fecal pellets were
rinsed with fresh water, dried for 24 h at 70 °C, and then burned in a furnace at
555 °C to eliminate organic matter, and finally at 1,000 °C to determine calcium
carbonate (CaCO3). They found that bioerosion rates of E. viridis were higher in
the shallow areas (1–3 m), areas of high densities. Average densities of E. viridis
ranged from 0.77 to 62 ind m-2 and size ranged from 2.01 to 2.44 cm at both reefs.
They found the average daily individual bioerosion rate to be 0.18 g day-1 and the
mean annual bioerosion rates between 1.14 to 4.14 kg m-2 year-1. Bioerosion
rates measured in this study were lower than that reported for many other species
of sea urchins, for example D. antillarum, endolithic sponges and parrotfishes.
However, they found that bioerosion activity of E. viridis had an important effect
on the morphology on the reefs and that the high densities of E. viridis at Palmas
might account for the degradation of coral at this site.
14.2.6 Physiology
Holothuria (Selenkothuria) glaberrima (Selenka) as a Novel Model System

for Cellular/Molecular Studies
The laboratory of Dr. José E. García-Arrarás at the University of Puerto Rico has
made use of a local echinoderm species, the sea cucumber Holothuria glaberrima,
as a novel model system to probe the phenomenon of tissue and organ regeneration.
Holothuria glaberrima (Fig. 14.6) is a species common in costal waters that can be
Fig. 14.7 Holothuria glaberrima specimen a before and b after evisceration
found in rocky shores with high wave action and is usually in close associations
with the sea urchin Arbacia punctulata (Lamarck). Although the original goal of
the laboratory focused on the regeneration of the digestive tract, spin-off research
from the laboratory has produced important information on holothurian anatomy,
neurobiology, immunology, evolution and genomics among other research areas.
Here we will summarize some of the findings that have resulted from this research.
In summary, the studies in H. glaberrima have provided information of interest
to many research areas other than the field of regenerative biology. Ongoing
studies of the transcriptome of nervous and intestinal systems are expected to
increase the interest of other colleagues in related fields to expand the number of
investigations using not only this species but also other holothurians.
Evisceration
Holothurians are well known to eviscerate their internal organs when submitted to
different types of stress. Evisceration in the laboratory is induced by injections of
KCl, although it can occur spontaneously in foul water or low oxygen conditions
(Fig. 14.7). Holothuria glaberrima provides a classic example of this phenomenon
by eliminating most of the visceral organs through the cloaca (García-Arrarás et al.
1998). The eviscerated organs include the intestinal system, which breaks at the
esophageal-small intestine junction and at the large intestine-cloaca junction.
Other organs associated with the intestine, including the hemal system and the
right respiratory tree, are also eliminated. Similarly, most of the gonads (if the
animal is in the reproductive season) are ejected but the gonadal base remains.
The same process has been documented in another local species, H. (Halodeima)
mexicana (Ludwig) (unpublished results).
Anatomy
Research into the regeneration process has provided anatomical descriptions of

several structures, tissues and organs. The most extensively studied has been the
digestive tract, which includes a short esophagus, a long descending and ascending
Fig. 14.8 a Diagram of a

transverse section of one of
the five radial nerves found
within the sea cucumber body
wall.
b Immunohistochemistry of a
radial nerve using one of the
nervous tissue markers
developed in the laboratory
(see Díaz-Balzac et al. 2007)
small intestine and a descending large intestine that connects to the cloaca (Díaz-
Miranda et al. 1995, García-Arrarás et al. 1998). In addition, many of the inves-
tigations provide important information on the neuroanatomy of sea cucumbers,
including a description of the nervous system in the digestive tract and tube feet
and the discovery of new nerve plexi within the connective tissue, (García-Arrarás
et al. 2001; Díaz-Balzac et al. 2007, 2010a, b) thus making H. glaberrima a much
needed model for understanding neural connections in holothurians.
Neurobiology
Studies on H. glaberrima have provided important information on the nervous

system. The main focus of these studies has been the development and use of
neural markers that serve to identify and characterize components of the nervous
system (Fig. 14.8). They have been extensively used in neuroanatomical studies as
described above. Many of the markers are antibodies that recognize epitopes in
cells of the nervous system. Some of these markers are against vertebrate and
invertebrate molecules such as acetylated a-tubulin (García-Arrarás and Viruet
1993), neuropeptides (García-Arrarás et al. 1991a, b; Díaz-Miranda et al. 1995,
1996) or yet unknown proteins (García-Arrarás et al. 2001; Díaz-Balzac et al.
2007; Mashanov et al. 2010b). The work on the nervous system also permitted the
characterization of the first holothurian neuropeptide (Díaz-Miranda et al.
Díaz-Miranda et al. 1992) and studies on its possible function in the intestinal
system (Díaz-Miranda and García-Arrarás 1995). More recently, markers for
holothurian glia have been developed and used to characterize this little studied
component of the echinoderm nervous system (Mashanov et al. 2009, 2010a).
Intestinal Regeneration
The formation of a new intestine in H. glaberrima following evisceration has been

the main subject of a series of publications from the García-Arrarás group,
including several reviews (García-Arrarás and Greenberg 2010; García-Arrarás

and Dolmatov 2010; Mashanov and García-Arrarás 2011). The initial description
of the process was published over a decade ago (García-Arrarás et al. 1998). Many
of the subsequent articles describe cellular events that take place during the pro-
cess, including extracellular remodelling (Quiñones et al. 2002), cell migration
(Cabrera-Serrano and García-Arrarás 2004; García-Arrarás et al. 2006), myogen-
esis (Murray and García-Arrarás 2004), cell dedifferentiation (Candelaria et al.
2006), cell division (García-Arrarás et al. 1998), apoptosis (Mashanov et al. 2010a,
b) and epithelial to mesenchymal transitions (García-Arrarás et al. 2011). Many of
the cellular events are now well understood and have been integrated in a working
model (see Mashanov and García-Arrarás 2011, García-Arrarás et al. 2011). In
brief, following evisceration cells of the mesentery mesothelium, particularly
myocytes, dedifferentiate and migrate as a cell layer toward the free end of the
mesentery, where the intestine was previously attached, prior to evisceration.
As cells migrate, some undergo proliferation while others ingress into the con-
nective tissue and provide cells for the formation of the intestinal rudiment. This
gives rise, initially, to a solid rod of tissue that extends from the oesophagus to the
cloaca. Luminal cells from the oesophagus and cloaca migrate into this tissue rod
forming the luminal layer and thus, the lumen of the intestine. Once the initial
rudiment is formed, cells from the mesothelium differentiate, giving rise to the
muscle cells (Murray and García-Arrarás 2004) and possibly to neurons. A com-
parison to other regenerative processes, such as wound healing of the body wall,
has shown that the same cellular events are common to both (San Miguel-Ruiz and
García-Arrarás 2007).
In recent years, experiments have aimed at understanding the molecular basis of
intestinal regeneration. Many of these experiments are summarized below. The
experimental data indicate a large number of genes are associated with intestinal
regeneration; over 70 % of the studied sequences showed some type of differential
expression between regenerating and normal intestines (Ortiz-Pineda et al. 2009).
Nervous System Regeneration
One of the main research interests of the García-Arrarás laboratory has been the
regeneration of the nervous system. The main reason is that this type of regen-
eration is highly limited in vertebrates and most other chordates. Nonetheless,
echinoderms with their amazing regenerative potential should provide excellent
models to study nervous system regeneration. Many of the investigations carried
out in the laboratory centered on this topic. Thus, neuronal markers were devel-
oped and the neuroanatomy of the holothurians was described. Even early
experiments on intestinal regeneration (García-Arrarás et al. 1998) were initiated
with an interest in the enteric nervous system, that is, the nervous system of the
digestive tract. Results have shown that H. glaberrima can regenerate its enteric
nervous system and that this regeneration occurs concomitantly with the regen-
eration of the intestine itself (García-Arrarás et al. 1999). Additional experiments
were done to demonstrate that the central nervous system of holothurians, the so-
called radial nerve cords, could also regenerate (San Miguel et al. 2009). Results
showed that following transection, nerve fibers could re-grow and bridge the injury
gap forming a new scar-free nerve cord in about a month time. A process of
neurogenesis was also documented that produced new neurons within the regen-
erated cord. Ongoing experiments are aimed at characterizing the genes associated
with this regeneration process.
Genomics
The laboratory began to study the molecular basis of intestinal regeneration in the
late 1990s. Initially the work was done on a gene-by-gene strategy, where the
selected genes were either candidate genes, such as Hox genes (Méndez et al.
2000) or genes identified by differential display techniques comparing regenerat-
ing and normal intestines (Roig-López et al. 2001). In this way, several genes of
interest were characterized in the holothurian and were shown to be overexpressed
during intestinal regeneration. Among these were serum amyloid A (Santiago et al.
2000) and ependymin (Suarez-Castillo et al. 2004).
With the advent of high throughput molecular analyses, the number of genes
associated with intestinal regeneration was multiplied. First, three cDNA libraries
were prepared from normal and regenerating intestines and over 5,000 EST
(Expressed Sequence Tags) were identified (Rojas-Cartagena et al. 2007). These
ESTs represent sequences of mRNAs (and therefore gene sequences) that were
expressed in the tissues at the time of dissection. This was the first attempt to
obtain large genomic information from a holothurian. These ESTs sequences were
then probed in a custom-made microarray in order to determine which genes were
differentially expressed during the regeneration process (Ortiz-Pineda et al. 2009).
As mentioned earlier, a large number of genes (*73 %) were found to be dif-
ferentially expressed at P \ 0.05 probability. At more stringent conditions
(P \ .001) the percentage was still highly significant (*40 %). These studies have
identified holothurian genes possibly associated with wound healing, cell prolif-
eration, differentiation, morphological plasticity, cell survival, stress response,
immune challenge, and neoplastic transformation. Among those that have been
described are cytoskeletal genes, such as actins, and developmental genes, such as
Wnt and Hox genes that show interesting expression profiles during regeneration.
More recently large-scale 454 pyrosequencing techniques have been applied to
study regenerative processes and have identified over 2,000,000 ESTs that cluster
into *80,000 contigs (Mashanov et al. in prep.). The tissue used for these analyses
was the radial nerve cord of normal and regenerating animals following the
transection of their cord. Thus, when the data are fully analyzed, the expectation is
to obtain a large percent of the transcriptome of normal and regeneration radial
nerve cords.
It is important to emphasize that the sea cucumber H. glaberrima has, thus,

become the holothurian with the largest number of genes described, and a refer-
ence for the genomic analysis of other members of its class.
Immune System
One of the research areas that has benefited indirectly from these regeneration
studies is the field of echinoderm immunology. The reason is that regeneration
follows the loss of tissues and this loss is usually associated with some traumatic
event causing exposure of the animal’s internal environment to the outside media.
Thus, the immune system is activated to deal with the possible invasion of
pathogens. For example, in the case of evisceration, the cloaca ruptures and as the
internal viscera are ejected, the coelomic cavity is exposed to the sea water.
The outcome of these events is that activation of the regenerative process could
be occurring along with activation of the immune system. Cells and genes that
might be associated with regeneration might in reality be involved in the immune
response or vice versa. Alternatively the possibility exists that the two events are
somehow coupled and that immune associated cells and molecules are also par-
ticipating in the regenerative process.
Lipopolysaccharide (LPS), a component of the bacterial cell wall, has been
used to study if genes associated with regenerative processes are also associated
with immune activation. LPS causes a coelomocyte-mediated immune response in
H. glaberrima, inducing an approximately threefold increase in coelomocyte
phagocytic activity (Santiago-Cardona et al. 2003). A large number of immune-
related genes were identified from the EST database (Ramírez-Gómez et al. 2008)
and many of these were shown to be associated with LPS-induced activation
(Ramírez-Gómez et al. 2009).
These studies have prompted a further in depth study into how different
pathogens might be able to activate different components of the immune system, in
particular different sub-populations of coelomocytes (Ramírez-Gómez et al. 2010).
In summary, H. glaberrima has also served to study the immune responses in
holothurians and has provided important information on fundamental aspects of
the cellular and molecular immune response (Ramírez-Gómez and García-Arrarás
2010).
Molecular Evolution
Echinoderms are a key group evolutionary group of animals. They are invertebrate
deuterostomes closely related to chordates. As such, they can provide key infor-
mation on the evolution of genes and their associated roles. Experiments with
H. glaberrima have already documented the evolution of particular genes. Some of
these data can be found associated with genomic studies (Rojas-Cartagena et al.
2007; Ramírez-Gómez et al. 2008; Ortiz-Pineda et al. 2009). Other genes have
Table 14.1 Number of species per classes in Puerto Rico in different kind of ecosystems
Class # Coral reefs Seagrass beds Mangroves Rocky shores Sandy beaches
species
Crinoidea 14 11 – – – –
Asteroidea 25 6 2 2 5 12
Ophiuroidea 49 30 20 13 5 10
Echinoidea 16 5 6 1 7 12
Holothuroidea 29 10 18 4 12 17
Total 133 62 46 20 29 51
been studied more in depth and their evolutionary associations have been well-
described. This is the case for Hox genes (Méndez et al. 2000, serum amyloid
A protein (Santiago et al. 2000) and ependymin (Suarez-Castillo et al. 2007). The
latter is an excellent example of how studies in holothurians have served to
establish evolutionary patterns. In this case, prior to our study, ependymins were
thought to be a vertebrate-specific family. Its characterization in H. glaberrima
showed it was present in various invertebrate groups.
14.3 Distribution and Diversity of Echinoderms
Based on Hendler et al. (1995), Benavides-Serrato (2006), Pawson et al. (2009)

and the database of the National Museum of Natural History (NMNH) in Puerto
Rico island, 133 species of echinoderms (Table 14.1) are distributed along dif-
ferent kinds of ecosystems through the island.
According to Weil (2005), Puerto Rico harbors a large diversity of habitats
along the coast and on the insular platform. Estimates from recent mapping of the
coral reef ecosystems and associated communities and habitats along Puerto Rico
by NOAA, indicated a total area of 5,010 km2. Of these, 757 km2 were covered by
coral reefs and colonized hard bottom habitats (15.1 % of the total area), 625 km2
by seagrass beds (12.8 %), 97 km2 by hard bottoms dominated by macroalgae
(1.9 %) and 73 km2 by mangrove forests (1.4 %).
According to (García-Arrarás et al. 1998), the insular shelf of La Parguera, on
the southwest coast of Puerto Rico, is characterized by an extensive development
of coral reefs, seagrass beds and mangrove forest. The dry, warm and relative
stable climate, low wave energy, high water transparency, relatively wide shelf
and oligotrophic offshore waters, are some of the factors that contribute to the
conditions of the marine ecosystem of La Parguera. Interactions among coral reef,
seagrass beds, and mangrove communities provide for a highly productive,
structurally complex, and biologically diverse ecosystem. Mean surface salinity is
35.2 %, but varies from 34 to 37 %. Mean temperature is 28.4 °C, varying from
25 °C to 31 °C.
Coral reefs in La Parguera are distributed in three parallel lines: (1) inshore, (2)
mid-shelf and (3) outer shelf (Almy and Carrión-Torres 1963, Acevedo and
Morelock 1988). The length and the wide range of the reef’s depths produce
physical, chemical and biological gradients providing a highly productive, struc-
turally complex, and biologically diverse ecosystem (Morelock et al. 1977,
Acevedo and Morelock 1988). The most common mangrove species is Rizophora
mangle (L.) Laguncularia racemosa (L. Gaertn. f.) is less common. Extensive sea
grass beds are well established in La Parguera. Thalassia testudinum (Banks ex
König) and Syringodium filiforme (Kützing) are the most common and widely
distributed species. Also, most extensive sea grass beds are found at a depth of two
meter around mangrove islands (García et al. 2003).
With the goal to characterize the echinoderms fauna in two major communities
in La Parguera, Benavides-Serrato (2006) collected and described echinoderms
associated with coral reefs and seagrass communities during several field trips
(February of 2005 to January of 2006) at eleven different localities (Fig. 14.4).
As a result of this study, 32 species were found at La Parguera. Thirty-one
species of echinoderms were identified from sampling stations from the seagrass
beds. The most abundant classes were Echinoidea (mean abundance varied
between 0.22 ind m-2 at Caballo Blanco to 5.89 ind m-2 at San Cristobal, rep-
resented by L. variegatus, E. lucunter and E. viridis) and Ophiuroidea (mean
abundance varied from 0.31 ind m-2 at Caballo Blanco to 3.99 ind m-2 at San
Cristobal, principally represented by Ophiocoma echinata (Lamarck), Ophiactis
savignyi (Müller and Troschel), Ophiothrix (Ophiothrix) angulata (Say) and
Ophiothrix (Ophiothrix) orstedii (Lütken).
Twenty-four species of echinoderms were identified from the coral reef com-
munity. Ophiuroidea had the highest number of species (14). The composition of
the different classes at different depths showed that ophiuroid and echinoid species
were widely distributed from depths of 0 to 20 m. Crinoids only appeared at
depths greater than the 10 m. On the other hand, asteroids were absent from the
shallow water habitats (0–5 m). The crinoids Davidaster discoidea and D. rubi-
ginosa (Pourtalès) were only found at The Buoy at a depths of 20–30 m.
14.3.1 Asteroidea Distribution
A total of seven species of sea stars within the Order Valvatida have been doc-
umented in Puerto Rico. These include Oreaster reticulatus (Linnaeus) in the
family Oreasteridae. This is probably one of the most significant and ubiquitous
sea stars of seagrass communities in Puerto Rico. It has been commonly docu-
mented across the entire shelf on seagrass bottoms and sandy patches from nearly
sea level to about 25 m depths, but is particularly abundant on shallow water
(depth \ 10 m) seagrass communities adjacent to Vieques, Culebra Island, La
Cordillera, and Fajardo, off northeastern Puerto Rico. Populations have locally
disappeared over the last three to four decades from many locations across the
main island, probably as a combined impact of long-term habitat decline as a result

of chronic water quality degradation. Also, this species was largely exploited in the
past by artisans for souvenirs.
Ophidiaster guildingii (Gray), in the family Ophidiasteridae, is very uncommon
and has been documented only from shallow water (depth of 2–5 m) fringing and
patch reefs off Cabo Rojo and Culebra Island. This is a very cryptic species,
usually hidden in reef crevices and overhangs. The common comet sea star,
Linckia guildingi (Gray), is also a rare species due to its cryptic behavior, but has
been documented from coral reefs and rocky reef habitats from Culebra Island,
Humacao, Fajardo, Luquillo, Vega Baja, and Cabo Rojo, at depths ranging from
about 5 to 20 m. Four species have been also documented in the family Astro-
pectinidae, including Astropecten articulatus (Gray), A. duplicatus (Koehler), and
A. antillensis (Lütken). These three species can be typically found buried beneath
the surface of sandy bottoms at variable depths, from nearly sea level to more than
20 m. They can be occasionally to commonly observed on sandy and muddy
bottoms off Humacao, Naguabo, Ceiba, Fajardo, La Cordillera, Vieques Island and
Culebra Island, off eastern Puerto Rico. But A. antillensis and A. articulatus have
also been observed off Patillas in the southeast, and off Luquillo, Río Grande,
Carolina, San Juan, and Vega Alta, along the northern coast. In addition, Astro-
pecten marginatus (Gray) was also found in high abundance in Luquillo Beach,
Puerto Rico and shares the habitat with Mellita lata (Clark) (Kenk 1944). The
geographic range of A. marginatus comprises Venezuela, Trinidad, Guiana, and
Brazil. They are located as far south as 27°300 N.
There are also two species within order Spinulosida, family Echinasteridae. The
conical spined sea star, Echinaster sentus (Say), has been sporadically observed
from variable shallow water bottoms (depth \ 5 m), including seagrass commu-
nities, fringing reefs, mixed subaquatic vegetation habitats (algae ? seagrass), and
rocky outcrops from Culebra Island, La Cordillera, and Fajardo. The thorny sea
star, Echinaster echinoporus (Lamarck) was observed only once from Culebra
Island by Hernández-Delgado on a shallow water (depth \ 1 m) transitional
rubble-seagrass-mangrove bottom.
14.3.2 Ophiuroidea Distribution
Six species belonging to four families in the order Ophiurida have been docu-
mented. These include Sigsbeia conifera (Koehler) (family Hemieuryalidae). The
rose lace coral brittle star is a highly cryptic, very small and very rare brittle star
species apparently only associated with rose lace coral, Stylaster roseus (Pallas).
Hernandez-Delgado observed only one individual within a reef crevice in Culebra
Island during a night dive in 2002 clinging to a S. roseus colony at a depth of 2 m.
The blunt-spined brittle star, Ophiocoma echinata (Lamarck) (family Ophiocom-
idae) is a common inhabitant of rubble areas in shallow, protected reef lagoons.
They typically hide during the day under rocks. The reticulated brittle star,
Ophionereis reticulata (Say) (family Ophionereididae) share nearly similar habi-

tats with O. echinata, though they prefer sandy bottoms and hide under reef
overhangs and under rock and coral slabs. This is one of the most common species
of brittle stars across the Puerto Rican shelf, at depths ranging from nearly sea
surface to well beyond 30 m. it prefers sandy bottoms at the edge of coral reefs,
under coral overhangs or rubble.
There are also three ubiquitous brittle stars from family Ophiodermatidae,
which include Ophioderma rubicundum (Lütken), O. apressum (Say), and
Ophiothrix suensonii (Lütken). The first species can be typically found hidden
during the day under coral heads and reef crevices but foraging at night across
open areas of reef bottoms. During coral mass spawning events they become very
active and rapidly climb coral heads, sea fans (Gorgonia spp.) and other octocorals
approximately two minutes before mass spawning begins. This behavior can be
used as an indication of the beginning of spawning and could probably result from
some form of chemical reception of signals from corals. In the case of
O. apressum, its behavior and typical habitat range is fairly similar to that of
O. rubicundum, while O. suensonii is typically hidden during the day within
sponges but can be commonly seen active at night, often climbing different
octocoral species.
There is also an important species of basket star, Astrophyton muricatum
(Lamarck) (order Phrynophiurida). It can be commonly found across the region,
particularly in coral reefs with abundant sea fan populations and at least moderate
water circulation. They often coil into a tight rounded shape attaching to gorgo-
nians or sponges, but open widely to filter plankton at night by orienting its body
and spreading its arms perpendicular to the prevailing current.
14.3.3 Echinoidea Distribution
The distribution and abundance of Echinometra lucunter and E. viridis were

examined on seven coral reefs near the southwest of Puerto Rico (McGehee 1992).
Urchin distribution was strongly correlated with patterns in water motion, depth,
and substrate size among the reefs. However, the two species of Echinometra
occupy different habitats. Echinometra lucunter was most abundant in the shallow
forereef zone while E. viridis was more common in habitats with less water
movement and smaller substrate particle sizes, such as deep forereef and nearshore
sites. McGehee (1992) observed that water movement may be the best variable to
predict abundance and distribution of the two urchins. As for depth, E. lucunter
was found in shallow water at depths of 2–3 m, and E. viridis was found at depths
greater than 3 m. Also, E. lucunter can be extremely abundant (e.g., 50–125 ind
m-2) on subtidal emergent eolianite platforms along the northern coast (Hernán-
dez-Delgado, unpub. data).
As of 1940, Mellita lata was ranked the most common echinoderm of Puerto
Rico (Kenk 1944). The distribution of this species occurred in the uppermost
sediments layers on sandy shallow beaches along the entire circumference of the
island. Mellita lata was extremely numerous along the north-northeast coast
(Luquillo, San Juan, Guayama) and northwest and west coast (Aguadilla, Añasco,
Mayagüez). The moves by rhythmical movements of the long spines of the body,
marginal spines, interambulacra spines 1, 2, 3, and 4, and posterior lunule spines.
Kenk (1944) recorded about 35 strokes of the spines per minute at 26.5 °C. Other
members of order Clypeasteroida and family Mellitidae include Mellita quin-
quiesperforata (Leske) and Leodia sexiesperforata. Both are also commonly found
partially buried under sediments on sandy beaches, but particularly on bottoms
with high abundance of organic material (e.g., Naguabo, Humacao, Yabucoa).
Diadema antillarum Distribution
Before the mass mortality D. antillarum densities were high at many locations
around Puerto Rico, San Juan 13.8 ind m-2 (Bauer 1980), La Parguera 12.7 ind
m-2 (Craft 1975), Guayanilla 3.0 ind m-2 (Vicente and Goenaga 1984), and Jobos
Bay 2.6 ind m-2 (Rivera and Vicente 1976). Lugo (2004) measured the density
and size structure of D. antillarum at five reefs in La Parguera: Turrumote, Mata la
Gata, Cayo Enrique, Laurel, and Media Luna. The median density for all locations
was 0.08 ind m-2. The highest mean urchin density (0.13 ind m-2) was at Mata La
Gata, followed by Laurel (0.30 ind m-2) and Media Luna (0.15 ind m-2). No adult
D. antillarum were found at Turromote. The highest mean densities of D. antil-
larum were in the shallow fore-reef zone (depth from 0 to 3 m). He found that the
average size (± SD) for all sites was 6.71 cm (± 0.25). The size varied with
depth: 6.95 cm in shallower areas, 6.48 at intermediate depths, and 3.37 cm at
deeper reefs.
Shortly after Lugo (2004), Weil et al. (2005) described the decrease in the
population of D. antillarum from 1984 until 2001 and the population status
(abundances, distribution and size structure) after 17 years after the mass mortality
in La Parguera. They also found that overall the D. antillarum populations were
unevenly distributed and low across all sites (0.83–1.55 ind m-2) compared to pre-
mortality densities. They also observed more urchins in shallow water areas
(depth \ 3 m). Urchins were significantly higher in shallow reef platforms
(0.89–1.98 ind m-2) (depths from 0 to 3 m), and the upper fore-reef (0.56–2.33 ind
m-2) habitats (depths from 3 to 7 m), compared to deeper (depths [ 7 m) habitats
(0.01–0.88 ind m-2). In contrast, El Atravesado reef had higher urchin densities at
the deeper (depths from 8 to 10 m) habitats, along with high abundance of juve-
niles. A number of factors such as, planktonic settlement, differential morality of
juveniles, higher spatial heterogeneity, and presence of refuges could influence
D. antillarum distribution. Populations in the sea grass mounds were dominated by
midsize (6 cm) to large individuals (9 cm). Within reefs, the average size was
medium to large. Small sized individuals were more common in shallow reef
waters than on the Thalassia mounds. They observed a high abundance of juve-
niles in protected, back reef and lagoonal reef habitats. Higher number of
aggregations and a higher number of urchins per aggregation were correlated with
low complexity (rugosity) habitats. This supports the idea that this behavior pro-
vides protection. They also concluded that recruitment is not limited to highly
complex, calcium carbonate habitats. Although average densities were well below
pre-mass-mortality densities in Puerto Rico, results of this study indicate that
D. antillarum seem to be making a slow come back in La Parguera.
More recently, Ruiz-Ramos et al. 2011) measured the population recovery of
D. antillarum at 26 locations around Puerto Rico during 2003–2004. Seven sites
were surveyed in the northern region, five in the western region, four in the
southern and eastern regions and six around Culebra Island. Adults were absent in
11 of the 26 localities measured. The highest densities of D. antillarum occurred in
Culebra and in the northern region. Arrecife El Banderote, Culebra had the highest
densities of D. antillarum (0.91 ind m-2) followed by Bahía Tamarindo and Bahía
Sardinas, with 0.70 and 0.51 ind m-2, respectively. They believe that the con-
tinued low densities could be due to a number of factors such as diminished larval
supply, recruitment failure, post-recruitment mortality, currents or lack of an
appropriate habitat due to chronic environmental degradation. Also, larger indi-
viduals were more common in Culebra and the northern region. As for water
transparency, the mean value around the island was 6.2 m. Water transparency was
greater at the northern sites and sea urchin density was positively correlated with
water transparency. The benthic communities of the northern region differed
significantly from the other locations. They found the highest mean percent cover
of coral was in Culebra (12.6 ± 8.4) and the lowest to the east (0.1 ± 0.3). Cover
of macroalgae was greater on reefs in more turbid areas. A positive relationship
was observed between D. antillarum density and percent coral cover. In conclu-
sion, they found that densities in their study were relatively higher than many other
wider Caribbean reef systems. However, they believe that population recovery of
D. antillarum seems remote.
There is also another important member of order Diadematoida, the magnificent
urchin, Astropyga magnifica (Clark). This is a very rare species in shallow water
areas. It can be often found at depths well below 30 m, but occasionally occurs in
shallow water (depth of 1 m) across seagrass communities. There are also two
species in the family Echinidae (order Temnopeuroida), Lytechinus variegatus,
and Tripneustes ventricosus. Both species are common on shallow seagrass hab-
itats and protected backreef lagoons. Lytechinus variegatus appears to be more
abundant on seagrass habitats of the northern coast (e.g., Condado Lagoon, Vega
Baja), while T. ventricosus is more ubiquitous. Both species tend to cover their test
with fragments of algae and seagrass blades. There is a single member of family
Cidaridae, the pencil urchin, Eucidaris tribuloides (Lamarck). This is a fairly
common reef urchin that often hides under sheltered zones It can often be observed
on rubble and seagrass areas. There are two members of the family Brissidae
(order Spatangoida), Meoma ventricosa ventricosa (Lamarck), and Clypeaster
rosaceus (Linnaeus). Meoma v. ventricosa can be commonly found buried under
sandy bottoms or sparse seagrass bottoms, while C. rosaceus often camouflages its
body under seagrass blades, algae and seagrass blade fragments.
14.3.4 Holothuroidea Distribution
There are only a few species of non-pelagic sea cucumbers that are known to
execute rapid movements. The species that are able to make rapid movements are
members of the order Apodida. Astichopus multifidus (Sluiter) is a member of the
order Aspidochirotida. Specimens of this group are characterized by dish-shaped
tentacles and respiratory trees. The genus Astichopus is easily recognized because
it is very large and soft and with tube feet uniformly covering both the dorsal and
ventral sides of the body. Astichopus was most abundant on the northwestern
coast, at Crashboat Landing, Puerto Rico. Specimens have been seen at depths of
20–40 m at different times near Isabela, Camuy and Arecibo, Puerto Rico. There
were a few individuals collected from La Parguera, Puerto Rico in areas of dense
turtle grass (Thalassia testudinum). Glynn (1965) stated that Astichopus possibly
breed in the late autumn or early winter in Puerto Rico. Astichopus is very active.
The most frequent means of progression when the animal was left undisturbed is a
slow crawl. Glynn (1965) examined the environmental effects on movements.
Sudden changes in sea water temperature (3–4 °C) resulted in walking, bounding
and rolling movements more effectively than other environmental changes
(salinity, oxygen availability, light regimes, etc.). Specimens were attracted to
natural and artificial light of low intensity (B5 foot-candles) and did not demon-
strate any clear tendencies to geotactic or thigmotactic behavior. Increased activity
of this sea cucumber does not release any toxic substance.
Euapta lappa (Müller) can be also commonly found on shallow (3–10 m) to
moderately deep (20–25 m) reefs. They are soft-bodied and are commonly found
at the base of reef patches. They usually hide in reef crevices during the day and
emerge at dusk, moving around adjacent sandy and rubble bottoms. Several other
species in the family Holothuridae (Order Aspirochirotida) can be found across
seagrass, sandy and rubble bottoms in northern and eastern Puerto Rico, including
Holothuria mexicana, Holothuria (Thymiosycia) thomasi (Pawson and Caycedo)
and Actinopygia agassizzi (Selenka). Also individuals of Isostichopus badionotus
(Selenka) (family Stichopodidae) have been documented from Culebra Island.
14.3.5 Echinoderm Larval Distribution
Williams and García Sais (2010) described the temporal and spatial distribution
patterns of echinoderm larvae in La Parguera, Puerto Rico. Temporal variability of
echinoderm larval abundances was examined from a series of monthly tows
between Cayo Collado and Cayo San Cristobal in La Parguera. In this study
11,921 echinoderm larvae were collected. Echinoidea larvae were the numerically
dominant taxa, representing 52.5 % of the total. Amongst the sea urchin assem-
blage, 20 % were D. antillarum larvae. Ophiuroidea larvae represented 45.6 % of
the total. The combined abundance of Holothuroidea and Asteroidea larvae was
Fig. 14.9 The relative

abundance (%) of
echinoderm classes to the
total larval collection
less than 2 % of the total (Fig. 14.9). Larval composition found in this study was
similar to the adult population structure of echinoderms found in la Parguera
reported in Benavides-Serrato (2006) study. Echinoidea and Ophiuroidea larvae
were collected in all 20 monthly samplings. Holothuroidea and Asteroidea larvae
were absent on approximately two-thirds of the samples. A possible explanation
for the lack of these larvae may be due to under-sampling associated with low
abundance of adults and short larval duration. No seasonal trend of early-stage
larvae was observed in this study. However, higher abundances of later-stage
larvae of Echinoidea and Ophiuroidea peaked during the summer and fall. Echi-
noidea, Ophiuroidea and Holothuroidea larval abundances all peaked at the shelf-
edge. Larval abundances of Asteroidea showed an increasing pattern of abundance
from inshore to offshore, with a peak at the oceanic station. The depth of the
substrate at the shelf-edge and the lack of benthic and planktonic predators may
account for the higher abundance of echinoderm larvae at this location. Also,
internal waves that break at the shelf-edge bring colder, more nutrient rich waters
to the surface, which would positively influence zooplankton abundance. They
collected more echinoderm larvae using the 64 lm mesh net than with the 202 lm
mesh net. Echinoderm larval abundance may be underestimated when 202 lm
and/or greater mesh net sizes are used. Larval patchiness also tends to increase in
collections when the 202 lm mesh net was used. The larval distribution in the
temporal and spatial study was found to be patchy/aggregated. The overall vari-
ations and inconsistent peaks may be a result of the patchiness of these larvae.
Echinoderms have not been cultured nor fished in Puerto Rico.

14.5 Threats
14.5.1 Mass Mortalities of Echinoderms
Mass mortalities have been reported as far back as 1965, when Glynn (1968)
described the mass mortalities of echinoids and other reef flat organisms concur-
rent with midday and low water exposures in La Parguera, Puerto Rico. All dead
urchins and some other echinoderms were counted and removed from the sampling
plot at Caracoles Reef in La Parguera over a two day period. Population sizes of
living organisms were determined from m2 quadrat counts. Tolerance of desic-
cation was also investigated by exposing 25 individuals of each of the five echi-
noid species (T. ventricosus, L. variegatus, E. lucunter, Brissus unicolor (Leske),
and D. antillarum on coral rubble in open sunlight (unprotected) over a period of
six hours. The same desiccation experiment was performed, but this time the sea
urchins were provided with some protection. In addition, Glynn (1968) investi-
gated the effects of solar radiation on the sea urchin survival. Fifteen individuals of
three species of sea urchins were exposed to sunlight in water baths just deep
enough to cover the animal. The most severe mortality of echinoids was during
April 14, 1965. Tripneustes ventricosus experienced the highest mortality
(86.4 %), L. variegatus 64 %, B. unicolor 25.9 % and E. lucunter 1.2 %. Lyte-
chinus variegatus and T. ventricosus were found dead in shallow water zones and
had not been exposed to drying. Before the mass mortality L. variegatus had
an aggregated mean density of 57.6 ind m-2, T. ventricosus 13.6 ind m-2.
Echinometra lucunter was very abundant, numbering [1,000 individuals.
Diadema antillarum and B. unicolor were scarce but showed a significant increase
in densities by spring 1966. There was also a noticeable recovery of T. ventricosus
(35.6 ind m-2) and L. variegatus (539.6 ind m-2) the following fall.
There was two other mass die offs on May 16 and June 28, 1965 reported by
Glynn (1968). On those episodes, there were two species of ophiuroids that were
affected seriously by extreme temperatures (39 °C), Ophioderma appressa (Say)
and Ophiocoma echinata. During the summer months (May–August) there were a
series of minus tides during midday when heat and desiccation approached
extreme levels. Glynn (1968) frequently observed temperatures exceeding 32 °C
and, during May and June 1965, extreme temperatures as high as 39 °C.
Other echinoderm species involved in these three mass mortalities were:
D. antillarum, Ophiomyxa flaccida (Say), Ophiothrix angulata, Ophiocoma riisei
(Lütken), Ophiactis savignyi, Holothuria (Halodeima) mexicana, and H. (Thymi-
osycia) arenicola (Semper). For the desiccation experiment, the loss in weight as a
function of exsiccation showed a linear relationship for both protected and
unprotected individuals. The loss in weight by B. unicolor increased to 70 % from
the initial wet weight after 6 h. Echinometra lucunter had the highest loss of
weight (50 % of initial wet weight after 6 h). All the unprotected urchins died after
3 h of exposure except E. lucunter. For the protected urchins, T. ventricosus had
the highest resistance to desiccation (80 % of the initial wet weight). For the solar
radiation experiment, five urchins died after 48 h of recovery, three in the

unprotected group and two in the protected group.
The second mass mortality event of echinoderms in Puerto Rico was between
1983 and 1984. Lessios (1995) stated that the 1983–1984 mass mortality of D.
antillarum was the most extensive and severe die-off ever recorded for a marine
invertebrate. Before 1983, the presence of this species was common (22 ind m-2)
on coral reefs (Lessios 1995). After the massive die-off, populations were drasti-
cally reduced by 95–100 % in many Caribbean locations (Lessios 1995). It was
theorized that a water-borne pathogen transported by surface currents was the main
cause of this mass mortality (Lessios et al. 1984, Phinney et al. 2001). In addition,
one report states that ballast water from ships was also linked with the spread of
the urchin-killing pathogen (Bak et al. 1984). The spread of the massive mortality
is believed to have originated from Punta Galeta, Panama and proceeded in two
different fronts. The first front went east from Panama to Tobago. The second front
was faster and went east of Costa Rica, North of Cayman Island, Belize, Mexico,
Jamaica, Gulf of Mexico, Florida, and to Bermuda. In February of 1984, an area of
3.5 million square kilometers was affected by this mass mortality (Lessios 1988).
Vicente and Goenaga 1984 observed the first diseased D. antillarum in January
1984 at Laurel reef, La Parguera, Puerto Rico. Before the mass mortality, densities
were reported from 2.6 ind m-2 (Jobos Bay) to as high as 13.8 ind m-2 (San Juan).
In Guayanilla densities before the mass mortality were 3.0 ind m-2 and after were
as low as 0.13 ind m-2 (Vincent et al. 1985).
Bunkley-Williams et al. (1986) reported the third recognized mass mortality of
echinoderms in Puerto Rico waters. This study described the isolated mortalities of
two sea urchins, Astropyga magnifica and Eucidaris tribuloides, during the winter
of 1984–1985 in the Northwest coast of Puerto Rico. An aggregation of 68
A. magnifica, at depths between 0.6 and 1.2 m deep with dropping spines and
rolling in the waves with the surge were observed at Crashboat Beach, Aguadilla
on December 8 and 12, 1984. These signs seemed very similar to those witnessed
during the D. antillarum 1983–84 die-off. Astropyga pulvinata (Lamarck) located
in deeper waters seemed healthy and there was an absence of sediment on tests. No
other species in the vicinity showed signs of disease or stress. Therefore, pollutants
were not thought to be a factor in this urchin mortality. In January 1985, there was
another mass mortality of E. tribuloides at Survival Beach near old Ramey Air
Force Base, Aguadilla. Six thousand to 8,000 sea urchins were observed washed
up on the beach. These die-offs were during times of rough waves and stormy
winter weather. No further mortalities were witnessed. These two mass mortalities
of A. magnifica and E. tribuloides are unexplained events.
Williams (1991) reported mass mortality outbreaks of T. ventricosus in Puerto
Rico between January and February 1995. The new die offs progressed more
slowly (6–8 days) compared to the D. antillarum die-off during 1983–84
(3–4 days). One thousand T. ventricosus were found dead at La Parguera
(Williams et al. 1996). This mortality was not localized but island wide. They also
witnessed urchins dying near Humacao and Fajardo. No samples were taken or
preserved. There was variation in symptoms. Many urchins were floating near the
surface while they still were able to direct spines toward tactile stimulation. The
gas production and lost of spines suggests that this species was involved in a
disturbance, primary or secondary pathogen. A similar die-off was reported
between September and November 1990 in La Parguera (Colon-Jones 1993). This
die-off was due to air exposure and/or higher seawater temperatures. The extent,
significance and frequency of these die-offs must be better understood in order to
identify the cause and their relationship with any other disturbance affecting their
habitat.
14.6 Recommendations
The knowledge of echinoderms in Puerto Rico has been scarce and focused
principally to ecological studies. Although inventories have been developed, this is
incomplete, especially in the north of the island. The focus of research has been
mainly located in La Parguera, Puerto Rico. The facilities located on the Isla of
Magayues (UPR at Mayagüez) are conveniently located for scientists to carry out
research in La Parguera region. Research needs to be extended and increased in
other zones of the Archipelago, especially the little islands: Mona, Culebra, Vi-
eques and Desecheo. In order to increase the research of these echinoderms, it is
necessary to encourage to students and scientists to study this phylum, especially
phylogeny, taxonomy, biogeography and functional aspects.
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Chapter 15
Echinoderms of the Canary Islands, Spain
José Carlos Hernández, Sabrina Clemente, Fernando Tuya, Angel

Pérez-Ruzafa, Carlos Sangil, Leopoldo Moro-Abad and Juan José
Bacallado-Aránega
J. C. Hernández (&) S. Clemente

Departamento de Biología Animal (Ciencias Marinas), Universidad
de La Laguna, Biodiversidad, Ecología Marina y Conservación,
c/Astrofísico Francisco Sánchez sn, La Laguna, 38206 , Tenerife, Canary Islands, Spain
e-mail: jocarher@ull.es
S. Clemente
e-mail: msclemen@ull.es
F. Tuya
Departamento de Biología, Universidad de Las Palmas de Gran Canaria,
Las Palmas de Gran Canaria, Campus Tafira s/n, Las Palmas de G.C.,
Canary Islands 35017, Spain
A. Pérez-Ruzafa
Departamento de Ecología e Hidrología, Universidad de Murcia, Murcia, Spain
C. Sangil
Departamento de Ecología e Hidrología, Universidad de La Laguna,
Biodiversidad, Ecología Marina y Conservación, c/Astrofísico Francisco Sánchez sn,
La Laguna, 38206 , Tenerife, Canary Islands, Spain
L. Moro-Abad
Servicio de Biodiversidad del Gobierno de Canarias, Santa Cruz de Tenerife,
Canary Islands, Spain
J. J. Bacallado-Aránega
Museo de Ciencias Naturales de Tenerife, c/Fuente Morales s/n, 38004,
Santa Cruz de Tenerife (Canarias), Canary Islands, Spain

472 J. C. Hernández et al.
15.1 Introduction
15.1.1 Geographical and Geological Context of the Canary Islands
The Canary Islands are located in the Northeast Atlantic, between 27°–29° N and
14°–18° W. The Archipelago consists of seven major islands, four smaller islets
and numerous rocky outcrops (Table 15.1, Fig. 15.1). The islands are arranged
longitudinally along 450 km. Fuerteventura Island is only 90 km from the African
continent, La Palma is almost 400 km from it (Fernández-Palacios and Martín
Esquivel 2001a). The islands have a land surface area of 7,490 km2 and 1,501 km
of coastline. The land surface and length of coastline vary in the different islands;
Tenerife is the largest in area and length of coastline (2,034 km2, 336 km), while
El Hierro has the smallest surface (269 km2) and La Gomera the shortest length of
coastline (97 km).
From a geological point of view, the Canary Islands belong to the group of
‘‘oceanic islands’’, they emerge from the ocean floor and have their own structures
(insular edifice) completely independent of the continents. In fact, the Canary
Islands, except for Lanzarote and Fuerteventura that share the same insular edifice,
had independent origins from the others. The size of the islands’edifice is espe-
cially relevant since over 80 % of most islands is under water. On average, insular
edifices rise from a depth of 3,500 m. The depth is greater than 4,000 m for La
Palma and El Hierro (Carracedo 1988; Carracedo et al. 2001).
The origin of the islands dates back to about 20 million years, but there are also
important variations in age between islands (Table 15.1). In fact, while in some
islands the eruptive periods have ceased several million years ago, in others it has
continued up to the present, as indicated by the large number of historical erup-
tions (\500–600 years old). A good example of this volcanic activity is the recent
eruption off El Hierro Island that started in October 2011. The difference in the
ages between islands has important implications for the substrate morphology and
topography, both inland and underwater, since erosion and remodeling of insular
edifices are higher in older islands (Carracedo et al. 2001).
The abyssal plain between the islands is dotted by a large number of submarine
mountains and volcanoes. Most rise only a few hundred meters above the ocean
floor, but there are two mountains, the ‘‘Banco de La Conception’’ located north of
Lanzarote and the ‘‘Banco de Amanay’’ southwest of Fuerteventura, whose summit
is less than 100 m deep. In the past, when sea level was lower, these mountains
were above sea level and were true islands (Fernández-Palacios 2004). The current
submarine volcanism is now considered more important than terrestrial activity,
since recent seismic studies and dating of lava flows indicate the presence of active
submarine volcanoes in the western region of the archipelago (Carracedo 2001).
Due to the volcanic nature of the islands, their edifices have a significant slope,
thus the island platforms (shallow bottoms of gentle slope located between 0 and
100 m depth) are limited in area compared to the land surface (Mitchell 1998;
Masson et al. 2002). The extension of the island platforms largely depends on the
15 Echinoderms of the Canary Islands, Spain 473
Table 15.1 Geographic data of the Canarian Arquipelago, incluiding islets (*). Source: Instituto
Geográfico Nacional, Ministerio de Fomento (http://www.ign.es), Carracedo et al. (2001)
Island Area (km2) Coastal perimeter (km) Age
(mill. years)
Lanzarote* 885 237 15,5
Fuerteventura* 1.664 310 20,5
Gran Canaria 1.560 252 14,5
Tenerife 2.034 336 7,5
La Gomera 370 97 12
La Palma 708 162 1,5
El Hierro 269 107 0,8
geological age of the islands and on the physicochemical characteristics of erupted

materials. The older islands (Fuerteventura, Lanzarote, Gran Canaria, La Gomera)
have been exposed longer to the action of erosive agents (waves, wind, runoff) and
therefore they have a wider platform than the younger islands (Tenerife, La Palma,
El Hierro) (Mitchell et al. 2003). In the case of El Hierro, the youngest island, the
edifice has hardly been remodeled and the insular platform is very small or non-
existent (Fig. 15.1b). Within each island, the extent of the platforms may differ.
They usually show greater development in the northern faces of the islands, which
are areas more highly exposed to erosive agents (Mitchell et al. 2003).
Along with erosion and the creation of offshore platforms, other underwater
structures and events have been recognized, such as rifts and massive landslides
(Urgeles et al. 1999; Masson et al. 2002; Mitchell et al. 2002). Rifts are the growth
axes of the islands that constitute their structural basis since they concentrate the
greatest amount of volcanic activity. Landslides are sudden and intense events that
generate large avalanches, caused by the gravitational effect due to the instability
of the insular edifice or by volcanic eruptions. Landslides detected in the archi-
pelago can extend into the ocean for over 100 km and constitute a volume of more
than 100 km3 (Watts and Masson 1995). These are not strange phenomena in
oceanic islands, or in the Canary Islands, and they represent typical stages of
consolidation and settlement processes of insular edifices (Gee et al. 2001a, 2001b;
Mitchell et al. 2002).
The morphology of the coastline and the continental shelf is highly variable
throughout the Canarian Archipelago and even in each island. However, several
shapes or substrates are more frequent than others, in relation to the geologic age
and its interaction with environmental factors (waves, currents, wind, etc.). The
coastline on the older islands has been heavily modified. It is homogeneous in
shape and hardly scalloped, representing the typical shape of a backward-stepping
coast. Beaches of organic origin are abundant, but cliffs and cobble beaches are
also frequent. Intertidal platforms, typically of small amplitude, show greater
development in the eastern islands. In contrast, in younger western islands where
the processes of coastal erosion coexist with volcanism, a more complex littoral,
richer in shapes, is found. The coastline is fringed with numerous points and rock
Fig. 15.1 Geographical position and bathimetric map of the Canarian Arquipelago. Island
platforms between 0–100 m depth are in white. Source Estudio Hidrográfico y Oceanográfico de la
Zona Económica Exclusiva de Canarias (ZEE-03), Ministerio de Defensa del Gobierno de España
ledges, as well as small shoals, but cliffs are the dominant forms. Beaches are
scarce, smaller, and mostly of volcanic origin (Yanes 1990).
In the submerged platforms sandy bottoms dominate, a pattern that increases
with depth because the sedimentation is favored by the reduction of hydrody-
namics with depth. On the other hand, the percentage of surface occupied by sandy
bottoms differs in each island. In older islands sedimentation seems to be favored
by the low slope. The heterogeneity (roughness) of the bottoms is particularly high
in younger islands, where the steeper slope of their bottoms along with the pres-
ence of submarine lava flows creates highly complex environments (Yanes 1990).
15.1.2 Oceanography
From an oceanographic point of view, the Canary Islands are a region of great
interest due to the interaction of different phenomena: the upwelling off the
African coast, the cold Canary Current, waves and trade winds. The combination
of these phenomena creates a complex mosaic of environments throughout the
archipelago, with peculiarities in the physical–chemical conditions of its waters.
These processes cool the waters of the archipelago that would otherwise have
higher temperatures in relation to the latitude (Brito et al. 1984; Aguilera et al.
1994; Fernández-Palacios and Martín-Esquivel 2001a; Fernández-Palacios et al.
2001b). The waters of the archipelago are situated on the eastern limit of the North
Atlantic subtropical gyre, occupying an intermediate position between the rich
cold upwelling waters of the northwest African coast and the warm and oligo-
trophic waters of the open ocean, giving rise to the so-called Transitional Coastal
Zone (Braun and Molina 1988; Barton et al. 1998; Barton and Arístegui 2004).
The surface seawater temperature ranges from minimum values of 17–19 °C in
April-March to maximum values of around 23–25 °C in September–October. This
surface temperature regime is slightly cooler than expected at the islands’ latitude
in the Atlantic Ocean. This is due to the cold Canary Current and the upwelling off
the African coast (Braun and Molina 1988; Molina et al. 1996; Hernández-Guerra
and Nykjaer 1997; Tejera et al. 2002; Marrero-Díaz et al. 2006).
The upwelling off the nearby African coast has its origin in the action of trade
winds from the northeast over the northwestern coasts of Africa. This causes, fol-
lowing Ekman’s theory (Ekman1905), the emergence of deep waters perpendicular
to the direction of the winds and towards the archipelago. Upwelled seawater is
colder, lower in salinity and richer in nutrients compared to surface water. The
incidence of this phenomenon over the archipelago has an important spatial and
temporal variation. The eastern islands, the closest to it up-welling, are more affected
by this process. Its influence on central and western islands is less and mainly due to
the incidence of the so-called upwelling filaments. Temporally, upwelling is more
regular in those months when the trade winds are more intense and constant (May–
August) (Braun and Molina 1988; Nykjaer and Van Camp 1994; Hernández-Guerra
and Nykjaer 1997; Tejera et al. 2002; Barton and Arístegui 2004).
The cold Canary Current, which flows NNE–SSW, is a branch of the Gulf
Current, a cyclonic system of currents that carries warm waters from the Caribbean
Sea to the northeast Atlantic Ocean. As it crosses the Bay of Biscay, this current
begins to cool, becoming a cold current when it reaches the Canaries. The average
speed of this current is 0.1–0.2 ms-1, but undergoes major seasonal changes
reaching minimum values of 0.025 ms-1 in summer. Each island acts as a barrier
during the flow of the current throughout the archipelago. This causes changes in
speed and creates mesoscale phenomena such as cyclonic and anticyclonic eddies
that produce sinking and upwelling events of small amounts of waters, and down-
stream warm wakes in the south and southwest regions of some islands (Braun and
Molina 1988; Arístegui et al. 1997; Tejera et al. 2002; Barton and Arístegui 2004).
Fig. 15.2 Sea surface temperatura (SST) map of the Canarian Arquipelago and surrounding
area. Source Dr. Antonio García Ramos, Departamento de Biología de la ULPGC
The different interactions of upwelling and, to a lesser extent, of the cold

current of the Canary Islands along the archipelago creates a gradient in temper-
ature (&2 °C), salinity, nutrients and primary production between the eastern and
western islands (Fig. 15.2). In this sense, the waters off the eastern islands have
more affinity to the waters of the temperate regions, while those off the western
islands are more similar to the waters of the tropics (Braun and Molina 1988;
Barton et al. 1998; Davenport et al. 2002; Barton and Arístegui 2004).
The open nature of the coasts of the Canary Islands results in a great proportion
of the littoral being exposed or semi-exposed to wave action. Bays and coves
protected from the waves are much more localized along the islands and are a
small part of the coast. The waves that affect the archipelago consist of two
substantially different types: wind waves and swells. Wind waves are dominant
over the swells. Wind waves, from the NNE and NE, have a mean speed of
18–22 kmh-1. They are produced by the action of trade winds over the seawater
surrounding the islands, creating waves that follow the direction and intensity of
the winds. The mean annual height of these waves is 1.4 m, with a frequency
of 9.5 s-1. Swells from NNW and NW are produced by storms from remote areas
of the North Atlantic. They create perturbations that travel along the ocean for
distances of up to 3,500 km. This kind of wave movement shows a marked
seasonal pattern, being more frequent from October to March. The mean annual
height of waves is between 2 and 3 m, with a frequency of 18 s. The prevailing
northern component in wave action at the Canary Islands makes the northern
coasts of the islands more exposed than the southern coasts (Braun and Molina
1988; Pérez et al. 2003; Yanes et al. 2006).
The climatic regime of the islands is characterized by the predominance of

trade winds from the northeast that affects the islands when they are under the
influence of the Azores anticyclone. These winds have an average speed of 20–22
km h-1. They are not constant throughout the year, undergoing major seasonal
variations. In summer they reach their maximum frequency, occurring around
90–95 % of the days. In fall the Azores anticyclone gets weaker, producing
periods of calm. However, it is in winter and spring when there are minimum
levels of trade winds, with an incidence of less than 50 % of the days. During this
period low pressures from the Atlantic Ocean are frequent, producing winds from
the northwest, southwest or southeast, as well as warm and dry invasions of
Saharan weather conditions towards the east and southeast (Marzol 1988).
The tidal regime of islands is semidiurnal as in the rest of the Atlantic region,
with two high tides and two low tides each day. Usually two consecutive high tides
and two low tides have the same amplitude, experiencing a change during the year.
The maximum tidal amplitude occurs during the spring and fall equinoxes,
reaching about 2.5 m, whereas minimum values are recorded during the summer
and winter solstices, not exceeding 0.7 m of amplitude. During the rest of the year
the average tidal amplitude usually ranges between 1.2 and 1.3 m, but local
variations may occur due to the morphology of the coast, the depth, or to particular
climatic conditions of winds and waves in favor or against tidal waves (Braun and
Molina 1988; Yanes 1990).
15.1.3 General Characteristics of Communities and Ecosystems
The seabed of the Canary Islands is morphologically highly complex with a rich
and diverse biota. The presence of certain biological populations and their struc-
ture are determined by the interplay of five factors: insularity, topography of the
bottoms, great length of coastline, oceanographic conditions and geographical
location. Since the Canary Islands originated from the seabed, isolated from other
regions, its biota necessarily came from nearby continents or islands. One could
expect to find in the islands the same patterns of communities and ecosystems
structures as in nearby regions. However, despite the great dispersal ability of
marine species, many species fail to cross ocean barriers. Therefore a biased
representation of the biota appears in the archipelago. The volcanic nature of the
islands determines the pronounced slope of the bottoms and the small extent of
sublittoral platforms that abruptly terminates onto the islands’ slopes. This implies
that the extension on which benthic communities can develop is spatially very
limited. However, the long coastline, along with the rich and diverse coastal
morphology, allows the occurrence and development of numerous different hab-
itats and ecosystems. On the other hand, the waters surrounding the islands are
oligotrophic that limits production in the environment (Braun and Molina 1988).
From a biogeographic point of view, the geographical location of the archi-
pelago has resulted in the coexistence of both temperate and tropical species (Brito
et al. 2001; Sansón et al. 2001; Brito and Ocaña 2004; Tuya and Haroun 2009). It
has been noted that the main system of currents that affects the archipelago is the
cold Canary Current, a descending branch of the Gulf Current that flows from the
north of the Iberian Peninsula towards the southeast. On its way through the
European Atlantic coast, it becomes progressively richer in propagule and larvae
of different species. Some of these species successfully reach the islands and become
established if conditions are favorable. Therefore, most of the species found in the
Canary Islands have their origin in the Atlantic-Mediterranean region. To a lesser
extent, American tropical and subtropical species have also colonized the Canary
Islands through this current. In many cases, the islands constitute the only region in
the eastern Atlantic Ocean where these species are found. Although there is no clear
system of currents that connects the islands with the tropical southern African coast,
some species from this region have colonized the Canaries. Probably southeast
African weather conditions occasionally counteract the Canary Current, favoring the
arrival of these species. Biota common to the Saharan coast (the region of mainland
Africa nearest to the Canary Islands) are scarce, mainly because the oceanographic
conditions generated by the upwelling of cold waters near the continent area are
very different from the conditions of the Canaries (Aguilera et al. 1994).
This set of variables defines the characteristics displayed by coastal popula-
tions, communities and ecosystems. Because the structure, composition and
functioning of ecosystems show characters in between those of temperate and
tropical systems, they are considered subtropical. The richness, diversity, disparity
and originality of the populations are very high and trophic relationships among
species are complex in many cases. The size of populations is usually small, due to
both the low productivity of the waters and the spatial restrictions of habitats. In
this sense, it should be emphasized that all benthic ecosystems, with the exception
of abyssal plain ecosystems, lack of continuity throughout the archipelago because
they are confined to the insular edifices and seamounts. The surface occupied by
benthic ecosystems depends directly on features inherent to the islands’ edifices
(e.g. coastal ecosystems between 0 and 100 m depth are larger on some islands
than in others). In this sense, according to the different availability of habitats and
to the oceanographic conditions at the islands, the population size of a species can
be different in each of them. On the other hand, given that most species are linked
to the bottoms, the non-continuity of ecosystems means that the only way of
exchange between populations is by larval dispersal. Since few species (only some
species of fish and probably also some cephalopods) have adults that are able to
move between islands, genetic connectivity between the islands’ populations in
turn depends on factors such as the distance between islands, the speed and
direction of currents, and characteristics of each species (e.g. dispersal ability of
the larvae). The population size of each island is also directly related to different
levels of human exploitation. The east to west oceanographic gradient across the
archipelago is also responsible for the distribution of biota. In the western islands,
with warmer waters, species with tropical affinities are more abundant, while
species with temperate affinities are more abundant in the eastern islands of
temperate waters (Brito et al. 2001; Sangil et al. 2007). Finally, the proximity of
the oceanic waters to the coast, and given that the Canary Islands are among one of
the main routes of migratory species (e.g. tunids), the pelagic systems strongly
interact with the benthic-littoral systems. Moreover, since great depths are occur
near the coast, deep-water species that migrate to shallow areas during their diel
rhythms and are temporarily integrated into coastal ecosystems (Aguilera et al.
1994). In this sense, it is also important to emphasize that some deep fauna occur
in shallow caves along the coast (Sangil 2007).
15.2 Research
The following section provides a historical perspective on the development of

studies on echinoderms in the Canarian Archipelago. We describe the shift in the
scientific strategy for the study of this zoological group and as some data to
quantity published material.
Although the echinoderm fauna of the Atlantic is well known, much of the
research has focused mainly on Western North Atlantic coastal waters. The
European Eastern Atlantic coasts have also been well studied (see Koehler 1921;
Mortensen 1925, 1927; Høiseater 1990; Hansson 2001), while Macaronesian
archipelagos (Azores, Madeira, Salvajes, Canary Islands and Cabo Verde) are less
known. Some studies have been done on the coasts of the Azores and Portugal
coasts (Barrois 1888; Nobre 1930, 1931, 1938; Chapman 1951; Marques 1983;
Pereira 1997). De Jesus and Abreu (1998) studied Madeira although sampling was
restricted to soft bottoms between 20 and 100 m depth. Recently, the echinoderm
fauna of the shallow rocky bottoms (0–25 m depth) of the Macaronesian archi-
pelagos has been surveyed in the context of the ‘‘Macaronesia 2000 Project’’
promoted by the Museo de la Naturaleza y el Hombre from Tenerife (Pérez-Ruzafa
et al. 2002) and Cabo Verde (Pérez-Ruzafa et al. 1999, 2003a, Entrambasaguas
2008; Entrambasaguas et al. 2008).
Echinoderms are one of the groups of marine animals that have received con-
siderable research effort in the Canaries. Because of its economic impact, research
interest has primarily concerned, and sometimes even monopolized, by one species:
the sea urchin Diadema aff. antillarum. This species have been recently described
and named as Diadema africana (Rodríguez et al. in press.). It is worthwhile to note
that researchers have used either D. antillarum or D. aff. antillarum to refer to the
Eastern-Atlantic species. We recommend the use of D. africana when referring to
eastern-Atlantic populations and have used this name throughout the text.
15.2.1 Pioneering Studies
The occurrence of echinoderms in the Canary Islands has been noted by a few
authors (Greef in Theel 1886; Panning 1939) and by expeditions in these waters.
Expeditions visiting the archipelago at the end of the 19th century and the first half
of the 20th include the Challenger, Travaillerur and Talisman, Princesse Alice,
Michael Sars, The Java-Sud Africa Expedition. As an example, Barrois (1888)
studied the echinoderm faunas of Azores and Madeira and made comparisons with
faunas from Canary Islands and Cabo Verde. Mortensen (1943) provided one of
the first references on echinoderms of the Canaries where he refers to the presence
of long-spined black sea urchins during his visit to Las Palmas (Gran Canaria) at
the beginning of 1930s. Reports from other foreign naturalists also emphasized,
several decades ago, large densities of this sea urchin in the shallow rocky subtidal
of the Canaries (Johnston 1969). This sea urchin (misidentified as Centrostephanus
longispinus) was even pointed out as ‘‘guilty’’ of the lack of algae in the rocky
subtidal through overgrazing (Johnston 1969).
15.2.2 1980s and 1990s-Establishing the Taxonomic Baselines

and the First Bionomical, Biogeographical and Ethological
Approaches
The first studies carried out by researchers from institutions on the Canary Islands
to describe the species composition of the echinoderm fauna are from the early
1980s. A team from the University of La Laguna, led by J.J. Bacallado, imple-
mented the research projects Bentos I and Bentos II (Bacallado et al. 1982). These
works focused on the collection, identification and taxonomic description of the
main animal groups of the marine bottoms of the Canary Islands, including
echinoderms (Bacallado et al. 1984, 1985). As a result of these projects the ori-
ginal checklist was improved (Bacallado et al. 1988).
The Bentos project was the base for studies on holothurians (Pérez-Ruzafa
1984, 1984a, b; Pérez-Ruzafa et al. 1985a, b, 1992a, b; Pérez-Ruzafa and Marcos
1987), including their spatial distribution (Pérez-Ruzafa et al. 1984a), biogeo-
graphical relationships (Pérez-Ruzafa et al. 1984b; Pérez-Ruzafa and Marcos
1987) and behavior (Pérez-Ruzafa et al. 1987). A morphological description of
spines for eight common species of echinoids was published by Moreno-Batet
et al. (1980). Of particular relevance for echinoids, was the publication of a
monograph on the main biological characteristics of Diadema africana (Bacallado
et al. 1987). This work reinforced previous observations of this research group
(Brito et al. 1984) on the important role this sea urchin seemed to play in the
structure and organization of shallow rocky bottoms of the archipelago. Impor-
tantly, this environmental issue took on a social dimension with the publication of
the book by Aguilera et al. (1994) about environmental sustainability of the
Canaries. These documents constituted the baseline on which subsequent works
were carried out, including taxonomic revisions and additions to all classes of
echinoderms.
15.2.3 The ‘‘recent’’ Past-An Explosion of Studies on Echinoderm

Ecology at the Start of the 21st Century
The end of the 20th is linked to an increase in studies on the ecology of echino-
derms from the Canaries. Of special importance is the project Macaronesia 2000
supported by the Museo de Ciencias Naturales de Tenerife and directed by J.J.
Bacallado. This project was done between 1998 and 2006 and was a great advance
for echinoderm studies in the Macaronesian islands (Azores, Madeira, Salvage
Islands, Canary Islands and Cape Verde). We have to emphasize the importance of
the doctoral thesis of L. Entrambasaguas (2009) and other works derived from this
project (Pérez-Ruzafa et al. 1999, 2002, 2003a; Entrambasaguas et al. 2008).
These and previous studies on taxonomy had their final zenith with the com-
pletion and subsequent publication of the BIOTA book, which compiled all tax-
onomic citations published until 2003 for each group of marine animals from the
Canarian Archipelago (Pérez-Ruzafa et al. 2003b). This document identified 91
species of echinoderms living in the waters of the Canary Islands (including two
subspecies): 15 echinoid species, 18 asteroids, nine crinoids, 15 ophiuroids and 34
holothurian species (Sect. 15.3 and Appendix).
Without doubt, the sea urchin D. africana was the center of attention for studies
on benthic ecology: high density of the sea urchin and the subsequent formation of
‘‘urchin barren grounds’’ (blanquizales—white bottoms with no erect macroalgal
cover as a result of overgrazing) catapulted the species to the spotlight of benthic
research in the Canary Islands. As a result, studies covered a suite of biological
and ecological topics: colonization of artificial substrates and interactions of this
sea urchin with other elements of the subtidal biota (Herrera 1998), patterns of
circadian movements and feeding preferences (Tuya 2002), reproductive biology
and genetic diversity (Garrido 2003), gametogenesis, larval development, settle-
ment and recruitment patterns (Hernández 2006), predation as a mechanism that
controls populations, including the role of marine protected areas (Clemente
2007), trophic ecology (Cabanillas 2009), and processes determining structure and
organization of benthic communities (Ortega 2010). Two projects: ‘‘Estudio de la
biología y ecología del erizo Diadema antillarum y de las comunidades algales de
sucesión en diferentes zonas de blanquizal del Archipiélago Canario’’ and
‘‘Canarias, Costa Viva’’, as well as monitoring studies to evaluate the effective-
ness of marine protected areas throughout the Canaries, have provided consider-
able data on the ecology of D. africana during 2001–2004. In particular, a large
body of research has described the relationships between size and abundance
patterns, and the connection between the abundance and size patterns of D. afri-
cana populations and algal and fish assemblages.
The end of the 20th century saw a drastic shift in the way science was evaluated
in Spain: a consolidation of peer-reviewed international indexed journals (included
in international databases where journals are ranked according to their scientific
relevance) as the main output of scientific publication. Therefore, the publication
of articles in peer-reviewed indexed journals on aspects related to the echinoderms
Fig. 15.3 Number of published articles in international, indexed, peer-reviewed journals on

aspects related to the sea urchin Diadema africana in the Canaries
from the Canaries has increased considerably in the last years (Fig. 15.3). These
works include some of the biological and ecological topics derived from the
doctoral theses and the projects noted above. Here, we compile these studies on the
biology and auto-ecology of D. africana, including reproductive biology (Garrido
et al. 2000; Hernández et al. 2006a), feeding preferences (Tuya et al. 2001),
nocturnal movements and ‘‘homing’’ behavior (Tuya et al. 2004a), seasonality of
settlement patterns and its coupling with the oceanographic scenarios of the
Canaries (Hernández et al. 2006b, 2010a), as well as technical topics related to the
in situ tagging to monitor movement (Tuya et al. 2003; Clemente et al. 2007a) and
predation (Clemente et al. 2007b). Another group of studies analyzed the spatial
and temporal variability of populations (Tuya et al. 2006a; Hernández et al.
2008a), including the effect of a set of environmental factors (e.g. turbulence
(Tuya et al. 2007), the topography of the reef and sedimentation (Clemente and
Hernández 2008), respectively human-induced factors such as fishing effort
(Hernández et al. 2008a) and the establishment of protection frameworks over the
coast (Hernández et al. 2008b). A large body of research analyzed the way D.
africana affects the composition, abundance and organization of algal assemblages
(Tuya et al. 2004b, 2004c, 2005). Two studies analyzed how the populations of
this echinoid interact with other parts of the animal biota, including the abundance,
diversity and organization of macro-invertebrate assemblages (Tuya et al. 2006b;
Ortega et al. 2009). Finally, recent investigations have focused on the way pre-
dation, mainly exerted by fishes, shape the abundance and size patterns of D.
africana in marine protected areas (Clemente et al. 2009, 2010) and non-protected
zones (Clemente et al. 2007b). Comparison of protected areas (marine reserves)
and zones subject to prominent exploitation that have qualitative and quantitative
distinct fish assemblages, in conjunction with previous observations (Tuya et al.
2004b; Hernández et al. 2008a), has been used to demonstrate the key role that
predation, chiefly induced by overexploited fish species, exert in the control of

populations of D. africana. Consequently, overexploitation of sea urchin predators
has been directly linked to large densities of D. africana, which causes the direct
elimination of erect algal beds and results in the prevalence of rocky unvegetated
bottoms in the rocky subtidal (‘‘urchin barren grounds’’). The disappearance of
these algal beds limits benthic primary production, as well as the amount of food
and shelter these algal habitats provide for associated invertebrates and fishes. This
exacerbates, even more, the deterioration of marine biodiversity in the shallow
subtidal waters of the Canarian Archipelago. Last, but not least, there is a positive
relation between the increases in densities of D. africana and the recent increase in
sea water temperature (Hernández et al. 2010a). This work is based on a historical
compilation of sea water surface temperature data from 1948 to the present and a
continuous collection of data on settlement, recruitment and densities from 2001 to
2008. The major conclusion is that the increase in sea water temperature, coupled
with the prevalence of ‘‘urchin barren grounds’’, enhanced the survivorship of D.
africana recruits. This work predicts an increase in the densities of this sea urchin
as well as the extension of ‘‘urchin barren grounds’’ under the predicted scenario of
raising temperatures. In conclusion, the rise in sea water temperature has been a
relevant factor in conjunction with overfishing of sea urchin predators to enhance
densities of the sea urchin.
The high level of social interest about the effect of high densities of D. africana
has not only increased the publication of scientific manuscripts. Publications have
disseminated the characteristics of the sea urchin to society in non-technical lan-
guage (Brito et al. 2004; Tuya et al. 2004d; Hernández et al. 2009). These pub-
lications have tried to raise awareness on the necessity to implement conservation
strategies to guarantee the sustainability and conservation of benthic communities
in the Canary Islands since high densities of the sea urchins are directly connected
with perturbed bottoms. Indeed, increasing interest on preserving the rocky sub-
tidal reefs of the Canaries has been consolidated by the development of the project
Diademar (Instituto Canario de Ciencias Marinas), which is exploring ways to
commercially exploit the high densities of the sea urchin, mainly for human
consumption of gonads, as a way of population control. In addition, the Vice-
consejería de Medio Ambiente and the Viceconsejería de Pesca of the Canary
Government and several insular administrations (Cabildos) have implemented a
program to periodically eliminate this sea urchin in large experimental plots across
the entire archipelago. The goal is to preserve algal beds (but see Sect. 15.5).
Other echinoderms have not received the same focus of attention within the
scientific community. There are only a few studies on other species of sea urchins,
such as the genetic diversity (Calderon et al. 2008) and a description of the
juvenile phases of Paracentrotus lividus (Hernández et al. 2005), and the spatial
distribution patterns of P. lividus and Arbacia lixula across depth gradients in the
shallow subtidal (Tuya et al. 2007). Other works focused on some ecological
aspects, such the relation between densities of D. africana and other species of
echinoderms (Ortega et al. 2009), particularly the holothurians (Tuya et al. 2006b).
The echinoderm fauna from the Canary Islands is presently composed of 127
species, nine of Crinoidea, 25 of Asteroidea, 42 of Ophiuroidea, 26 of Echinoidea
and 28 of Holothuroidea. Of these, 71 species inhabit the littoral zone, at depths
less than 50 m (one of Crinoidea, 16 of Asteroidea, 24 of Ophiuroidea, 13 of
Echinoidea and 17 of Holothuroidea) (see Appendix and pictures on Figs. 15.4–8
for the most common species). This biodiversity is similar to that reported for the
Mediterranean (147 species in total, 101 in shallow water) or the British Isles (119
and 71 respectively). It is higher than the reported in other archipelagos such as
Cabo Verde (66 species in total and 42 in shallow waters) but this is due probably
to differences in sampling effort.
The present status of the echinoderm fauna of the Archipelago needs a major
revision, because many citations come from old works and indirect references,
some species reported could be misidentifications. For example, Holothuria
(Lessonothuria) poli, was reported for the Canary Islands by Ludwig (Greef, in
Theel 1886), Herouard (Tortonese 1965) and for Palma by Tortonese (1965). We
have never found the species in our sampling of the different islands. In the same
way, the cluster composed of Holothuria (Holothuria) stellati, Holothuria (Hol-
othuria) tubulosa, Holothuria (Holothuria) mammata and Holothuria (Holothuria)
dakarensis, which have been cited for the archipelago, must be reduced to Hol-
othuria (Holothuria) mammata after the taxonomic and genetic review of the
group in Atlanto-Mediterranean waters by Borrero-Pérez et al. (2009). According
to their results, H. dakarensis would be restricted to the Cape Verde Islands and
the coast of West Africa, H. mammata to the Atlanto-Mediterranean area and H.
tubulosa to the Mediterranean Sea. Holothuria stellati was considered a junior
subjective synonym of H. tubulosa.
The small holothuroid Holothuria (Holothuria) helleri was reported by Pérez-
Ruzafa (1984) and Pérez-Ruzafa et al. (1984a) from samples of coral branches at
102 m depth, but the validity of this species must be reviewed as it could be
confused with juveniles of other Holothuria species.
Some confusion has existed with species with anfiatlantic distributions. Lessios
et al. (2001) divided Diadema antillarum into Western and Eastern Atlantic clades
from a molecular genetic analysis. This could also be the case for the small
brooding holothuria Neocnus incubans, reported at the Canary Islands as its only
locality of its distribution in the Atlantic Ocean (Pérez-Ruzafa et al. 1984a; Pérez-
Ruzafa et al. 1984b).
Shallow rocky bottoms are inhabited by few species. The more conspicuous are
the sea stars Ophidiaster ophidianus, Coscinasterias tenuispina and Marthasterias
glacialis, the holothuroid Holothuria (Platyperona) sanctori, and the sea urchins
Diadema africana, Arbacia lixula and Paracentrotus lividus. The three sea urchins
coexist at all depths down to 30 m, but some vertical zonation can be observed in
their distribution (Tuya et al. 2007). Paracentrotus lividus is herbivorous and
dominates in shallower areas, from 0 to 5 m depth, with algal cover and well
Fig. 15.4 a Antedon bifida (Pennant, 1777); b Thethyaster subinermis (Philippi, 1837) (Photo: J.
Barquín); c Chaetaster nodosus Perrier, 1875; d Asterina gibbosa (Pennant, 1777); e Hacelia
attenuata Gray, 1840; f Narcissia canariensis (d’Orbigny, 1839); g Ophidiaster ophidianus
(Lamarck, 1816); h Echinaster sepositus (Retzius, 1783)
Fig. 15.5 a Coscinasterias tenuispina (Lamarck, 1816); b Marthasterias glacialis (Linnaeus,

1758);c Astropartus mediterraneus (Risso, 1826); d Ophiolepis paucispina (Say, 1825); e
Ophioderma longicaudum (Retzius, 1789); f Ophiocomina nigra (Abildgaard, 1789); g
Ophiopsila aranea Forbes, 1845; h Amphipholis squamata (delle Chiaje, 1829)
Fig. 15.6 a Ophiothrix fragilis (Abildgaard, 1789); b–c Stylocidaris affinis (Philippi, 1845); d
Centrostephaus longispinus (Philippi, 1845); e Diadema africana (Rodríguez, Hernández,
Clemente 2010); f, g Arbacia lixula (Linnaeus, 1758); h Coelopleurus floridianus Agassiz, 1872
Fig. 15.7 a Sphaerechinus granularis (Lamarck, 1816); b–c Paracentrotus lividus (Lamarck,
1816); d Genocidaris maculata Agassiz, 1869; e Echinocyamus pusillus (Müller, 1776); f, g
Brissus unicolor (Leske, 1778); h Echinocardium cordatum (Pennant, 1777)
Fig. 15.8 a Neocnus incubans Cherbonnier, 1972; b Holothuria mammata Grube, 1840; c, d
Holothuria arguinensis Koehler and Vaney, 1906; e, f Holothuria sanctori delle Chiaje, 1823; g
Euapta lappa (Müller, 1850); h Leptosynapta inhaerens (Müller, 1776)
illuminated environments. Arbacia lixula is more abundant at slightly deeper

waters (1–15 m) and in sciaphilous assemblages, probably having a slightly higher
trophic level than P. lividus. Finally, D. africana is the most abundant species,
reaching densities up to 240 ind. per 100 m2 covering large extensions of un-
vegetated rock. It can reach depths up to 100 m. A less abundant species that can
be confused with Diadema is another long-spined sea urchin C. longispinus. The
sea star Hacelia attenuata can also be found on rocky and detritic bottoms.
On rocky substrata, but close to soft bottoms, the holothurians H. (H.) mammata
and Holothuria (Lessonothuria) arguinensis are common. The daily feeding
behavior of these species in Canary Islands has been described by Pérez-Ruzafa and
Marcos (1987). Holothuria (Vaneyothuria) lentiginosa lentiginosa inhabits rocky
and sandy bottoms deeper than 20 m. Holothuria (Panningothuria) forskali has
been collected on rocky substrata at 348 m depth at Tenerife Island (Pérez-Ruzafa
et al. 1987). This is the southernmost and greatest depth reported for the species.
Pérez-Ruzafa (1984) made the first report of the small incubating holothurian
Neocnus incubans in the Atlantic from the Canary Islands, it was always found on
Cystoseira spp. on the upper infralittoral. The small, incubating ophiuroid, Am-
phipholis squamata, is common in photophilous algae.
The ophiuroid Ophiothrix (Ophiothrix) fragilis, Ophiocomina nigra, Oph-
iomyxa pentagona and Ophioderma longicauda are common under stones and in
crevices. The nocturnal anfiatlantic apodid holothurian Euapta lappa is common in
cryptic habitats and is particularly abundant in the western part of the archipelago
where the waters are warmer. Other species of ophiuroids on rocky bottoms of the
archipelago are Ophioconis forbesi, Ophioderma appressum, Ophiocoma pumila,
Ophiactis savignyi and Ophiactis virens.
The sea stars Luidia ciliaris and Astropecten aranciacus and the sea urchin
Echinocardium cordatum occur on sandy bottoms. The sea urchin Brissus uni-
color, occurs on sandy bottoms under stones. The sea stars Astropecten irregularis
irregularis and Peltaster placenta, the ophiuroids Ophiura (Dictenophiura) carnea
and Ophiura texturata, the sea urchins Genocidaris maculata, Schizaster cana-
liferus, Spatangus purpureus, Plagiobrissus costai and the holothuroids H. (H.)
mammata and Thyone fusus occur on sandy and muddy bottoms in shallow water
and Eostichopus regalis and Stichopus tremulus in deeper water.
The sea urchin Echinocyamus pusillus and the ophiuroid Ophiura grubei are
common on detrictic bottoms. The ophiuroid Ophiacantha setosa can be found on
gorgonians.
15.3.1 Biogeographic Relationships of the Echinoderm

Fauna of the Canary Islands
The origin of the echinoderm fauna of the Canary Islands is mainly warm-tem-
perate (28 %) and with a wide thermal distribution (26 %) (Fig. 15.9). The strictly
Fig. 15.9 Origin and affinities of the echinoderm fauna of the Canary Islands in relation with the
thermal regions in the Atlantic described by Briggs (1996)
south boreal component is represented only by Mesothuria connectens reported for

the archipelago by Mortensen (1927) from material collected by the Talisman
expedition in 1975 at 2,518 m. However, the South-Boreal-Temperate component
constitutes the 19 % of the species. It includes the shallow water species Antedon
bifida, Luidia ciliaris, Asterina gibbosa, Echinaster sepositus, Henricia sangui-
nolenta, Ophiocomina nigra, Echinocyamus pusillus, Aslia lefebrei, Pawsonia
saxicola, Stichopus tremulus, Holothuria forskali and Leptosynapta inhaerens, and
the deep water species Ophiomitrella clavigera, Amphilepis norvegica, Ophiactis
abysicola, Araeosoma fenestratum and Araeosoma histrix. The tropical (13 %) and
tropical-warm temperate faunas (13 %) are well represented. Except for Euapta
lappa which occurs at depths between 10 and 20 m and Ophiolepis paucispina,
which reaches a maximum depth of 37 m, all strictly tropical species present at the
archipelago occur in deep waters or over a wide range of depths. These include
Astropecten hermatophilus (15-1,500 m), Narcisia canariensis (25–155 m), Or-
easter reticulatus and Astrophyton muricatum (0–70), Ophiomisidium pulchellum
(1,380–1,830 m), Ophiocoma pumila (0–368 m), Ophionereis sexradia (18–130),
Plesiodiadema antillarum (651–3,111), Diadema africana (0–400), Bathyplotes
pourtalesi (134–1,477 m) or Psychropotes semperiana (1,143–5,600 m). All of
these, except Astropecten hermatophilus, Narcisia canariensis and Ophionereis
sexradia.
The tropical-warm temperate component is represented mainly by Eastern
Atlantic-Mediterranean species as Astropecten aranciacus, Thethyaster subiner-
mis, Chaetaster longipes, Peltaster placenta, Hacelia attenuata, Ophidiaster
ophidianus, Ophiomyxa pentagona, Astropartus mediterraneus, Ophiacantha
bidentata, Ophiacantha setosa, Ophiacantha valenciannensi, Ophiura (Dicteno-
phiura) carnea, Ophiura grubei, Ophioderma longicaudum, Ophiopsila
Fig. 15.10 Relationship between the number of shared species of the different biogeographic
areas in the Atlantic and the Canary Islands and the distance to the archipelago. This relationship
is negative emphasizing the importance of oceanic currents and connectivity to explain the
composition of the fauna in oceanic islands. White circles: shallow water species (\50 m); Grey
circles: all depths
guineensis, Ophiactis virens, Ophiothrix maculata, Centrostephaus longispinus,

Brissopsis atlantica mediterranea, Holothuria (Holothuria) dakarensis, Holothu-
ria (Platyperona) sanctori, Holothuria (Vaneyothuria) lentiginosa lentiginosa and
Molpadia musculus. There are also some anfiatlantic species as Coscinasterias
tenuispina, Ophioderma appressum, Ophiactis savignyi, Arbacia lixula, Geno-
cidaris maculata, Brissus unicolor, Benthodites lingua and Psychropotes depressa.
There are few analyses on the distribution and biogeography of Atlantic echi-
noderms (Clark and Downey 1992). The relationships of the echinoderm fauna of
the Canary Islands with other Atlantic faunas were analyzed for the class Holo-
thuroidea (Pérez-Ruzafa et al. 1984b) and in the context of South-Western Med-
iterranean echinoderm fauna relationships (Pérez-Ruzafa and Marcos 1987).
The shallow water echinoderm fauna of Canary Islands is a consequence of its
oceanography and location in the Atlantic. The number of species shared with
other geographical areas is directly related to distance (Fig. 15.10). Therefore, of
the 71 species inhabiting the archipelago above 50 m depth, the highest number
corresponds to the Mediterranean (55 species), Portugal (50), the Cantabric (43)
and Mauritania and Guinea Gulf region (40). The low number shared with Mor-
occo (36) is probably due to the scarcity of works in this region. In fact, the
number of species increases to 54 when deep water fauna is considered because
most of the deep water expeditions which passed through the archipelago sampled
in both areas.
The lower number of shallow water shared species is with the Caribbean (12),
Brazil (13), and Cabo Verde Archipelago (20). In the case of the American coasts,
this can be explained because of the distance. But in the case of Cabo Verde, the
explanation has to do with the oceanographic currents system that precludes the
flux of tropical species from south to north in the Eastern Atlantic.
Fig. 15.11 Contribution to the echinoderm fauna of the Canary Islands of the different faunal
groups according to the biogeographic regions and provinces described by Briggs (1996). Figures
indicate the number of species shared and the percentage of the total
A 56 % of the species present in the archipelago has a wide distribution in the

Eastern Atlantic area (Fig. 15.11). Furthermore, 27 species (23 %) are from the
Lusitanian province (five from the Mediterranean, 13 from the Atlantic and nine
which are present in both areas). Three species (Astropecten hermatophilus,
Narcisia canariensis, Ophionereis sexradia) are shared exclusively with the
tropical Eastern Atlantic region (from Cabo Verde Islands and the Cape Verde on
the mainland to Angola (Briggs 1996). Only one species, the deep water holo-
thurian Mesothuria connectens, is exclusively from the Eastern-Atlantic Boreal
region. Finally, 13 species (13 %) are characteristic of the Caribbean-West Indian
and Brazilian provinces in the Western Atlantic.
Only 13 species have been reported exclusively from the Canarian Archipelago
waters: the sea stars Lonchotaster tartareus, Thoracaster cylindratos, Colpaster
scutigera, Freyastera tuberculata, the ophiuroids Ophiomusium serranum, Ophi-
omusium vallidum, Ophiozonella molesta, Ophiura inermis, Amphiura grandis, the
sea urchins Cidaris blakei, Salenocidaris rarispina, Aceste bellidifera and the
holothurian Peniagone ferruginea. All are deep water species occurring at more
Fig. 15.12 Cluster tree representing the affinities of the echinoderm faunas between different
geographic regions in the Atlantic. Distances were constructed with the Jaccard index for
presence-absence data. Note the close relationship between distances and differences in mean
annual surface water temperatures as showed by the satellite image at left
than 900 m depth. Diadema africana has been described for Madeira, Canary
Islands and Cape Verde, but as this species had been reported until now as Dia-
dema antillarum or Diadema aff. antillarum, a review of the distribution of the
species in the Eastern Atlantic must be done.
The application of the Jaccard index to the echinoderm fauna of these men-
tioned regions (including shallow and deep sea faunas) produces two main groups
(Fig. 15.12). The first includes British Islands, Cantabric Sea and Portugal coasts
and the Atlantic coast of Morocco, the Mediterranean and the Canary Islands. The
second group links the Caribbean and Brazil on one hand with Cabo Verde and the
Mauritanian and Guinean Gulf on the other. These groups represent the four major
thermal regions in the Atlantic surface waters: South Boreal, Cold-temperate,
Warm-temperate and Tropical (Briggs 1996). The tropical zone is delimited by the
20 °C isotherm for the coldest month.
15.4 Echinoderm Ecology: Implications for Conservation
Echinoids are by far the most abundant group of echinoderms of shallow waters of
the Eastern Atlantic Archipelagos. Few detailed ecological studies have focused
on other taxa in this region. Comparisons of spatial distribution of sea urchins A.
lixula, P. lividus and D. africana show a consistent depth-dependent partitioning
pattern on rocky reefs, in which the latter is the competitive dominant species both
at the Canary Islands (Tuya et al. 2007) and Madeira Island (Alves et al. 2001).
This depth gradient in the vertical distribution of the three sea urchin species has
been related to their distinct morphological characteristics, and consequently, their
different resistance to natural hydrodynamic forces (Tuya et al. 2007). Arbacia
lixula and P. lividus seem to be morphologically adapted to minimize the chances
of being dislodged by high water motion at the intertidal and first few meters of the
subtidal and coexist in relatively high densities only in the shallowest depths (\5
m). Densities of D. africana, less adapted to support intense water movement,
increase with depth. The result is a general pattern in the vertical distribution of
these sea urchins in the warm-temperate rocky reefs of the eastern Atlantic Ocean
(Alves et al. 2001, Tuya et al. 2007), in which wave exposure and island slope that
affects wave energy are major environmental factors determining densities of these
key herbivorous sea urchins (Hernández et al. 2008a). Moreover, protected coasts
have calmer waters that promote retention of urchin larvae (Hernández 2006).
Larvae at exposed sites larvae tend to be carried away. The result is that areas with
higher hydrodynamics show greater urchin density in a deep band (15–20 m) just
below the algal stand, whereas along sheltered coasts algal beds the urchins occur
only at the first meters depth where more food is available (Hernández et al.
2008a).
Because of the biological and ecological features, such as size, motility, feeding
preferences or longevity, several urchin species play a key role, even more
important than other herbivores, in controlling epibenthic communities and in turn
of the overall structure of coastal communities (Lawrence and Sammarco 1982).
The main consequence of this overall phenomenon of intense grazing activity is a
shift from large areas of rocky reefs covered by complex erect macroalgal beds to
overgrazed substrates (‘urchin barrens’ or ‘barren grounds’) dominated by
encrusting algae and some sessile invertebrates.
The influence of echinoids grazing activity on rocky reefs communities of the
eastern Atlantic oceanic islands, particularly in the Canary Islands, have been well
studied (Alves et al. 2001, Tuya et al. 2004c; Hernández et al. 2008b). These are
relatively small barren grounds produced by P. lividus in tidepools, and by A.
lixula on shallow bottoms and around artificial jetties (JC Hernández pers. obs.).
However, barren grounds resulting from the intense grazing activity of D. africana
are more extensive and commonly spread throughout the entire region (Aguilera
et al. 1994; Brito et al. 2004; Tuya et al. 2004c), reaching up to 50 m depth and
covering about 75 % of the total littoral rocky bottoms of the Canary Islands
(Barquín et al. 2004). Densities of this species can reach more than 12 ind m-2,
with severe consequences (Brito et al. 1984, 2004; Alves et al. 2001; Tuya et al.
2004b; Hernández 2006). Field monitoring and experimental manipulations of
urchin densities have shown its major effects on algal communities (Hernández
et al. 2008a). At moderate densities, sea urchin grazing can stimulate plant pro-
duction and promote species diversity. Because of this sea urchins should be
considered an integral part of ecosystem dynamics (Sammarco 1982; Valentine
et al. 1997). However, densities of as few as 4 urchins m-2 can remove up to
70–80 % of algal cover (Hernández et al. 2008a) and dramatically reduce
macroalgal diversity, resulting in the formation of impoverished sea urchin-

dominated areas denuded of all but encrusting algae (Lawrence 1975; Sangil et al.
2006a, 2006b). Compared to typical urchin barrens elsewhere in the world
(Chapman 1981; Pearse and Hines 1987; Valentine and Johnson 2005; Guidetti
and Dulcic 2007), barrens generated by D. africana in the eastern Atlantic appear
to be maintained by relatively low numbers of sea urchins (starting at 2 ind m-2).
Probably, the oligotrophic condition of waters surrounding the archipelago pro-
vides limited nutrient resources that are not sufficient to maintain extremely high
urchin densities. However, species movement and food consumption rates, which
involve grazing large areas of rocky substrate at night (Tuya et al. 2004a), allow
comparatively lower urchin densities to maintain a barren habitat denuded of most
erect macroalgae biomass.
Loss of macroalgal beds due to D. africana grazing activity produces not only
lower species diversity (Herrera 1998; Garrido 2003) but also a lack of habitat
suitable for fish feeding and reproduction (Brito et al. 2004; Tuya et al. 2005;
Clemente 2007). Macroalgae constitute a major energy source that maintains the
food web, and are the main biological engineers on rocky reefs of the Canary
Islands, which lie between tropical and temperate waters and has no coral reefs.
Consequently, algal cover provides the habitat that supports the whole system. It
constitutes a well-known indicator of benthic conservation status (Hernández et al.
2008b). Another good example of a third-order change that occurs as a conse-
quence of the development of urchin barrens is the distribution and abundances of
large holothurians, linked to specific type of habitats across shallow-waters of the
eastern Atlantic (Tuya et al. 2006b). This fact could be related to variations in the
availability of particulate organic matter across habitats. Benthic grazers, such as
sea urchins, can transform up to 70 % of the consumed algae to detrital fecal
matter (Vadas 1977; Koike et al. 1987; Frantzis and Gremare 1992; Lison de Loma
et al. 2000; Mills et al. 2000; Mamelona and Pelletier 2005). High densities of D.
africana on barren grounds likely produce large amounts of fecal pellets, as a
source of fresh particulate organic matter (Lawrence 1975; Wotton 1994; Law-
rence and Klinger 2001; Levinton et al. 2002; Mamelona and Pelletier 2005).
Consequently, holothurians were significantly more abundant in D. africana bar-
rens compared with vegetated habitats, a pattern that was strongest for the
numerically dominant species, Holothuria sanctorii (Tuya et al. 2006b).
As a major herbivore, D. africana is recognized as having a key ecological role
that threatens the resilience of marine systems in the eastern Atlantic, with the
subsequent establishment of undesired conservation states (Hernández et al.
2008a). The reported increasing frequency of overgrazing events, concomitant
with loss of associated ecosystem goods and services, generates a serious envi-
ronmental problem that constitutes a key threat to coastal marine ecosystems and
local communities of the islands. Hence, the necessity of managing the populations
of D. africana to prevent overgrazing management has become apparent, espe-
cially in areas of the Canary Islands where algae-associated services (such as
fishing stocks) are important for livelihood sustenance. Algal stands are very
restricted in space in the Canary Islands and have shown a temporal regression
related to the demographic explosion of the sea urchin during recent years (Cle-
mente 2007; Hernández et al. 2008a). High percentage cover is usually attained
only in shallow water stands at exposed sites where hydrodynamics prevent urchin
activity (Alves et al. 2001; Tuya and Haroun 2006), at certain marine protected
areas (Clemente 2007; Hernández et al. 2008a), or at barely perturbed areas
associated with a delayed anthropogenic development such as isolated and less
populated islands (Clemente et al. 2009).
Many factors are thought to be involved in the recent expansion of D. africana
populations and regression of algal beds at the eastern Atlantic Archipelagos. We
can indicate an array of environmental factors that promote high urchin densities,
such as substrate topography and diversity and certain oceanographic events that
can influence sea urchin settlement, recruitment and survival (Hernández et al.
2008a). In addition, several anthropogenic factors, such as the number of islanders
and tourists per coastal perimeter, the number of operational fishing boats and the
overfishing of the sea urchins’ natural predators have been associated with
increased urchin abundances (Hernández et al. 2008a; Clemente et al. 2009).
Diadema africana is a termophilus species, which probably constitutes a
competitive advantage for its proliferation at the sublittoral latitude of the Canary
Islands in contrast to sea urchins of temperate affinities such as P. lividus. This has
been especially relevant in the last decades, when a trend of progressive sea water
warming has occurred in the region as a consequence of global climate change
processes (Brito 2008). In fact, a long-term study has detected a clear relationship
between sea surface temperature and D. africana settlement (Hernández et al.
2010a). Urchin settlement was higher during warmer years. Temperatures above
24 °C produced major settlement pulses and higher post-settlement survival rates
in barren areas. However, some variability across habitats indicates resilience of
the systems. While sea water temperature is a major factor influencing D. africana
density in urchin barrens because it enhances post-settlement survival, specific
factors in healthy macroalgal beds constrain settlement, increases post-settlement
mortality and controls sea urchin populations (Hernández et al. 2010a). The study
shows that warmer summer temperatures (temperatures at settlement) have been
more frequent since the early 19800 s with a parallel increase in urchin population
densities (Hernández et al. 2008a). Assuming climate change predictions (IPCC
2007) for the future are correct, the strong positive relationship between thermal
history and post-settlement survival of this ecologically important species suggests
that barren grounds will become more extensive in the future as seawater tem-
peratures warm in relation to anthropogenic activities. Nonetheless, mass mortality
events, as found for P. lividus in 2003 (Girard et al. in press), could became more
frequent due to warmer waters. In this sense, we would like to point out that
researchers have observed mortality events of D. africana in Madeira and Gran
Canaria islands in 2010 (F. Alves, M. Kaufman and F. Espino pers. com.,
respectively). However, it is too soon to know the cause of this recent phenomenon
that, initially, seems to be very spatially restricted. In both cases, urchins were
infected by the bald sea urchin disease, and possibly originated by the bacteria
Vibrio alginolitycus (Hernández et al. 2010b).
Regulation of sea urchin populations also depends on certain trophic interac-

tions. Experimental studies have identified key fish species that successfully prey
upon D. africana and have the potential to control their populations (Clemente
2007; Clemente et al. 2011). This has important implications for coastal man-
agement. By controlling herbivore densities, and thus the amount of intact mac-
roalgal cover, predatory fish indirectly aid in preserving ecosystems service or
capacity to provide locally important fishing resources and coastal productivity in
sublittoral rocky reefs. The predatory fish observed are mainly species very sus-
ceptible to fishing (Clemente 2007). Consequently increased fishing activities have
been associated with lower numbers of these predatory species and higher urchin
densities in the Canary Islands (Clemente et al. 2009; 2011) and elsewhere (Sala
et al. 1998; Guidetti 2006; Guidetti and Dulčić 2007). Additionally, fishing
decreases the mean size of fish. This adds to the loss of species capable of con-
suming large sea urchins and resulting in a smaller escape by size from predation
that favour their proliferation. The removal of fish predators by severe overfishing
in most areas of the Canarian Archipelago have led to an urchin population out-
break driven only by settlement and recruitment in the absence of any kind of
predatory or top-down control (Hernández 2006; Clemente 2007). This ecological
disequilibrium, mediated by trophic cascade processes, has major consequences
for the structure and function of the entire benthic community.
Because top-down effects on D. africana at the eastern Atlantic archipelagos
depend on specific predatory species and their abundances within the food web,
preserving the diversity of fish predators must be a priority of conservation efforts
(Clemente et al. 2010). However, at this time this scientific knowledge has not
been fully taken into consideration by policy makers and no protection measures
have been directly applied to key predators of the sea urchin. There is only one
kind of effective management tool in force that can indirectly contribute to the
control of sea urchin populations. This is marine protected areas in which fishing
restriction measures are implemented to allow fishing stocks to recover (Her-
nández et al. 2008a). As elsewhere around the world (Sala et al. 1998; Babcock
et al. 1999; McClanahan et al. 1999; Guidetti 2006), studies in the Canary Islands
have stressed the value of marine protected areas as a tool for ecosystem resto-
ration; not only for fishes and fishing resources that increases predation on urchins
and indirectly aids regeneration of benthic communities (Hernández et al. 2008a;
Clemente et al. 2009). In fact, the three marine protected areas within the Canary
support higher densities of fish predators and higher levels of predation intensity
on D. africana compared to equivalent non-protected locations (Clemente 2007;
Clemente et al. 2011).
Regardless of the complexity of factors involved in regulation of overpopula-
tions of D. africana may be achieved by aiding the recovery of the predatory fish
community with appropriate fishing bans and protection measures. Current
knowledge about the species’ population dynamics has critical implications for the
conservation of rocky reef ecosystems in the eastern Atlantic Islands. They add to
a growing body of research suggesting that the incidence of barrens and their
associated impacts are related to human disturbance (Steneck 1998; Jackson et al.
2001). However, active management actions directed to control urchin population

have been limited to developing a strategy of manually removing sea urchin by
scuba divers, providing evidence that erect macroalgae have the potential to
recolonize encrusting macroalgae barrens (Brito et al. 2004), but of doubtful
outcome in the long-term. Combined perturbations caused by overfishing and
progressive sea water warming are likely to result in an environment increasingly
favourable to the sea urchin, which seems difficult to control with such small-scale
actions. Conservation policies need to focus efforts on specifically protecting
healthy macroalgal bed habitats and the predatory fishes within them, because they
are predicted to increase post-settlement mortality of D. africana. Such actions
could improve ecosystem function by retaining the equilibrium between sea
urchins and macroalgae.
Pre-hispanic human settlement in the Canary Islands, commondly known as

Guanches (Majos-Lanzarote and Fuerteventura; Canarii-Gran Canaria; Guanches-
Tenerife; Benahoritas-La Palma; Bimbaches-El Hierro) usually consumed sea
urchins. Indications of pre-hispanic human consumption have been discovered in
various shells accumulations locations in the islands. However sea urchin species
have been identified only at the ‘‘Guinea deposit’’ in Frontera-El Hierro (Martín
Oval et al. 1985). The species most consumed was P. lividus, probably because of
its abundance in intertidal rocky areas. Nowadays, this sea urchin is still consumed
by the islanders. However it remains only in particular coastal areas as a tradition
and no commercial exploitation exists. Presently, and probably then, this sea
urchin is also used as fish bait for fishing traps or to attract fishes to hook lines. In
some coastal locations this is the only known use. Today, other sea urchin species,
such as Arbacia lixula, Diadema africana and Sphaerechinus granularis, are also
used as bait. Arbacia lixula and D. africana are collected by snorkeling while S.
granularis, due its deeper distribution, is a bycatch of trammel nets. The number of
urchins used for this purpose is not known and is very difficult to monitor due to
the diversity of discharging places, many of them located at inaccessible sites.
Paracentrotus lividus is commercially exploited in several European countries
such as Ireland, France, Portugal, Italy, Greece and Spain. In the Spanish Penin-
sula, it is mainly exploited in the northwest (Galicia), north (Cantabria, Asturias y
Pais Vasco), northeast (Cataluña), and in the south (Andalucía). Although, it is
locally consumed in all these Spanish Comunidades, France is the main consumer
of sea urchins in Europe and the second consumer in the world after Japan
(Grosjean 2001). At the present, some of the most important fisheries in Europe
like the ones in Ireland and Brittany have collapsed (Le Direach 1987; Andrew
et al. 2002). In Spain, fishing activities regulations depend on the ‘‘Comunidad
Autónoma’’. While Galicia has a well developed sustainable fishing activity
(Catoira-Gòmez 1992), others like Asturias, Cantabria and Andalucía have a
growing semi-professional activity. However, the general overfishing situation of

the traditional resources could favour an increase of sea urchin fisheries in the near
future (Thorpe et al. 2000). At this time accurate stock studies are needed to
promote appropriate regulations for a sustainable fishing activity and prevent the
collapse of this resource.
Human fishing activity effects on P. lividus populations, although it has not
been studied in the Canary Islands, are evident in many intertidal habitats of the
islands compared to remote or marine protected areas locations. In these areas, of
human use, P. lividus is scarce and is restricted to inaccessible deep cavities in the
basaltic volcanic rock. In the Canary Islands, fishing activities in the intertidal are
regulated. Only Sport Fishing Licence holders can fish for shellfish during
weekends and holidays. No number or close season regulations exist. In our
opinion, this is an insufficient measure to regulate shellfish fishing activities that
favors overfishing in many intertidal benches. This situation has focused the
attention of managers who have recently funded a National Project to optimize and
promote P. lividus culture in Galicia and in the Canary Islands. This project aims
to improve the technical and biological aspect of larval culture and to develop
culture systems at different locations along the Spanish coast to provide juveniles
for farms or repopulation. This project was supported by the Ministerio de Medio
Ambiente y Medio Rural y Marino through the Junta Asesora de Cultivos Marinos
(JACUMAR). It has contributed to the improvement of spawning and fertilization,
larval culture and metamorphosis-settlement.
In addition to human collection, other factors such as temperature and com-
petition also modify the abundance of P. lividus. The Canary Islands are the
southern-most limit of this temperate species. It is more abundant off the eastern,
colder islands. Although, there are no specific studies that assess the temporal
abundance of this sea urchin, we have noticed that anomalous warm summers like
occurred in 2003 were associated with localized mass mortalities at several
intertidal and subtidal locations in Tenerife (Girard et al. in press). The sea water
warming trend in the Canarian Arquipelago (Hernández et al. 2010a) will probably
increase the frequency of these mortality events in the future, which would
decrease the intertidal populations. Bathymetric segregation of P. lividus occurs in
the littoral area. It is more abundant in shallow waters between 1–5 m (Tuya et al.
2007). This bathymetric segregation is mainly caused by competition with the
grazing of the abundant D. africana which has greatly reduced food resources and
has limited P. lividus distribution to shallow macroalgae stands refuges. Otherwise
it could reach deeper waters as in the Mediterranean Sea. In summary, the
depletion of macroalgae stands by D. africana, sea water warming and human
collection is modifying the potential distribution of P. lividus in the archipelago.
In recent years, due to the outbreak of D. africana, local management agencies
have become interested in the development of a commercial sea urchin fishery. At
the same time, these agencies are proposing reducing activity of D. africana at
some sites to gain in the conservation status of rocky bottoms of the Canary
Islands. However, both strategies have the potential to restore local unproductive
barren grounds to algal stands (Brito et al. 2004). These strategies should be
combined with other conservation measures such as the exclusion of urchin pre-
dators from fishing and protection of algal stand ecosystems. Combining these
strategies would ensure that the ecosystem would keep the balance between
urchins and algae. Urchin removals alone are inadequate (but see Sect. 15.4). In
general, the management of urchin outbreak cases around the world has consisted
in urchin removal. However better results have been always obtained when this is
done inside a marine protected area (Tegner and Dayton 1977; McClanahan et al.
1996). This kind of management measures has been proposed for the Canary
Islands several times (Clemente 2007; Hernández et al. 2008a; Clemente et al.
2009; Clemente et al. 2011).
These combined actions would potentially decrease the frequency and duration
of urchin overgrazing but limited knowledge of direct and indirect effects of these
ecosystem scale actions makes it difficult to assess the applicability and sustain-
ability of this method. Therefore we call for governmental responsibility and rigor
when planning this kind of action. Uncontrolled removal of urchins during the
planned fishery could cause urchin populations to ‘‘crash’’ with potential eco-
logical and economic side effects. The recent ‘‘boom and bust’’ trend of sea urchin
fisheries around the word (USA, Mexico, Canada) (Andrew et al. 2002; Uthicke
et al. 2009), most of them considered nowadays as overexploited, indicates the
need for more information on basic population parameters of species when
implementing removal actions (Russell 2000). Rigorously evaluating population
demographics has, in the case of the species D. africana, both ecological and
economic significance, especially now that it is beginning to be commercially
harvested. We encourage seeking sustainable solutions to the increasing popula-
tions of the sea urchin D. africana, trying to contribute to the correct management
and conservation of the fragile marine resources of the Canary Islands. However,
we have to develop standards for the commercial exploitation of this species of
what we lack today.
15.6 Recomendations
We have divided our recommendations into three major sections: biodiversity,

ecology and fisheries. These constitute the main research lines that currently drive
research on echinoderms in the Canarian Institutions (universities and research
institutes). Moreover, in each section we have pointed out research gaps that
should be considered for future studies in the archipelago.Biodiversity
– Accurate information about spatial distribution of echinoderm fauna is needed
to precisely identify species habitats and depths ranges.
– Genetics studies of some taxonomic groups are needed to clarify biogeo-
graphical patterns and taxonomic identification problems (e.g. Arbaciella ele-
gans is thought to be small Arbacia lixula).
– Field explorations should be extened to deeper habitats such as rhodoliths and

deep coral communities. These have been poorly studied and probably contain a
rich echinoderm fauna.
Ecology
– Population dynamics of key sea urchin species should be monitored, including
their influence in triggering transition phases between algal stands and unpro-
ductive urchin barren grounds.
– The effectiveness of marine protected areas as tool to prevent and revert urchin
barren situations should be studied.
– The effects of sea water warming and acidification on the calcification processes
in larvae and adults of echinoderms and its relevance on population dynamics
should be analyzed. The influence of environmental parameters is of special
importance for species of echinoderms that modify the habitat and whose
activity can greatly affect the whole system.
– The magnitude of the recent mortality events of D. africana in Maderia and the
Canary Islands, and any related potential ecosystem wide effects should be
determined.
Fisheries and aquaculture
– Tagging-recapture studies should be done to determine age, growth rate and
natural mortality of sea urchins. These studies, combined with other population
surveys, will provide baseline data to develop an adequate strategy of coastal
management for the planned fishery of D. africana.
– Experimental reduction of sea urchin densities inside Marine Protected Areas
that are still affected by dense urchin barren grounds should be done to identify
effective strategies that could promote ecosystem transitions toward productive
macroalgal stands. The potential reconstruction of the trophic structure at
marine protected areas, where fishing activities have ceased during several
years, may favour the control of newly settled urchins after reduction in density.
– Techniques need to be developed for production of juvenile sea urchin and sea
cucumbers through laboratory cultures for aquacture or repopulation.
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Yanes A (1990) Morfología litoral de las islas Canarias Occidentales. Secretariado Publicaciones
Universidad de La Laguna, La Laguna
Yanes A, Marzol MV, Romero C (2006) Characterization of sea storms along the coast of
Tenerife, the Canary Islands. J Coastal Res 48:124–128
Chapter 16
Latin America Echinoderm Biodiversity
and Biogeography: Patterns and Affinities
A. Pérez-Ruzafa, J. J. Alvarado, F. A. Solís-Marín, J. C. Hernández,

A. Morata, C. Marcos, M. Abreu-Pérez, O. Aguilera, J. Alió, J. J.
Bacallado-Aránega, E. Barraza, M. Benavides-Serrato, F. Benítez-
Villalobos, L. Betancourt-Fernández, M. Borges, M. Brandt, M. I.
Brogger, G. H. Borrero-Pérez, B. E. Buitrón-Sánchez, L. S. Campos,
J. R. Cantera, S. Clemente, M. Cohen-Renfijo, S. E. Coppard, L. V.
Costa-Lotufo, R. del Valle-García, M. E. Díaz de Vivar, J. P. Díaz-
Martínez, Y. Díaz, A. Durán-González, L. Epherra, M. Escolar, V.
Francisco, C. A. Freire, J. E. García-Arrarás, D. G. Gil, P. Guar-
deras, V. F. Hadel, A. Hearn, E. A. Hernández-Delgado, A. Herrera-
Moreno, M. D. Herrero-Pérezrul, Y. Hooker, M. B. I. Honey-Esca-
ndón, C. Lodeiros, M. Luzuriaga, C. L. C. Manso, A. Martín, M. I.
Martínez, S. Martínez, L. Moro-Abad, E. Mutschke, J. C. Navarro,
R. Neira, N. Noriega, J. S. Palleiro-Nayar, A. F. Pérez, E. Prieto-Ríos,
J. Reyes, R. Rodríguez-Barreras, T. Rubilar, T. I. Sancho-Mejías, C.
Sangil, J. R. M. C. Silva, J. I. Sonnenholzner, C. R. R. Ventura, A.
Tablado, Y. Tavares, C. G. Tiago, F. Tuya and S. M. Williams
A. Pérez-Ruzafa (&) A. Morata C. Marcos

Departamento de Biología Animal Biodiversidad, Ecología Marina y Conservación
(Ciencias Marinas), Facultad de Biología, Universidad de La Laguna, Tenerife, islas
Canarias, Spain
e-mail: angelpr@um.es
A. Morata
e-mail: aixamaria.morata@um.es
C. Marcos
e-mail: cmarcos@um.es
J. J. Alvarado S. M. Williams
Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa
Rica, San Pedro, San José, 11501-2060, Costa Rica
S. M. Williams
e-mail: stcmwilliams@gmail.com

512 A. Pérez-Ruzafa et al.
F. A. Solís-Marín
Colección Nacional de Equinodermos ‘‘Ma. E. Caso Muñoz’’, Laboratorio de Sistemática y
Ecología de Equinodermos, Instituto de Ciencias del Mar y Limnología (ICML),
Universidad Nacional Autónoma de México (UNAM), Apdo, 70-305 04510, Mexico, D.F.,
Mexico
J. C. Hernández S. Clemente
Departamento de Biología Animal (Ciencias Marinas), Universidad de La Laguna,
Biodiversidad, Ecología Marina y Conservación, c/Astrofísico Francisco Sánchez sn, La
Laguna, Tenerife, Canary Islands 38206 , Spain
e-mail: jocarher@ull.es
M. Abreu-Pérez
Instituto de Oceanología, Ministerio de Ciencias Tecnología y Medio Ambiente (CITMA), ,
Cuba
O. Aguilera
Museu Paraense Emilio Goeldi, Coordenação de Ciências da Terra e Ecologia,
Departamento de Geociências Av. Perimetral, 1901 Terra Firme, CEP, PA, 66077-830,
Brazil
e-mail: orangel.aguilera@gmail.com
J. Alió
Centro de Investigaciones Agrícolas de Sucre y Nueva Esparta, Instituto Nacional de
Investigaciones Agrícolas, Estado Sucre, 6101, Cumaná, Venezuela
e-mail: jalio@inia.gob.ve
J. J. Bacallado-Aránega
Museo de La Naturaleza y el Hombre de Tenerife, Santa Cruz de Tenerife, islas Canarias,
Spain
E. Barraza
Gerencia de Vida Silvestre, Ministerio de Medio Ambiente y Recursos Naturales, San
Salvador, El Salvador
e-mail: ebarraza@marn.gob.sv
M. Benavides-Serrato G. H. Borrero-Pérez
Instituto de Investigaciones Marinas y Costeras—INVEMAR, A.A 1016, Santa Marta,
Colombia
G. H. Borrero-Pérez
e-mail: giomar_borrero@invemar.org.co
F. Benítez-Villalobos J. P. Díaz-Martínez
Instituto de Recursos, Universidad del Mar, Carretera a Zipolite Km 1.5, Puerto Ángel,
Oaxaca, Mexico
e-mail: fvb@angel.umar.mx
J. P. Díaz-Martínez
e-mail: lobi2@hotmail.com
L. Betancourt-Fernández A. Herrera-Moreno
Programa EcoMar, Inc., Santo Domingo, Dominican Republic
e-mail: ongprogramaecomar@yahoo.com
M. Borges
Museu de Zoologia, Universidade Estadual de Campinas, Campinas-SP, 13083-863, Brazil
e-mail: borges@unicamp.br
16 Latin America Echinoderm Biodiversity and Biogeography 513
M. Brandt
Department of Ecology and Evolutionary Biology, Brown University, Providence, RI, USA
e-mail: margarita.brandt@gmail.com
M. I. Brogger M. I. Martínez A. Tablado
Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’-CONICET, Buenos Aires,
Argentina
e-mail: mbrogger@bg.fcen.uba.ar
M. I. Martínez
e-mail: mmartinez@macn.gov.ar
A. Tablado
e-mail: tablado@macn.gov.ar
B. E. Buitrón-Sánchez
Departamento de Paleontología, Instituto de Geología UNAM, Mexico, DF 04510, Mexico
e-mail: blancab@servidor.unam.mx
L. S. Campos
Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro,
Rio de Janeiro-RJ, 21941-971, Brazil
e-mail: campos-lucia@biologia.ufrj.br
J. R. Cantera K. R. Neira
Departamento de Biología, Universidad del Valle, A.A. 25360, Cali, Colombia
e-mail: jaime.cantera@correounivalle.edu.co
R. Neira
e-mail: rneira56@yahoo.com
M. Cohen-Renfijo
Centre d’Oceanologie de Marseille, Université de la Méditerranée, Marseille Cedex, France
e-mail: ablocsberg@gmail.com
S. E. Coppard
Smithsonian Tropical Research Institute, PO BOX 0843-03092, Balboa, Ancón, Panamá
e-mail: CoppardS@si.edu
L. V. Costa-Lotufo
Departamento de Fisiologia e Farmacologia, Instituto de Biologia, Universidade Federal do
Rio de Janeiro, Rio de Janeiro-RJ, 21941-971, Brazil
e-mail: lvcosta@secrel.com.br
R. del Valle-García
Instituto de Oceanología, Ministerio de Ciencias Tecnología y Medio Ambiente (CITMA), ,
Cuba
e-mail: rosa.denisse1@hotmail.cominformatica@giron.sld.cu
M. E. Díaz de Vivar
Facultad de Ciencias Naturales, Universidad Nacional de la Patagonia San Juan Bosco,
Puerto Madryn, Argentina
e-mail: mediazdevivar@gmail.com
Y. Díaz A. Martín
Laboratorio de Crustáceos Peracáridos. Centro de Biodiversidad Marina. Departamento de
Estudios Ambientales - INTECMAR, Universidad Simón Bolívar, Caracas, 1080, Venezuela
e-mail: yusdiaz@usb.ve
L. Epherra T. Rubilar
Centro Nacional Patagónico-CONICET, Puerto Madryn, Argentina
e-mail: epherra@cenpat.edu.ar
T. Rubilar
e-mail: rubilar@cenpat.edu.ar
M. Escolar
Instituto Nacional de Investigación y Desarrollo Pesquero, Mar del Plata, Argentina
e-mail: marianaescolar@inidep.edu.ar
V. Francisco
Laboratorio de Ecología de Ecosistemas de Arrecifes Coralinos, Centro de Investigación y
de Estudios Avanzados del Instituto Politécnico Nacional, Mérida, Yucatán, Mexico
C. A. Freire
Departamento de Fisiologia, Universidade Federal do Paraná, Curitiba-PR, 81531-990,
Brazil
e-mail: cafreire@ufpr.br
J. E. García-Arrarás E. A. Hernández-Delgado R. Rodríguez-Barreras
Department of Biology, University of Puerto Rico, Río Piedras, PO Box 23360, San Juan,
00931-3360, PR, USA
e-mail: jegarcia@hpcf.upr.edu
E. A. Hernández-Delgado
e-mail: coral_giac@yahoo.com
R. Rodríguez-Barreras
e-mail: ruber.rodriguez@hotmail.com
D. G. Gil
Instituto de Desarrollo Costero, Universidad Nacional de la Patagonia San Juan Bosco,
Comodoro Rivadavia, Argentina
e-mail: gil_damian@hotmail.com
P. Guarderas
GEOcentro, Universidad San Francisco de Quito, Cumbayá, Ecuador
e-mail: pagigv@yahoo.com
V. F. Hadel C. G. Tiago
Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao-SP, 11600-000,
Brazil
e-mail: vafhadel@usp.br
C. G. Tiago
e-mail: cltiago@usp.br
A. Hearn
Department of Wildlife, Fish and Conservation Biology, University of California, Davis,
CA, USA
e-mail: arhearn@ucdavis.edu
M. D. Herrero-Pérezrul
Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico Nacional, Ave. IPN S/n
Col., Playa Palo de Santa Rita, CP, 23096, La Paz, BCS, Mexico
e-mail: dherrero@ipn.mx
Y. Hooker
Laboratorio de Biología Marina, Departamento de Ciencias Biológicas y Fisiológicas,
Universidad Peruana Cayetano Heredia, Av. Honorio Delgado 430, Urb. Ingeniería, S.M.P,
Lima, Peru
e-mail: hookery@yahoo.com
Y. Hooker
Unidad Marino Costera, Servicio Nacional de Áreas Naturales Protegidas (SERNANP),
Ministerio del Ambiente, Calle Diecisiete No 355, Urb. El Palomar—San Isidro, Lima, Peru
M. B. I. Honey-Escandón E. Prieto-Ríos
Posgrado en Ciencias del Mar y Limnología, Instituto de Ciencias del Mar y Limnología
(ICML), UNAM, Apdo, 70-305 04510, Mexico, D.F., Mexico
C. Lodeiros
Instituto Oceanográfico de Venezuela, Universidad de Oriente y Fundación para la
Investigación y Desarrollo de la Acuicultura del Edo. Sucre, 6101Cumaná, Venezuela
e-mail: cesarlodeirosseijo@yahoo.es
M. Luzuriaga
Laboratorio de Plancton e Ictioplancton, Instituto Nacional de Pesca del Ecuador,
Guayaquil, Ecuador
e-mail: mluzuriagav@yahoo.com
C. L. C. Manso
Laboratório de Invertebrados Marinhos, Universidade Federal de Sergipe, Itabaiana-SE,
49500-000, Brazil
e-mail: cynthia@phoenix.org.br
S. Martínez
Departamento Evolución de Cuencas, Facultad de Ciencias, Iguá 4225, 11400, Montevideo,
Uruguay
e-mail: smart@fcien.edu.uy
L. Moro-Abad
Consorcio Insular de La Reserva Mundial de La Biosfera La Palma, Servicio de
Biodiversidad del Gobierno de Canarias, ,
E. Mutschke
Laboratorio de Ecología y Ciencias Ambientales, Universidad de Magallanes, ,
e-mail: erika.mutschke@umag.cl
J. C. Navarro
LECROY S.A., Km 2.5, Avenida Carlos Julio Arosemena, Bodega No.5, Guayaquil,
Ecuador
e-mail: jcnavarrogo@gmail.com
N. Noriega
Laboratorio de Biología Marina y Laboratorio de Bioquímica, Universidad Simón Bolívar,
Caracas, 1080, Venezuela
e-mail: nicidanoriega@gmail.com
J. S. Palleiro-Nayar
Centro Regional de Investigación Pesquera, Instituto Nacional de Pesca, Apartado postal
1305, Ensenada, BC, México
e-mail: juliopalleiro@yahoo.com.mx
A. F. Pérez
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos Aires,
Argentina
e-mail: analiafperez1@gmail.com
E. Prieto-Ríos
Facultad de Ciencias Biológicas, Biología, Universidad Nacional Mayor de San Marcos,
Mesa de partes. Ciudad Universitaria de San Marcos, Av. Venezuela s/n, Lima 1, Peru
16.1 Introduction
The first attempt to study the diversity and biogeography of echinoderms from
Latin America was done by Maluf (1988a, b). She identified 630 species of
echinoderms from southern California to southern Peru (Central Eastern Pacific or
J. Reyes
Laboratorio de Zoología de Invertebrados, Universidad del Zulia, Maracaibo 4001 y
Fundación para la Investigación y Desarrollo de la Acuicultura del Edo. Sucre, Cumaná,
6101, Venezuela
e-mail: jenyreyesl@yahoo.es
T. I. Sancho-Mejías
Museo de Zoología, Escuela de Biología, Universidad de Costa Rica, San José, Costa Rica
e-mail: sancho.ekin02@gmail.com
C. Sangil
Departamento de Ecología e Hidrología, Universidad de La Laguna, Biodiversidad,
Ecología Marina y Conservación
J. R. M. C. Silva
Departamento de Biologia Celular e do Desenvolvimento, Instituto de Ciências Biomédicas,
Universidade de São Paulo, São Paulo-SP, 05508-990, Brazil
e-mail: jmcs@usp.br
J. I. Sonnenholzner
Universidad Laica Eloy Alfaro de Manabí, Departamento Central de Investigación, ,
e-mail: jsonnenholzner@hotmail.com
C. R. R. Ventura
Museo Nacional, Universidade Federale do Rio de Janeiro, Rio de Janeiro-RJ, 20940-040,
Brazil
e-mail: ventura@acd.ufrj.br
Y. Tavares
Faculdade de Ciências, Filosofia e Letras de Paranaguá, Universidade Estadual do Paraná,
Pontal do Paraná-PR, 83203-280, Brazil
e-mail: ytavares@ufpr.br
F. Tuya
Departamento de Biología, Universidad de Las Palmas de Gran Canaria, Las Palmas de
Gran Canaria, Campus Tafira s/n, Las Palmas de G.C., Islas Canarias 35017, Spain
A. Durán-González
Universidad Nacional Autónoma de México (UNAM), Apdo, 70-305 04510, ,
M. B. I. Honey-Escandón
Universidad Nacional Autónoma de México (UNAM), Apdo, 70-305 04510, Mexico, D.F.,
Mexico
e-mail: mbihoney@hotmail.com
CEP). Two-thirds of all CEP echinoderms occur on the continental shelf

(depth \200 m), with a low level of endemism. She found an increase in species
richness from higher to lower latitudes, with peaks of richness in the Gulf of
California, Panama and the Galápagos Archipelago, places that also have a higher
number of endemic species, and are also areas with more research. The Gulf of
California had a greater similarity with the tropics than the Pacific side of the Baja
California peninsula and southern California, while the oceanic island of Cocos
was more similar to the mainland than the Galápagos and Revillagigedo archi-
pelagos. Moreover, she indicated the presence of five faunal transition zones
related to large-scale abiotic parameters. Those transition zones were: (1) the
Galápagos Islands, (2) Gulf of Guayaquil, (3) Costa Rica–Panama, (4) the mouth
of the Gulf of California and (5) Central Baja California outer coast. Seventy
percent of all the 265 CEP shelf restricted species were endemic to the region,
12 % were of a Northeast Pacific affinity (Oregonian and Alaska provinces), 5 %
had a Southeast Pacific affinity (Magellanic and Peruvian provinces), and 13 %
were widespread species. Most species in this last category were transpacific more
than circumtropical, and were species associated with reefs and rocky shores
(Maluf 1988b).
On the Atlantic side, Price et al. (1999) analyzed a database of presence/
absence of the Asteroidea compiled by Clark and Downey (1992) that was based
on records collected over 150 years. They determined geographical patterns of
diversity and make comparisons between coastal and deep-sea diversity for 26
regions of the Atlantic. For 349 species, they found a higher degree of endemism
in coastal waters and a greater level of similarity between regions with increasing
depth. However, the number of species between bathymetrical categories was
similar: 199 species at depths between 0 and 200 m, 135 and 169 species between
200 and 500 m, and 198 species at depths [500 m. The greatest number of species
(81 species) was in the region between Florida and the Yucatan Peninsula, fol-
lowed by the Bahamas-Caribbean (77 species) and the Caribbean and Guyana
basins together with the Gulf of Mexico region (66 species). This last region
possessed the highest number of endemic species of all the Atlantic Asteroidea (14
species). Regions like Guyanas-Cape Frío, Cape Frío-Río de la Plata, Río de la
Plata-Tierra del Fuego-Falklands islands and the Canary islands had few endemic
species (richness: 48, 22, 43, 26 species respectively; endemism: 1, 1, 8, 0 species
respectively). According to Price et al. (1999) the disparity in sampling efforts is
an important factor which can influence interpretation of geographic patterns.
Alvarado and Cortés (2004), Alvarado et al. (2008, 2010) and Alvarado (2011)
studied the diversity of echinoderms in Central America, covering both Pacific and
Caribbean coast. On both coasts the class Ophiuroidea was the richest with 85
species on the Pacific side and 79 on the Caribbean coast. Panama was also the
richest country in the region on both coasts, with 253 species on the Pacific coast
and 154 species on the Caribbean coast. Alvarado et al. (2010) and Miloslavich
et al. (2010) made an extensive review of published records on echinoderms of the
Caribbean and reported a total of 433 species. Ophiuroidea was the richest class
with 148 species, followed by Asteroidea (116 species). Mexico and Colombia
were the richest countries with 182 and 180 species, respectively. In terms of
Caribbean ecoregions, the Southwestern Caribbean was richest (283 species),
followed by the Western Caribbean (268 species), the Greater Antilles (248 spe-
cies), the Southern Caribbean (151 species), and lastly the Eastern Caribbean (79
species).
Miloslavich et al. (2011) analyzed the marine biodiversity of South America
(including the Pacific coast of Costa Rica and Panama, and excluding the Carib-
bean coast from Colombia and Venezuela). The best known groups in the region
are fish, mollusks, crustaceans, echinoderms, cnidarians, and macroalgae. They
divided the region into five subregions: (1) Eastern Tropical Pacific (223 species of
echinoderms, 3.3 % of the total species, 4.4 spp./100 km of the coast, 51 % spp. in
OBIS–Ocean Biogeographic Information System), (2) Humbolt Current-Chile and
Peru (364 species of echinoderms, 3.6 % of the total species, 5.0 spp./100 km of
the coast, 38 % spp. in OBIS), (3) Patagonian shelf-Uruguay and Argentina (207
species of echinoderms, 5.5 % of the total species, 3.7 spp./100 km of the coast,
76 % spp. in OBIS), (4) North, South and East Brazilian shelves (254 species of
echinoderms, 2.8 % of the total species, 3.4 spp./100 km of the coast, 60 % spp. in
OBIS) and (5) tropical west Atlantic-Venezuelan Atlantic, Guyana, Suriname and
French Guyana (107 species of echinoderms, 3.9 % of the total species, 5.7 spp./
100 km of the coast, 84 % spp. in OBIS). According to their analysis with the
OBIS database, echinoderms have a low level of endemism (3.6 %) and seem to
have reached a relatively stable number with few new additions. This could be the
result of the lack of taxonomic expertise, limited funding for research, lack of
collecting effort, and limited access to sampling sites.
Sthör et al. (2012) presented a global biodiversity analysis of the class
Ophiuroidea, reporting 2,064 species. They divided Latin America into three
regions: East Pacific (EP), South America (SA) and West Atlantic (WA). West
Atlantic was the richest region with 335 species with 60.6% of them endemic to
the region, followed by EP with 186 species and 62.9 % endemics, and SA with
124 species and 24.2 % endemics. Moreover, according to their depth strata, the
bathyal stratum (depths between 200 and 3,500 m) was the richest in the three
regions (229 species in WA, 111 species in EP and 102 species in SA), followed
by the shelf stratum (depths between 0 and 200 m) (217 spp. in WA, 92 spp. in EP
and 79 spp. in SA). The abyssal (depths between 3,500 and 6,000 m) (16 spp. in
WA, 28 spp. in EP and 17 spp. in SA) and the hadal strata (depths [ 6,500 m) (0
sp. in WA, 1 sp. in EP and SA) were less rich.
On this chapter we analyze the database presented in the appendix of this book.
We investigate the current patterns of diversity by country and by class of echi-
noderms. In addition, we analyze their biogeographical, depth, and habitat or
substratum affinities.
Fig. 16.1 Biogeographical regions in central and south American coasts according to Briggs
(1995) and Clark and Downey (1992)
16.2 General Oceanographic Variables of Latin America

and the Canary Islands
Traditionally, the area has been divided into five biogeographical regions with nine
provinces (Briggs 1995) (Fig. 16.1). Western Atlantic Region, from Bermuda
(Southern Florida) to Cape Frío (Brazil), which includes the Caribbean, Brazilian
and West Indian provinces. In the Pacific Ocean, the Eastern Pacific Region
includes the Mexican, Panamanian and Galápagos Provinces, the Eastern South
America Region with the Peru-Chilean Province and Easter Island in the central
south Pacific. In the Eastern Atlantic, the Canary Islands belong to the Lusitania
Region.
Together they cover a wide biogeographical and climate range (Table 16.1).
They are affected by the main currents in the Atlantic and the Pacific oceans
(Fig. 16.2). Minimum temperature reaches -2 °C South of the Brazilian region
and Eastern South America. The highest temperature is reached in the tropical
520
Table 16.1 Climatic and trophic variables in the water column in the coastal biogeographical provinces of North, Central and South America and the
Canary Islands
Minimum Maximum Mean T ST0 ST50 MLD Photic Maximum Chl Minimum Chl Mean Chl Chl Productivity (g
T (°C)a T (°C)a (°C)a (°C)b (°C)b (m)b Depth (mg m-3)a (mg m-3)a (mg m-3)a (g m-1)b C m-2 d-1)b
(m)b
Caribean 5.3 31. 25.6 26.9 25.8 23.0 44.2 74.3 0.0 0.6 6.6 0.5
West Indian 2.8 31.2 27.3 26.9 25.8 23.0 44.2 74.3 0.0 0.3 6.6 0.5
Brazilian -2.0 30.5 24.6 27.6 26.7 14.5 38.7 74.3 0.0 0.4 55.8 1.9
EasternSouth -2.00 24.7 10.6 8.5 7.1 101.2 30.0 74.3 0.0 1.7 1.3
America
Canary 16.79 24.7 20.7 21.2 18.0 28.2 28.7 40.9 0.0 0.2 35.0 2.0
Islands
Easter Island 16.7 14.2 23.3 43.3 15.7 0.7
Chilean 16.7 14.2 23.3 43.3 15.7 0.7
Galápagos 27.6 24.2 22.5 53.3 4.3 0.3
Panamanian 27.3 21.0 10.2 39.2 11.3 0.9
Mexican 27.3 21.0 10.2 39.2 11.3 0.9
a
Mean values obtained from monthly satellite data for the period 1982–1991 for sea surface temperature and 2003–2010 for chlorophyll (http://
neo.sci.gsfc.nasa.gov)
b
Data from Longhurst et al. (1995), Longhurst (1998), compiled by Taeger and Lazarus (2010)
A. Pérez-Ruzafa et al.
Fig. 16.2 Main current systems affecting the American coasts. Orange arrows correspond to
warm currents, blue arrows correspond to cold currents. PNEC Pacific North Equatorial Current,
PNECC Pacific North Equatorial Countercurrent, PEUCC Pacific Equatorial Undercountercur-
rent, PSEC Pacific South Equatorial Current; CRCC Costa Rica Costal Current; CPC
Circumpolar current; ANEC Atlantic North Equatorial current; AECC Atlantic Equatorial
Countercurrent; ASEC Atlantic South Equatorial Current
areas of the Pacific and Caribbean, exceeding 31 °C. The thermal range between
summer and winter is greater than 25 °C in most areas and reaches 28.4 °C in the
West Indies. Canary Islands show less variability, with a range of only 7.8 °C
(Fig. 16.3a, b). In general, all the regions are productive. The Brazilian and Canary
Islands provinces appear to be among the most productive in the world (1.92 and
2.01 g C m-2day-1, respectively) (Taeger and Lazarus 2010). However, in the
case of the Canary Islands, this value is reached in the West African upwelling.
Productivity in the archipelago is less and maximum chlorophyll concentration
does not exceed 40.91 mg m-3. Minimum mean chlorophyll concentration is
reached in the Galápagos archipelago, with only 4.29 g m-3 (Fig. 16.3c).
The Caribbean, West Indian and Galápagos provinces are characterized by
warm waters, with the highest photic depth and with a high mixed layer depth and
low chlorophyll concentration and low productivity. The Panamanian and Mexican
Fig. 16.3 Mean annual minimum (a) and maximum (b) sea surface temperature (°C) for the
period 1982–1992 obtained from monthly satellite images. c Mean annual concentration of
chlorophyll a (mgm-3) for the period 2003–2010. Data source (http://neo.sci.gsfc.nasa.gov)
provinces share with the former the high temperature and photic depth, but have a
narrow mixed layer depth and higher productivity. The Chilean, Easter Island and
Fig. 16.4 Ordination of North, Central and South American biogeographical regions and
environmental variables in a biplot representation of the first axes of the Principal Component
Analysis (PCA) performed on the surface water column parameters according to Longhurst (see
Table 16.1). Key to abbreviations for environmental variables are in Table 16.1
Eastern South Atlantic provinces are characterized by cold and transparent waters,
with the highest mixed layer depth and low productivity (Fig. 16.4).
16.3 Diversity and Biogeographic Affinities
The echinoderm fauna of Latin America and the Canary Islands consist of 1,539
species, 82 species of Crinoidea, 392 species of Asteroidea, 521 species of
Ophiuroidea, 242 species of Echinoidea and 302 species of Holothuroidea. Six
hundred and twenty-seven species are found only along the Atlantic coasts of
Central and South America, 597 species are found only along the Pacific coasts
and oceanic islands and 19 species are found only in the Eastern Atlantic, in the
Canary Islands.
Species richness is highly variable among the different countries, from 20
species in Easter Island and 23 species along the Caribbean coast of Guatemala to
a maximum of 374 species in Cuba (Fig. 16.5a). However, the number of species
is highly dependent of the coast length of the countries (Fig. 16.6). The number of
species per 10 km of coast is 0.22 in Chile and 66.6 in Malpelo. Among bio-
geographical provinces (Fig. 16.7) the highest mean value is in the Panamanian
province with 11.88 (s.e. ± 7.4) species per 10 km. However, Permanova analyses
do not detect significant differences between provinces (P = 0.255).
The highest diversity (species richness (S), average taxonomic distinctness
(D+), total taxonomic distinctness (sD+), average phylogenetic diversity (Phi+)
and total phylogenetic diversity (sPhi+)) is in Caribbean and West Indian countries
(Atlantic coasts of Mexico, Guatemala and Cuba). This suggests they could be the
Fig. 16.5 Number of species (up) and percentage (down) of the five classes of echinoderms
along the Pacific and Atlantic coasts of North, Central and South American countries and the
Canary Islands. MEX Mexico; REV Revillagigedo Archipelago; GUA Guatemala; SAL El
Salvador; HON Honduras; NIC Nicaragua; CRC Costa Rica; COC Cocos Island; PAN Panama;
COL Colombia; MAL Malpelo Island; GAL Galápagos Archipelago; ECU Ecuador; PER Perú;
CHL Chile; PAS Eastern Islands. BEL Belize; VEN Venezuela; BRA Brazil; URG Uruguay, ARG
Argentina; MAV Malvinas Islands; CUB Cuba; HAI Haiti; RDO Dominican Republic, PRI Puerto
Rico; CAN Canary Islands; atl: Atlantic; car: Caribbean; pac: Pacific
origin of North Atlantic and Eastern Tropical Pacific echinoderm faunas. Cur-
rently, both areas (Caribbean and West Indies) are quite similar in temperature
regimes (maximum 31.21 °C in both areas, minimum 5.28 and 2.80 °C, mean
25.61 and 27.25 °C in the Caribbean and West Indian respectively), but differ in
chlorophyll concentration (0.64 and 0.28 mg m-3 respectively).
On the other hand, there are significant differences (Permanova P = 0.001) in
the percentage of the five classes of echinoderms between provinces (Fig. 16.5b).
Ophiuroidea is the dominant class in most regions, except in the Chilean province
that is dominated by Asteroidea, the Easter Island province that is dominated by
Echinoidea and the Mexican province that is dominated by Holothuroidea. Cri-
noidea is always the least abundant class. Their absence in the Easter Island region
and scarcity in the Eastern South America, Galápagos, Panamanian and Mexican
provinces should be noted. The absence of Crinoidea has been also emphasized in
other Atlantic archipelagos like Cabo Verde (Pérez-Ruzafa et al. 1999). It is also
Fig. 16.6 Relationship between the number of echinoderm species and shore length of the
Pacific and Atlantic coasts of the North, Central and South American countries and the Canary
Islands
Fig. 16.7 Standardized species richness per 10 km shore length in the biogeographical provinces
of North, Central and South American countries and in the Canary Islands. Error bars correspond
to s.e. of the mean
worthwhile to note the negative relationship that exists between the dominance of
holothurians in the faunas and the presence of crinoids (Fig. 16.8).
The echinoderm fauna of the different countries is consistent with their
belonging to one of the above mentioned regions and provinces. NMDS analyses,
performed with Bray-Curtis similarity calculated on species presence-absence
(Bray and Curtis 1957), spatially distributed the countries according to their
geographic location and importance to a given province (Fig. 16.9). The plot
shows two well-established clusters. One includes the Panamanian, Galápagos and
Fig. 16.8 Negative relationship between the percentage of species of crinoids and holothurians
of the North, Central and South American countries and in the Canary Islands
Fig. 16.9 Multivariate analysis nMDS plot of distance-related echinoderm species assemblage
structure from Latinamerica based on species composition data and Bray-Curtis similarity
measure. The faunal similarity between biogeographical provinces is represented by the relative
distance. MEX Mexico; REV Revillagigedo Archipelago; GUA Guatemala; SAL El Salvador;
HON Honduras; NIC Nicaragua; CRC Costa Rica; COC Cocos Island; PAN Panama; COL
Colombia; MAL Malpelo Island; GAL Galápagos Archipelago; ECU Ecuador; PER Perú; CHL
Chile; PAS Eastern Islands. BEL Belize; VEN Venezuela; BRA Brazil; URG Uruguay, ARG
Chilean provinces to the tip of the Eastern South American region. Another cluster
well separated region includes the Caribbean, West Indian, Lusitanica and Bra-
zilian provinces. Earlier studies (Maluf 1988a, 1991) and results presented here
suggest that the echinoderm faunas of the Panamic, Galápagos and the Chilean
provinces are biogeographically related. There is a low level of association among
echinoderm species from the Chilean fauna and Eastern South American
Fig. 16.10 a Ordination of countries in the representation of the first axes of the Detrended
Correspondence analysis (DCA) performed on the echinoderm species matrix. b Ordination of
species in the representation of the first axes of the Detrended Correspondence analysis (DCA)
performed on the echinoderm species matrix. Only species with a weight [40 % on the axis are
represented. MEX Mexico; REV Revillagigedo Archipelago; GUA Guatemala; SAL El Salvador;
HON Honduras; NIC Nicaragua; CRC Costa Rica; COC Cocos Island; PAN Panama; COL
Colombia; MAL Malpelo Island; GAL Galápagos Archipelago; ECU Ecuador; PER Perú; CHL
Chile; PAS Eastern Islands. BEL Belize; VEN Venezuela; BRA Brazil; URG Uruguay, ARG
biogeographic region. Chile is closer to Argentina and the Malvines than to Peru.
In fact, there are two biogeographical provinces, the Peru-Chilean and the South
America or Magellan.
The first two axis of the Detrended Correspondence Analyses (DCA), also
performed on country species composition, also show a geographical ordination,
Fig. 16.11 Composition of the echinoderm faunal assemblages of the biogeographical provinces
of North, Central and South America and the Canary Islands according to the thermal distribution
range of the species inventoried. a Caribbean, b Brazilian, c West Indian, d Canary Is.
(Lusitania), e Panamanian, f Galápagos, g Chilean, h Easter Island
Fig. 16.12 Number of echinoderm species shared by two countries as a function of the distance
between them. Distances have been calculated using the geographical coordinates at the midpoint
of their shorelines
Fig. 16.13 Ordination of

samples (biogeographical
areas) and environmental
variables (a) and families
(b) in the representations of
the first axes of the canonical
correspondence analysis
(CCA) performed on the
matrix containing the number
of species representing the
echinoderm orders and
families in the different
Atlantic biogeographical
provinces
Fig. 16.14 Ordination of samples (biogeographical areas) and environmental variables (a) and
species (b) in the representations of the first axes of the canonical correspondence analysis (CCA)
of Atlantic echinoderm species matrix using environmental data as explanatory variables. Key of
abbreviations for environmental variables and species are in Table 16.1
with North Atlantic locations on the second axis, Pacific countries on the positive
part of the first axis, and Southern Atlantic, and Southern Pacific countries as a
transition between the two main groups. The first axis, which determines the
separation of Atlantic and Pacific faunas and the gradient throughout the Magellan
region explain 29.6 % of total variance. The second axis explains only an addi-
tional 3.8 % (Fig. 16.10a).
The species represented in Fig. 16.10b are those that have a weight on the axis
higher than 40 %. The positive part of axis 1 is represented by species exclusive to
the Pacific and with a wide distribution in this ocean, like the sea urchins Tox-
opneustes roseus, Diadema mexicanum Echinometra vanbrunti, Eucidaris thou-
arsii, Tripneustes depressus, the sea stars Nidorellia armata, Ophidiaster
alexandri, Phataria unifascialis, the ophiuroids Ophiocoma aethiops and the ho-
lothuroids Holothuria (Halodeima) kefersteini, Holothuria (Platyperona) difficilis,
Isostichopus fuscus.
The left extreme of axis 1 is characterized by exclusively Atlantic species. In
the lower part of the axis 2 are widely distributed species and exclusively from the
western Atlantic coasts, like the crinoids Comactinia meridionalis meridionalis,
Davidaster discoideus, the ophiuroids Ophionereis reticulata, the sea star Luidia
clathrata, the sea urchin Echinometra viridis or the holothuroids Holothuria
(Halodeima) mexicana, Actinopyga agassizii.
Between both extremes are ubiquitous species, widely distributed in both
oceans, like the ophiuroids Ophiactis savignyi and Amphipholis squamata, the
Fig. 16.15 Ordination of samples (biogeographical areas) and environmental variables (a) and
species (b) in the representations of the first axes of the canonical correspondence analysis (CCA)
of Atlantic and Pacific echinoderm species matrix using environmental data as explanatory
variables. Key of abbreviations for environmental variables and species are in Table 16.1
holothuroids Holothuria (Thymiosycia) arenicola and Holothuria (Thymiosycia)

impatiens, and the sea star Linckia guildingi.
Along the axis 2 are anfiatlantic species, shared by the American coasts and the
Canary Islands, like the ophiuroids Astrophyton muricatum, Ophiocoma pumila,
Fig. 16.16 a Percentage of echinoderm species per class by bathymetric range in the Pacific
taxonomic list and b percentage of species present in one to six bathymetric intervals. N = 620
species
the sea stars Oreaster reticulatus, Nymphaster arenatus, the sea urchins Diadema
antillarum antillarum, Brissus unicolor and the holothuroid Euapta lappa.
Cosmopolitan species are an important component in all the biogeographical
regions (Fig. 16.11). Except the Chilean and the Easter Island provinces, which are
dominated by cold temperate species. The Central and South American regions are
dominated by tropical species and to a lesser extent, temperate species with a wide
distribution range. In the Canary Islands, the warm temperate component is also
important.
Similarity between country faunas depends on climatic and trophic conditions
and on geographical proximity. The number of shared species is highly dependent
on the distance separating two countries (Fig. 16.12). The diminishing proportion
of shared species between two areas with increasing geographical distance is an
obvious feature of natural systems (Hengeveld 1990; Huston 1994; Rosenzweig
1995; Brown and Lomolino 1998; Hubbell 2001).
Fig. 16.17 a Percentage of echinoderm species per class by bathymetric range in the Caribbean
and Atlantic taxonomic list and b percentage of species present in one to six bathymetric
intervals. N = 836 species
There are more islands in the Atlantic region (i.e. the Lesser Antilles). Geo-
graphical distance between islands might be an important source of similarity in
terms of species richness and taxonomic composition. Two islands may share the
same number of species not because they are similar in area and/or in diversity
because they are geographically close (Rosenzweig 1995), which allows individ-
uals to move easily from one island to the other.
Canonical Correspondence analyses performed on the number of species rep-
resenting the echinoderm orders and families in the different biogeographical
provinces (Fig. 16.13) are consistent with the previous non-canonical analyses on
species presence-absence data and climatic conditions. Figure 16.13 shows the
results obtained for Atlantic provinces. The environmental variables used were
mean seasonal and mean, minimum, maximum and range of variation annual
values obtained from monthly satellite data for sea surface temperature
(1982–1991) and for chlorophyll (2003–2010). Productivity values were obtained
structure from a the Pacific taxonomic list and b the Caribbean and Atlantic taxonomic list, based
on species presence/absence matrix by bathymetric interval using Bray-Curtis similarity measure
from Longhurst et al. (1995), Longhurst (1998), compiled by Taeger and Lazarus
(2010) (Table 16.1).
The first two axes account for 46.6 and 30.3 %, respectively, of the total var-
iance of species-environment relation. The first axis represents a gradient from the
Caribbean and West Indies to the south. The positive extreme of the first axis is
associated with the Eastern South America region and highest mean chlorophyll
concentration in all seasons and the lowest temperatures. It is represented by the
Fig. 16.19 a Percentage of echinoderm species per class by habitat type in the Pacific taxonomic
list and b percentage of species presents in one to seven habitat types. N = 473 species
families Heliasteridae, Stichasteridae, Ganeriidae, Myxasteridae and Prenasteri-

dae. The Myxasteridae is exclusive to the Eastern South America region and the
Prenasteridae is shared with the Chilean province.
In the negative part of this axis, associated with highest temperatures and low
chlorophyll concentration, are families with a wide distribution in the Atlantic but
absent in the Eastern South America region like Echinocyamidae. Some of them
like Brisingida, Ophidiasteridae, Saleniidae, Stichopodidae, Ophiodermatidae or
the class Crinoidea are even present along Pacific coasts. This indicates their
tropical origin and ancient links, prior to closure of the Isthmus of Panama. In
general, these families are represented by different species in each ocean. For
example, the sea star family Ophidiasteridae, is present in the area with 29 species,
14 in the Pacific, five in the Western Atlantic and nine in the Anfiatlantic. None is
shared between the Atlantic and Pacific coasts. Congeneric species are Ophidiaster
Fig. 16.20 a Percentage of echinoderm species per class by habitat type in the Caribbean and
Atlantic taxonomic list and b percentage of species presents in one to seven habitat types.
N = 630 species
alexandri, Ophidiaster bayeri, Tamaria floridae and Tamaria halperni in the

Western Atlantic, Ophidiaster agassizi, Ophidiaster ludwigi along the Pacific
coasts, and Tamaria obstipa and Tamaria stria. Ophidiaster guildingii and Oph-
idiaster ophidianus in the Anfiatlantic. Narcissia canariensis and N. trigonaria are
Anfiatlantic, Narcissia gracilis and the subspecies N. gracilis malpeloensis occur
in the Pacific.
The second axis represents a Western–Eastern gradient. Pentametrocrinidae,
Elpidiidae, Hyocrinidae and Spatangidae characterize the positive part of this axis.
Pentametrocrinidae is present in the Canary Islands and is not shared with south
American regions, Elpidiidae and Hyocrinidae are shared by the Canary Islands
and all the Pacific regions except Easter Island, but are absent in the South
American coasts. In the same way, Spatangidae is a family shared by the Canary
Islands and the Pacific Mexican coast but is absent from the other regions.
structure from the a the Pacific taxonomic list and b the Caribbean and Atlantic taxonomic list,
based on species presence/absence matrix by habitat category using Bray-Curtis similarity
measure
The negative part of this axis is represented by the families Solasteridae,

Echinometridae, Sclerodactylidae and the order Cassiduloida. All of them are
present in the Caribbean, West Indian and Brazilian provinces, and even in the
warm Pacific coasts. But they are not present in the Canary Islands or the Eastern
South America region. It indicates a common tropical origin, after the separation
of western and eastern Atlantic coasts but prior to closure of the Isthmus of
Panama.
The same analyses performed on species presence-absence data can be seen in
Fig. 16.14 for the Atlantic regions and Fig. 16.15 for Pacific and Atlantic Regions.
The figure includes only species fitting more than 50 % of the axis and weighting
more than 50 %.
As previously mentioned, the affinities between faunas are a consequence of the
combination of climatic and trophic factors, connectivity as a function of distance,
current patterns and historical processes. The partition of the variance to see the
relative influence of environmental variables and spatial organization (latitude and
longitude coordinates) was done following the methodology proposed by Borcard
et al. (1992). It shows that the covariation of space and environment reach 62.2 %
in the case of Atlantic provinces. The environmental variables alone are respon-
sible for 37.8 % of the variance. The covariation of space and environment of the
Atlantic and Pacific regions analyzed together account for only 11.4 % of the
variability while environmental variables explain 58.6 % of the variability. There
is 14.5 % variation that is purely spatial and an additional 15.5 % is of unex-
plained variation and stochastic fluctuations or due to historical processes linked to
the isolation of the faunas of the Caribbean and tropical Pacific after the closure of
the isthmus of Panama.
Furthermore, different environmental factors would be responsible for faunal
composition and species distribution at different spatial scales (Barry and Dayton
1991; Levin 1992; Pérez-Ruzafa et al. 2003; Entrambasaguas et al. 2008). Water
temperature is considered the most important influence on the global distribution
of marine animals and minimum temperature is usually the factor that determines
faunal similarities at small geographical scales (Steele 1983; Pérez-Ruzafa and
López-Ibor 1988; Pérez-Ruzafa et al. 2003). Factors such as predation (Tegner and
Dayton 1981; Sala 1997), settlement and recruitment (Young and Chia 1982; Ebert
1983; Hereu et al. 2004; Hernández et al. 2010), availability of trophic resources
(Menge 1992), disease epidemics (Hagen 1999; Dumont et al. 2004), or com-
petitive interactions (Hagen and Mann 1992), physical factors like substrate nat-
ure, bottom complexity, depth, wave exposition, etc. (Drouin et al. 1985; Tyler
et al. 2000; Entrambasaguas et al. 2008) or harvesting (Pfister and Bradbury 1996;
Hasan 2005) are of major importance in explaining small scale species distribution
and abundance.
16.4 Bathymetrical and Substrate Distribution
In the Pacific taxonomic list (see appendix) there were 620 species of echinoderms
with bathymetric information (Fig. 16.16) that represent 86 % of the species of the
list, while in the Caribbean and Atlantic taxonomic list (see appendix) there were
836 species, that represent 92 % of the species on the list (Fig. 16.17). In the
Pacific most species were reported at depths between 20 and 200 m (336 species),
30 % in the class Asteroidea (Fig. 16.16a). This class was also the predominant at
depths between 0 and 20 m (28 %). The class Ophiuroidea was the predominant at
depths between 200–1,000 m (35 %) and 1,000–2,000 m (42 %). The class Ho-
lothuroidea was predominant at depths between 2,000–4,000 m (36 %) and
4,000–6,000 m (55 %). Most species on the Pacific list were found only in one or
two bathymetric intervals (each 36 %) while just 2 and 1 % were found in five and
six bathymetric intervals respectively (Fig. 16.16b). In the Caribbean and Atlantic
list most species were also at depths between 20 and 200 m (568 species)
(Fig. 16.17a), 35 % in the class Ophiuroidea. This class also predominated at
depths between 0–20 m (36 %), 200–1,000 m (37 %) and 1,000–2,000 m (35 %).
The class Asteroidea was predominant at depths between 2,000–4,000 m (38 %)
and 4,000–6,000 m (41 %). Most of the species on the Caribbean and Atlantic list
were found only in two or one bathymetric intervals (41 and 25 %, respectively)
while only 1 and 0.2 % were found in five and six bathymetric intervals respec-
tively (Fig. 16.17b).
According to the depth categories in the Pacific and the Caribbean-Atlantic, the
similarity analysis showed four groups with 40 % resemblance (Fig. 16.18a, b).
The first group is composed by the depths from 0 to 20 m, 20 to 200 m and 200 to
1,000 m. These groups possesses the highest number of species (314, 336, 260
species respectively in the Pacific and 365, 568, 551 species in the Caribbean-
Atlantic), Margalef species richness (54.4, 57.5, 46.5 respectively in the Pacific
and 61.5, 89.2, 87.1 respectively in the Caribbean-Atlantic) and Shannon diversity
(5.7, 5.8, 5.5 respectively in the Pacific and 5.8, 6.3, 6.3 respectively in the
Caribbean-Atlantic). The second group is composed by depths from 1,000 to
2,000 m and 2,000 to 4,000 m, that possesses moderate values of number of
species (164 and 134 species in the Pacific and 207 and 114 in the Caribbean-
Atlantic), Margalef species richness (31.9 and 27.1 in the Pacific and 38.6 and 23.8
in the Caribbean-Atlantic) and Shannon diversity (5.1 and 4.8 in the Pacific and
5.3 and 4.7 in the Caribbean-Atlantic). The other two groups are composed by only
one bathymetric interval (4,000–6,000 m and 6,000–10,000 m) and possesses the
lowest values of all indices; number of species (55 and 8 species in the Pacific and
32 and 2 in the Caribbean-Atlantic), Margalef species richness (13.4 and 3.3 in the
Pacific and 8.9 and 1.4 in the Caribbean-Atlantic) and Shannon diversity (4.0 and
2.0 in the Pacific and 3.4 and 0.6 in the Caribbean-Atlantic). Of the first two
groups, the Caribbean-Atlantic always had higher values than the Pacific in all the
indices. But in deeper waters (groups 3 and 4), the Pacific was richer than the
Caribbean-Atlantic.
There were 473 species of echinoderms (66 % of the total) in the Pacific
taxonomic list (see appendix) with habitat or substrate information (Fig. 16.19) In
the Caribbean and Atlantic taxonomic list (appendix) there were 630 species
(70 % of the total) with habitat or substrate information (Fig. 16.20). Most of the
species in the Pacific were on rocky bottoms. Of the 294 species, 30 % are in the
class Asteroidea (Fig. 16.19a). This class was also the predominant on rocky and
rubble bottom (34 and 53 %, respectively). On coral reefs and in kelp forest, the
classes Ophiuroidea (33 and 36 %, respectively) and Holothuroidea (33 and 36 %,
respectively) were predominantd. Holothuroidea were predominant on muddy
bottoms (30 %). On sandy bottoms, Asteroidea (26 %) and Ophiuroidea (26 %)
were predominant. Most of the species in the Pacific list were found in only one
habitat (each 47 %) while only 0.2 % were found in five and seven habitats
(Fig. 16.19b). In the Caribbean and Atlantic list, most of the species were on sandy
bottoms (357 species) (Fig. 16.20a), 33 % in the class Asteroidea. This class also
was predominant on rubble (33 %) and muddy (30 %) bottoms. The class
Ophiuroidea was predominant on coral reefs (50 %), mangroves (62 %), seagrass
(43 %) and rocky bottoms (30 %). Most of the species in the Caribbean and
Atlantic lists were found in only one habitat (33 %) while only 3 % and 1 % were
found in six and seven habitats respectively (Fig. 16.20b).
There are three groups according to habitat categories in the Pacific
(Fig. 16.21a). The first group is composed of coral reefs, muddy, rocky and sandy
bottoms. These habitats possess the highest values of total species richness
(94–294), Margalef species richness (20.3–51.4) and Shannon Diversity (5.3–5.9).
The second group is composed by sea grass, mangroves and rubble bottom. These
habitats possess low values of total species richness (2–15), Margalef species
richness (1.4–5.2) and Shannon Diversity (0.7–2.7). The third group is composed
by the kelp forest that possesses low values of the total species richness (11),
Margalef species richness (4.2) and Shannon Diversity (2.3). In the Caribbean-
Atlantic, the nMDS indicates the presence of four groups (Fig. 16.21b). The first
one is composed of sandy and muddy bottoms that possess the highest values of
total species richness (357–333 species respectively), Margalef species richness
(60.4–57.1 respectively) and Shannon Diversity (5.9–5.8). The second group is
composed of rocky bottoms and coral reefs that possess high to moderate values of
total species richness (298–199 species respectively), Margalef species richness
(52.1–37.4 respectively) and Shannon Diversity (5.7–5.3 respectively). The third
group is composed of seagrass and rubble bottoms with moderate values of total
species richness (94–143 species respectively), Margalef species richness
(20.5–28.6 respectively) and Shannon Diversity (4.5–4.9 respectively). The last
group is composed of mangroves that possess the lowest values of total species
richness (29), Margalef species richness (8.3) and Shannon Diversity (3.4).
Acknowledgments We are grateful with all the authors that participated on this contribution.
We appreciate the help and extensive work of the students from the Laboratorio de Ecología y
Taxonomía de Equinodermos, Colección Nacional de Equinodermos Dra. María Elena Caso,
Instituto de Ciencias del Mar y Limnología from the Universidad Autónoma Nacional de México.
JJA wants to acknowledge Monica Chavez for preparing all the figures from this chapter and
CONACYT and CONICIT for all the economic support during the preparation and analysis of
this review.
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Appendix
Solís-Marín FA, JJ Alvarado, M Abreu-Pérez, O Aguilera, J Alió, JJ Bacallado-

Aránega, E Barraza, M Benavides-Serrato, F Benítez-Villalobos, L Betancourt-
Fernández, M Borges, M Brandt, MI Brogger, GH Borrero-Pérez, BE Buitrón-
Sánchez, LS Campos, J Cantera, S Clemente, M Cohen-Renjifo, S Coppard,
LV Costa-Lotufo, R del Valle-García, Y Díaz, ME Díaz de Vivar, JP Díaz-
Martínez, A Durán-González, L Epherra, M Escolar, V Francisco, CA Freire,
JE García-Arrarás, DG Gil, P Guarderas, VF Hadel, A Hearn, JC Hernández,
EA Hernández-Delgado, A Herrera-Moreno, MD Herrero-Pérezrul, Y Hooker,
MBI Honey-Escandón, C Lodeiros, M Luzuriaga, CLC Manso, A Martín,
MI Martinez, S Martínez, L Moro-Abad, E Mutschke, JC Navarro, R Neira,
N Noriega, JS Palleiro-Nayar, AF Pérez, A Pérez-Ruzafa, E Prieto-Rios, J Reyes,
R Rodríguez, T Rubilar, T Sancho-Mejía, C Sangil, JRMC Silva,
JI Sonnenholzner, CR Ventura, A Tablado, Y Tavares, CG Tiago, F Tuya,
SM Williams

in Latin America, DOI: 10.1007/978-3-642-20051-9,
Table A.1 Taxonomic list of the Echinoderms of the Pacific coast of Latin America
544
Depth (m) Habitat MEX REV GUA SAL HON NIC CRC COC PAN COL MAL GAL ECU PER CHL PAS
Class Crinoidea
Order Comatulida
Superfamily Antedonacea
Family Antedonidae
Antedon bifida (Pennant, 1777) x
Anthometra adriani (Bell, 1908) 70–1674 x
Eumorphometra fraseri John, 1938 410–687 x
Fariometra parvula (Hartlaub, rb x x
1895)
Florometra magellanica (Bell, 20–1017 rb,sb x x x
1882)
Florometra mawsoni A. H. Clark, 38–770 x
1913
Florometra parvula (Hartlaub, 589–1969 rb x
1895)
Florometra serratissima (A. 12–3234 rb x x
H. Clark, 1907)
Florometra tanneri (Hartlaub, 1895) 104–207 rb x x
Isometra hordea John, 1938 17–490 x
Isometra graminea John, 1938 x
Isometra vivipara Mortensen, 1917 79–845 x
Promachocrinus kerguelensis 20–2100 x
Carpenter, 1880
Solanometra antarctica (Carpenter, 287–1759 x
1888)
Family Zenometridae
Psathyrometra bigradata (Hartlaub, 741–823 x x
1895)
(continued)
Appendix
Table A.1 (continued)
Superfamily Notocrinacea
Appendix
Family Notocrinidae
Notocrinus virilis Mortensen, 1917 x
Notocrinus mortenseni John, 1938 x
Superfamily Tropiometracea
Family Thalassometridae
Thalassometra agassizii (Hartlaub, 596–1429 rb x x x
1895)
Order Hyocrinida
Family Hyocrinidae
Calamocrinus diomedae A. 717–1431 x x
Agassiz, 1890
Hyocrinus foelli (Roux & Pawson, 3030 rb x x
1999)
Ptilocrinus antarcticus (Bather, 450–500 rb x
1908)
Class Asteroidea
Order Paxillosida
Family Astropectinidae
Astropecten armatus Gray, 1840 0–160 mb,rb,sb x x x x x x x x
Astropecten armatus erinaceus 11–60 rb,sb x
Gray, 1840
Astropecten benthophilus Ludwig, 1408 x
1905
Astropecten brasiliensis 6–99 mb,sb x x
peruvianus Verrill, 1870
Astropecten exiguus Ludwig 1905 232–2136 x x
Astropecten fragilis Verrill, 1870 1–5 sb x x x
(continued)
545
546
Astropecten regalis Gray, 1840 0–204 mb,rb,sb x x x x
Astropecten sulcatus Ludwig, 95–121 x x x
1905
Astropecten verrilli de Loriol, 2–488 mb,rb,sb x x x x
1899
Astropecten ornatissimus Fisher, 278–286 rb,sb x
1906
Bathybiaster loripes Sladen, 1889 80–500 sb x
Dipsacaster eximius Fisher, 1905 377–971 mb,sb x x
Dytaster gilberti Fisher, 1905 1573–4335 mb,sb x x x x x
Leptychaster inermis (Ludwig, 732–1593 x x x x
1905)
Mimastrella cognata (Sladen, 10–2424 mb,rb x
1889)
Psilaster charcoti (Koehler, 1906) 20–3248 sb x
Psilaster pectinatus (Fisher, 1905) 1866 x
Psilaster sladeni Ludwig, 1905 1485–1618 x
Persephonaster armiger Ludwig, x
1905
Tethyaster canaliculatus (A. 23–300 rb x x x x
H. Clark, 1916)
Thrissacanthias penicillatus 55–1503 sb x x
(Fisher, 1905)
Family Goniopectinidae
Ctenodiscus procurator Sladen, 50–1050 mb x
1889
Ctenodiscus crispatus (Retzius, 10–1946 mb x x x
1805)
Luidia (Platasterias) latiradiata 10–12 mb,sb x
(Gray, 1871)
(continued)
Appendix
Luidia armata Ludwig, 1905 33–126 sb x x x
Appendix
Luidia asthenosoma Fisher, 1906 x

Luidia bellonae Lütken, 1865 4–55 rb,sb x x x
Luidia columbia (Gray, 1840) 0–220 mb,sb x x x x
Luidia ferruginea Ludwig, 1905 280 mb x x
Luidia foliolata (Grube, 1866) 3–55 mb,sb x x x x x
Luidia latiradiata (Gray, 1871) 1.5–12 mb,sb x x x x x
Luidia magellanica Leipoldt, 0–40 rb,sb x x
1895
Luidia phragma H. L. Clark, 1–386 sb x x x
1910
Luidia porteri A. H. Clark, 1917 110 x
Luidia superba A. H. Clark, 1917 3–250 mb,sb,rub x x x x x x
Luidia tessellata Lütken, 1859 18 sb x x x x
Family Porcellanasteridae
Eremicaster crassus (Sladen, 1570–6330 rb x x x x x x
1883)
Eremicaster pacificus (Ludwig, 1463–5780 mb,sb x x x x x x x
1905)
Eremicaster vicinus Ludwig, 5204–7200 mb x
1907
Porcellanaster ceruleus Wyville- 1158–6035 mb x x x
Thomson, 1877
Styracaster paucispinus Ludwig, 4240–4337 mb x
1907
Thoracaster cylindratus Sladen, 2600–5303 mb x x
1883
Family Pseudarchasteridae
Pseudarchaster discus Sladen, 117–400 mb,rb,sb x
1889
Pseudarchaster pectinifer 1180–3575 sb,rub,mb x x x x
Ludwig, 1905
547
(continued)
548
Pseudarchaster pulcher Ludwig, 700–1620 sb,rub x x
1905
Pseudarchaster pusillus Fisher, 98–800 sb,rub x
1905
Pseudarchaster verrilli Ludwig, 999 x
1905
Order Notomyotida
Family Benthopectinidae
Benthopecten acanthonotus Fisher, 1800–1936 rb x
1905
Benthopecten cognatus (Ludwig, 3058 x
1905)
Benthopecten pectinifer (Ludwig, 1485–2323 rb x x x
1905)
Benthopecten spinuliger (Ludwig, 1618–2323 x x
1905)
Calyptraster tenuissimus 0–732 sb x
Bernasconi, 1966
Cheiraster (Luidiaster) planeta 350–500 sb x
(Sladen, 1889)
Cheiraster (Luidiaster) californicus 488–512 mb x
Ziesenhenne, 1942
Nearchaster aciculosus (Fisher, 466–1903 mb,sb x x
1910)
Pectinaster agassizi Ludwig, 1905 790–2323 mb,rb x x x x x x
Order Valvatida
Family Acanthasteridae
Acanthaster planci (Linnaeus, 1758) 0–30 cr,rb x x x x x x x x
Family Asterinidae
Asterina fimbriata Perrier, 1875 0–300 mb,rb x
Meridiastra modesta (Verrill, 1870) 213 rb,sb x x x x x x
Patiria chilensis (Lütken, 1859) 2–40 rb,kf x x
Appendix
(continued)
Patiria miniata Verrill, 1913 0–300 cr,rb x x
Appendix
Patiriella calcarata (Perrier, 0–20 rb x x

1875)
Patiriella exigua (Lamarck, 0–3 x
1816)
Family Asterodiscididae
Amphiaster insignis Verrill, 0–128 cr,mb,rb,sb x x x x x x
1868
Paulia horrida Gray, 1840 33–121 rb x x x x x
Family Asteropseidae
Asteropsis carinifera (Lamarck, 3–60 cr,rb x x x x x x
1816)
Dermasterias imbricata (Grube, 2–260 rb x
1857)
Family Ganeriidae
Cycethra verrucosa (Philippi, 0–270 rb,sb x x
1857)
Ganeria falklandica Gray, 1847 0–145 rb,sb x
Ganeria hahni Perrier, 1891 0–135 rb,sb x
Family Goniasteridae
Ceramaster grenadensis 75–898 x x
patagonicus (Sladen, 1889)
Ceramaster patagonicus 10–898 rb,mb,sb x x x
(Sladen, 1889)
Ceramaster leptoceramus 382–1248 mb,rb x x
(Fisher, 1905)
Cryptopeltaster lepidonotus 188–1244 sb x x
Fisher, 1905
Hippasteria phrygiana 0–400 rb x
(Parelius, 1768)
Hippasteria falklandica Fisher, 225–1148 rb x
1940
549
(continued)
550
Hippasteria spinosa Verrill, 1909 50–512 rb x
Litonotaster tumidus H. L. Clark, 4066–5196 mb x
1920
Mediaster elegans Ludwig, 1905 1789 x x x
Mediaster transfuga Ludwig, 1905 840–900 sb x x
Mediaster tenellus Fisher, 1905 580–1192 sb x
Nymphaster diomedeae Ludwig, 702–1810 mb,sb x x x x x
1905
Pillsburiaster ernesti (Ludwig, 2149 x
1905)
Family Mithrodiidae
Mithrodia bradleyi Verrill, 1870 0–14 cr,rb x x x x x x x x x x
Family Odontasteridae
Acodontaster elongatus elongatus 8–400 rb,sb x
(Sladen, 1889)
Acodontaster elongatus granuliferus 74–841 rb,sb x
(Koehler, 1912)
Diplodontias singularis (Müller & 0–84 rb,sb x x
Troschel, 1843)
Odontaster meridionalis (E.A. 0–646 rb,sb x
Smith, 1876)
Odontaster penicillatus (Philippi, 6–400 rb,sb x
1870)
Family Ophidiasteridae
Leiaster coriaceus Peters, 1852 x
Leiaster glaber Peters, 1852 2–15 cr x
Leiaster teres (Verrill, 1871) 1–57 cr,rb x x x x x
Linckia columbiae Gray, 1840 0–100 cr,rb x x x x x
(continued)
Appendix
Linckia guildingi Gray, 1840 rb x x x
Appendix
Linckia multifora (Lamarck, x

1816)
Narcissia gracilis A.H. Clark, 56–91 rb x x x
1916
Narcissia gracilis malpeloensis 20–59 cr, rb x x
Downey, 1975
Ophidiaster agassizi Perrier, 1–75 rb x
1881
Ophidiaster ludwigi de Loriol, x x
1900
Pharia pyramidata (Gray, 1840) 0–130 cr,rb,sb x x x x x x x x x x
Phataria mionactis Ziesenhenne, x
1942
Phataria unifascialis (Gray, 0–50 cr,rb,sb x x x x x x x x x x x x
1840)
Tamaria obstipa Ziesenhenne, 67–79 x x
1942
Tamaria stria Downey, 1975 15–49 rb x x
Family Poraniidae
Porania (Porania) antarctica 0–900 mb,rb,sb x
E.A. Smith, 1876
Poraniopsis echinaster Perrier, 5–450 mb,rb,sb x
1891
Poraniopsis inflatus (Fisher, 48–1094 x x
1906)
Family Oreasteridae
Nidorellia armata (Gray, 1840) 0–183 cr,rb,rub x x x x x x x x x x x
Pentaceraster cumingi (Gray, 2–92 cr,rb,sb x x x x x x x x
1840)
Family Solasteridae
Lophaster furcilliger Fisher, 1905 86–4200 mb,rb,rub x x x x
551
Lophaster stellans Sladen, 1889 15–450 rb,sb x

(continued)
552
Depth Habitat MEX REV GUA SAL HON NIC CRC COC PAN COL MAL GAL ECU PER CHL PAS
(m)
Paralophaster antarcticus Koehler, 88–750 sb x
1912
Solaster regularis Sladen, 1889 5–800 rb,sb,mb x
Order Velatida
Family Korethrasteridae
Peribolaster folliculatus Sladen, 10–460 rb x
1889
Family Pterasteridae
Diplopteraster semireticulatus 300– rb,sb x
(Sladen, 1882) 1500
Diplopteraster verrucosus (Sladen, 0–470 sb x
1882)
Pteraster cf. diaphanus (Ludwig, 1410 x x
1905)
Pteraster gibber (Sladen, 1882) 7–460 rb x
Pteraster affinis Smith, 1876 80–130 rb x
Pteraster affinis lebruni Perrier, 1891 74–341 sb x
Hymenaster cremnodes H. L. Clark, 4335 mb x
1920
Hymenaster gracilis Ludwig, 1905 2418– x x x x
3241
Hymenaster pellucidus Thomson, 13–3240 mb x
1873
Hymenaster platyacanthus Ludwig, 2487– x x x
1905 2877
Hymenaster quadrispinosus Fisher, 778– mb x x x
1905 3240
Hymenaster trias H. L. Clark, 1920 5203 mb x
Hymenaster violaceus Ludwig, 1905 x
(continued)
Appendix
(m)
Appendix
Order Spinulosida
Family Echinasteridae
Echinaster (Echinaster) 18–28 rb x
parvispinus A. H. Clark,
1916
Echinaster (Othilia) 3–24 sb x x
aculeata (Gray, 1840)
Echinaster (Othilia) 1–55 cr,mb,rb x
spinulosus Verrill, 1869
Echinaster (Othilia) 0–18 rb x
tenuispinus Verrill, 1871
Echinaster cribella x
Lütken, 1871
Echinaster cylindricus x
Meissner, 1892
Echinaster panamensis x
Leipoldt, 1895
Henricia aspera Fisher, 487–570 rb x
1906
Henricia asthenactis 91–1250 rb x
Fisher, 1910
Henricia clarki Fisher, 226– rb x x
1910 2001
Henricia gracilis 267– rb x
(Ludwing, 1905) 1244
Henricia leviuscula 0–228 rb x
(Stimpson, 1857)
Henricia nana (Ludwig, 57–200 rb x x
1905)
(continued)
553
554
(m)
Henricia obesa (Sladen, 22–940 rb x x x
1889)
Henricia seminudus (A. 666 rb x x
H. Clark, 1916)
Henricia studeri Perrier, 0–430 rb,sb,rub x x
1891
Order Forcipulatida
Family Asteriidae
Anasterias antarctica 0–350 rb x
(Lütken, 1857)
Anasterias pedicellaris 0–120 sb x
(Köehler, 1923)
Anasterias spirabilis 34–54 x
(Bell, 1881)
Anasterias varium 0–350 rb x
(Philippi, 1870)
Astrometis sertulifera 11–156 cr,rb x x x
(Xantus, 1860)
Astrostole platei 0–20 rb x
(Meissner, 1896)
Astrostole paschae (H. rb x
L. Clark, 1920)
Coronaster marchenus 52–84 cr,rb,rub x x x x
Ziesenhenne, 1942
Cosmasterias lurida 0–650 mb,rb,sb x
(Philippi, 1858)
Diplasterias brandti 0–500 mb,rb,sb x
(Bell, 1881)
Distolasterias robusta 3334 x x x
(Ludwig, 1905)
(continued)
Appendix
(m)
Appendix
Evasterias troscheli x
(Stimpson, 1862)
Leptasterias pusilla 37 rb x
Fisher, 1930
Meyenaster gelatinosus 0–22 rb x x x
(Meyen, 1834)
Pisaster brevispinus 0–102 rb x
(Stimpson, 1857)
Pisaster giganteus 0–374 rb x
(Stimpson, 1857)
Pisaster ochraceus 0–8 rb x x
(Brandt, 1835)
Rathbunaster rb x
californicus Fisher, 1906
Sclerasterias alexandri 61–384 x x x x x
(Ludwig, 1905)
Sclerasterias heteropaes 18–457 rb x x
Fisher, 1924
Family Coscinasteridae
Psalidaster mordax 80–600 sb x
Fisher, 1940
Family Heliasteridae
Heliaster canopus intertidal rb x
Perrier, 1875
Heliaster cumingii 0–14 rb x
(Gray, 1840)
Heliaster helianthus 0–20 rb,sb x x x
(Lamarck, 1816)
Heliaster kubiniji 0–20 rb x x x
Xantus, 1860
(continued)
555
556
(m)
Heliaster microbrachius 0–20 rb x x x
Xanthus, 1860
Heliaster polybrachius 0–20 rb x x x
H. L. Clark, 1907
Heliaster solaris A. rb x
H. Clark, 1920
Labidiaster radiosus 5–450 rb x
Lütken, 1871
Family
Pedicellasteridae
Ampheraster 0–520 rb,rub x x
hyperonchus (H.
L. Clark, 1913)
Hydrasterias improvisus 1618– x
(Ludwig, 1905) 2418
Tarsaster cocosanus 245 x x
(Ludwig, 1905)
Tarsaster galapagensis 704 x
(Ludwig, 1905)
Family Pycnopodiidae
Pycnopodia 0–455 rb x
helianthoides (Brandt,
1835)
Family Stichasteridae
Allostichaster capensis 0–100 rb x
(Perrier, 1875)
Stichaster striatus Müller 0–80 rb,sb,kf x x
& Troschel, 1840
Family Zoroasteridae
(continued)
Appendix
(m)
Appendix
Cnemidaster nudus 1366– mb,rb x x

(Ludwig, 1905) 2600
Cnemidaster wyvillii x
Sladen, 1889
Myxoderma longispinum 980– mb,rb x x x x
(Ludwig, 1905) 2418
Myxoderma 395–650 mb,rb x
platyacanthum (H.
L. Clark, 1913)
Myxoderma qawashqari 0–723 sb x x
(Moyana & Larrain,
1976)
Myxoderma sacculatum 1000– mb,rb x
(Fisher, 1905) 1546
Zoroaster magnificus 3056– mb,sb x x x
Ludwig, 1905 3667
Zoroaster ophiurus 695– mb,sb x
Fisher, 1905 2226
Order Brisingida
Family Brisingidae
Astrolirus panamensis 48–2418 x x x x x
(Ludwig, 1905)
Hymenodiscus 4064 mb x
monacantha H. L. Clark,
1920
Hymenodiscus tenella 2418 x
(Ludwig, 1905)
Family Freyellidae
Freyastera benthophila 4064– mb x
(Sladen, 1889) 4667
(continued)
557
558
(m)
Freyella insignis 3182– x x x x
Ludwig, 1905 3279
Freyella pacifica x
Ludwig, 1905
Freyella propinqua x
Ludwig, 1905
Freyellaster scalaris (A. x
H. Clark, 1916)
Class Ophiuroidea
Order Euryalida
Family Asteronychidae
Asteronyx excavata 266– sb x
Lütken & Mortensen, 1236
1899
Asteronyx loveni Müller 152– cr,mb,sb x x x
& Troschel, 1842 2663
Asteronyx longifissus 266– x
Doederlein, 1927 1800
Astrodia plana (Lütken 717– x x x x
& Mortensen, 1899) 3058
Family
Asteroschematidae
Asteroschema rubrum 350–731 x
(Lyman, 1882)
Asteroschema sublaeve 1271 x x
Lütken & Mortensen,
1899
Family
Gorgonocephalidae
(continued)
Appendix
(m)
Appendix
Astrocaneum spinosum 4–183 cr,mb,sb,rb x x x x

(Lyman, 1875)
Astrodendrum 717 x
galapagensis A.
H. Clark, 1916
Astrodictyum panamense 23559 cr,sb x x x x x x
(Verrill, 1867)
Astrotoma agassizii 2–1180 rb x
Lyman, 1875
Gorgonocephalus 4–900 rb x
chilensis (Philippi, 1858)
Gorgonocephalus 1271 x
diomedeae Lütken &
Mortensen, 1899
Order Ophiurida
Family Ophiomyxidae
Ophiomyxa panamensis 60–293 rb,sb x x x x
1899
Ophiomyxa vivipara 15–399 rb x
Studer, 1876
Ophiolycus nutrix 70–4538 sb x
(Mortensen, 1936)
Family Amphiuridae
Amphilepis nuda x
Tommasi, 1976
Amphilepis patens 384– mb x x x
Lyman, 1879 4087
Family Amphiuridae
(continued)
559
560
(m)
Amphicontus minutus 9–22 x x
Hill, 1940
Amphichondrius 280–384 mb, sb x x x
granulatus (Lütken &
Mortensen, 1899)
Amphichondrius laevis 4–15 cr, rb x x
Ziesenhenne, 1940
Amphiodia assimilis 3334 x x x
(Lütken & Mortensen,
1899)
Amphiodia grisea 54 x x x
(Ljungman, 1867)
Amphiodia occidentalis 0–367 mb, sb x x
(Lyman, 1860)
Amphiodia oerstedi 0.2–1 mb,rb x x x x
(Lütken, 1856)
Amphiodia platyspina 350 rb x x
Nielsen, 1932
Amphiodia sculptilis 4–15 rb x x
Ziesenhenne, 1940
Amphiodia tabogae 0–364 rb x x x x x
Nielsen, 1932
Amphiodia urtica 9–80 rb,sb x x
(Lyman, 1860)
Amphiodia violacea x x x
(Lütken, 1856)
Amphiodia vicina H. 64 mb x
L. Clark, 1940
Amphipholis elevata 0–73 sb x x
Nielsen, 1932
(continued)
Appendix
(m)
Appendix
Amphipholis granulata 280–384 x

1899)
Amphipholis laevidisca x x
H. L. Clark, 1909
Amphipholis pugetana 4–1620 sb x x x
(Lyman, 1860)
Amphipholis puntarenae 0–508 mb x
(Lütken, 1856)
Amphipholis squamata 0–1200 cr,m,mb,sb,sg,rub,rb x x x x x x
(Delle Chiaje, 1828)
Amphioplus 4–1408 mb,sb x
(Amphioplus)
strongyloplax (H.
L. Clark, 1911)
Amphioplus 1207 mb x
(Amphioplus)
coniortodes H. L. Clark,
1918
Amphioplus daleus 2690– rb x x
(Lyman, 1879) 3219
Amphioplus 15–73 mb x
philohelminthius
Ziesenhenne, 1940
Amphioplus magellanica 35–174 x
(Mortensen, 1936)
Amphioplus textilis x
(Koehler, 1907)
Amphiura anomala 300– x
Lyman, 1875 1500
(continued)
561
562
(m)
Amphiura arcystata H. 6–849 mb,rb,sb x x x x
L. Clark, 1911
Amphiura eugeniae 0–582 x
Ljungman, 1867
Amphiura (Amphiura) 44–3017 sb x x x x
diomedeae Lütken &
Mortensen, 1899
Amphiura (Amphiura) 0–300 x
magellanica Ljungman,
1867
Amphiura (Ophionema) x
hexacantha Nielsen,
1932
Amphiura calbuca 0–300 x
Mortensen, 1952
Amphiura carchara H. 1587 mb,rb,sb x
L. Clark, 1911
Amphiura gastracantha 1207 mb,sb x
1899
Amphiura gymnogastra 549– x x
1899
Amphiura gymnopora 333 x
1899
Amphiura notacantha 1236 mb,sb x
1899
Amphiura otteri 80 rb x
Ljungman, 1872
Appendix
(continued)
(m)
Appendix
Amphiura polyacantha 1271 x x

1899
Amphiura papillata 702 x
1899
Amphiura princeps 0–300 x
Koehler, 1907
Amphiura seminuda 9–4096 rb,sb x
1899
Amphiura serpentina 770– x x
Lütken & Mortensen 1865
1899
Amphiura verticillata x
Ljungman, 1867
Microphiopholis 0–82 mb,rb,sb x x x x x
geminata (Le Conte,
1851)
Microphiopholis 0–137 mb,sb x x x x x
platydisca (Nielsen,
1932)
Microphiopholis x x
puntarenae (Lütken,
1856)
Ophiocnida californica 6–302 mb,rb,sb x
Ziesenhenne, 1940
Ophiocnida hispida (Le 0–794 cr,rb x x
Conte, 1851)
(continued)
563
564
(m)
Ophiophragmus 0–40 rb,sb x x
chilensis (Müller &
Troschel, 1843)
Ophiophragmus x
disacanthus
Ziesenhenne, 1940
Ophiophragmus 0–134 sb x x x x x x
marginatus (Lütken,
1856)
Ophiophragmus x x x
ophiactoides
Ziesenhenne, 1940
Ophiophragmus 0–134 x x x
paucispinus Nielsen,
1932
Ophiophragmus stellatus 18–73 x
Ziesenhenne, 1940
Ophiophragmus 0–128 sb x x x x x
tabogensis Nielsen, 1932
Ophiostigma tenue 1–101 rb x x x
Lütken, 1856
Triplodia abdita (A. x
M. Clark, 1970)
Family Hemieuryalidae
Amphigyptis perplexa 0–143 mb,rb,sb x
Nielsen, 1932
Ophiochondrus stelliger 73–439 x
Lyman, 1879
Sigsbeia laevis 55–64 x
Ziesenhenne, 1940
(continued)
Appendix
(m)
Appendix
Sigsbeia lineata Lütken 43–183 rb x x x

& Mortensen, 1899
Family Ophiacanthidae
Subfamily
Ophiacanthinae
Ophiacantha antarctica 90–4004 sb x
Koehler, 1900
Ophiacantha bathybia H. 1993 rb x
L. Clark, 1911
Ophiacantha contigua 1062– x x x
1899
Ophiacantha cosmica 147– mb,rb,sb x x x x
Lyman, 1878 4840
Ophiacantha costata 733– mb,sb x x x
1899
Ophiacantha cyrena A. 717 x
H. Clark, 1916
Ophiacantha deruens x
Koehler, 1899
Ophiacantha diplasia H. 9–1408 cr,mb,sb x
L. Clark, 1911
Ophiacantha frigida 1667– x
Koehler, 1909 3914
Ophiacantha hirta 1244– mb,sb x
1899
Ophiacantha 702– x x
inconspicua Lütken & 1485
Mortensen, 1899
565
(continued)
566
(m)
Ophiacantha marsupialis x
Lyman, 1878
Ophiacantha 519– mb,sb x x
moniliformis Lütken & 1244
Mortensen, 1899
Ophiacantha pacifica 2877 x
1899
Ophiacantha paucispina 2690 x
1899
Ophiacantha pentacrinus 434–598 sb x
Lütken, 1869
Ophiocantha phragma 13–644 rb,sb x x x x x
Ziesenhenne, 1940
Ophiacantha 183–549 rb x
quadrispina H. L. Clark,
1917
Ophiacantha 115– mb,rb,sb x
rhachophora H. 1552
L. Clark, 1911
Ophiacantha rosea 32–1538 rb,sb x
Lyman, 1878
Ophiacantha savagica 1171– x
Tommasi, 1976 1180
Ophiacantha sentosa 2067– mb x x
Lyman, 1878 5203
Ophiacantha similis A. 717 x
H. Clark, 1916
Ophiacantha spinifera 999– x
1899
Appendix
(continued)
(m)
Appendix
Ophiacantha vivipara 0–1097 rb,kf x

Ljungman, 1870
Ophiolebes mortenseni 717 x
A. H. Clark, 1916
Ophiotreta valenciennesi 549 x
valenciennesi (Lyman,
1879)
Subfamily
Ophiohelinae
Ophiotholia spathifer x
(Lyman, 1879)
Subfamily
Ophioplinthacinae
Ophiomitrella chilensis 25–300 x
Mortensen, 1952
Ophiophthalmus 51–2600 mb,rb,sb x x
normani (Lyman, 1879)
Ophiothamnus dupla 1171– x
Tommasi, 1976 1180
Ophiurothamnus laevis 1008 x
1899)
Subfamily
Ophiotominae
Ophiotoma paucispina 1643– mb x x x x
(Lütken & Mortensen, 4082
1899)
Ophiolimna antarctica 88–2750 x
(Lyman, 1879)
(continued)
567
568
(m)
Family Ophiactidae
Hemipholis cordifera 18–34 cr x x
Bosc, 1802
Hemipholis gracilis 34 mb x x x x x x x
Verrill, 1867
Histampica duplicata 125– cr,mb,rb x x x x x
(Lyman, 1875) 2870
Ophiactis asperula 0–576 rb,kf x x
(Philippi, 1858)
Ophiactis kroeyeri 0–60 rb,sb,mb,rub x x x
Lütken, 1856
Ophiactis savignyi 0–518 cr,m,rb,sg,rub x x x x x x x x x x x
(Müller & Troschel,
1842)
Ophiactis simplex (Le 0–302 cr,rb x x x x x x x x x
Conte, 1851)
Ophiactis plana Lyman, x x x
1869
Ophiactis profundi 1008 x
1899
Ophiopholis bakeri 9–1006 cr,rb x
McClendon, 1909
Family
Ophiochitonidae
Ophiochiton carinatus 589– sb,rub x x
1899
Ophiochiton fastigatus 733 x
Lyman, 1878
(continued)
Appendix
(m)
Appendix
Family Ophiocomidae
Ophiocoma aethiops 0–30.5 cr,rb,sb x x x x x x x x x x x x x x
Lütken, 1859
Ophiocoma alexandri 0–70 cr,rb,sb x x x x x x x x x x x x x
Lyman, 1860
Ophiocoma dentata x
Müller & Troschel, 1842
Ophiocoma erinaceus 8 x
Müller & Troschel, 1842
Ophiocoma longispina x
H. L. Clak, 1917
Ophiocomella schmitti 5–40 cr,rb x x x
A. H. Clark, 1939
Ophiocomella sexradia cr,rb x x
(Duncan, 1887)
Ophiocomina nigra 236–549 sb x
(Abildgaard, in O.F.
Müller, 1789)
Ophiopsila californica 33–201 rb x
A. H. Clark, 1921
Family
Ophiodermatidae
Diopederma danianum 7–137 cr,mb,rb,sb x x x x x
(Verrill, 1867)
Ophiocrytus granulosus 0–79 rb,sg x x
Nielsen, 1932
Ophioderma appressa 0–20 cr,rb x
(Say, 1825)
Ophioderma elaps x
Lütken, 1856
(continued)
569
570
(m)
Ophioderma panamense 0–20 cr,rb,sb x x x x x x x x x x
Lütken, 1859
Ophioderma 183 x x
pentacantha H. L. Clark,
1917
Ophioderma teres 0–54 cr,rb,sb x x x x x x x x
(Lyman, 1860)
Ophioderma vansyoci 15.2–27 rb x
Hendler, 1996
Ophioderma variegatum 0–110 mb,rb,sb x x x x x x x
Lütken, 1856
Ophioderma 4–9 rb,sb x
sodipallaresi Caso, 1986
Ophiopaepale diplax 0–230 mb,sb x x x
(Nielsen, 1932)
Ophiuroconis bispinosa 4–143 sb x x x
Ziesenhenne, 1937
Family
Ophionereididae
Ophionereis 73–155 x x x
albomaculata E.A.
Smith, 1877
Ophionereis annulata 0–229 cr,rb,sb x x x x x x x x x
(Le Conte, 1851)
Ophionereis dubia x
1842)
Ophionereis mb x
eurybrachiplax H.
L. Clark, 1911
(continued)
Appendix
(m)
Appendix
Ophionereis perplexa 0–73 cr,rb x x x x

Ziesenhenne, 1940
Ophionereis porrecta 0–30 rb x
Lyman, 1860
Ophionereis schayeri x
1844)
Family Ophiotrichidae
Ophiothrix galapagensis 0–549 cr,mb,rb,sb x x
1899
Ophiothela gracilis 7–9 cr,rb x x x
Nielsen, 1932
Ophiothela mirabilis 5–20 cr,mb,rb x x x x x x x x
Verrill, 1867
Ophiothrix magnifica 0–20 cr,mb,rb,sb x x
Lyman, 1860
Ophiothrix rudis Lyman, 0–1 rb,cr,sb x
1874
Ophiothrix spiculata Le 0–2059 cr,mb,rb,sb x x x x x x x x x x
Conte, 1851
Family Ophiuridae
Subfamily
Ophiolepidinae
Ophiolepis crassa 6–230 mb,rb,sb x x
Nielsen, 1932
Ophiolepis grisea H. 7–13 mb x x x x x x
L. Clark, 1940
Ophiolepis pacifica 0–18 cr,rb x x x
Lütken, 1856
(continued)
571
572
(m)
Ophiolepis plateia 1–12 cr,rb,sb,rub x x x
Ziesenhenne, 1940
Ophiolepis variegata 1–110 cr,mb,sb x x x x x
Lütken, 1856
Ophiomusium glabrum 878– mb,rb,sb x x x x x x x x
1899
Ophiomusium diomedeae 702– x
1899
Ophiomusium lymani 51–2906 mb,sb x x x x x x
Thomson, 1873
Ophiomusium variabile 267–902 mb,rb x x
1899
Ophioplocus esmarki 0–74 rb,sg x
Lyman, 1874
Ophioplocus hancocki x
Ziesenhenne, 1935
Ophiosphalma jolliense 17–1230 mb,rb,sb x x x
(McClendon, 1909)
Ophiozonella alba 1408– x x x
1899)
Ophiozonella clypeata x
(Lyman, 1883)
Ophiozonella contigua 2417– x x
1899)
Ophiozonella falklandica x
Mortensen, 1936
Appendix
(continued)
(m)
Appendix
Subfamily
Ophioleucinae
Ophiernus adspersus 770– mb,sb x x x
adspersus Lyman, 1883 1245
Ophiernus adspersus 770– x
annectens Lütken & 1158
Mortensen, 1899
Ophiernus polyporum 1207– x
1899
Ophiernus seminudus 840– mb,sb x x x
1899
Subfamily Ophiurinae
Amphiophiura abcisa 245– rb,sb x x x
1899)
Amphiophiura x
irregularis Ziesenhenne,
1940
Amphiophiura obtecta 2197 x x x
1899)
Amphiophiura oligopora 1152 mb x
(H. L. Clark, 1913)
Amphiophiura x
paucisquama
Ziesenhenne, 1940
Amphiophiura superba 51–1820 mb x
1899)
573
(continued)
574
(m)
Amphiophiura vemae 3739– x
Kyte, 1987 4124
Gymnophiura concava x
Tommasi, 1976
Gymnophiura mollis 2417– x x x
1899
Ophiocten amitinum 29–920 rb,sb,kf x
Lyman, 1878
Ophiocten hastatum 1159– x x x x x x x
Lyman, 1878 2877
Ophiogona doederleini x
(Koehler, 1901)
Ophionotus victoriae 5–752 mb,rb,sb x
Bell, 1902
Ophiomastus bulufonica 1171– x
Tommasi, 1976 1180
Ophiomastus molinae intertidal rb x
Castillo-Alarcon, 1968
Ophiomastus tuberculata 1171– x
Tommasi, 1976 1180
Ophiomisidium leurum x
Ziesenhenne, 1940
Ophiosteira koehleri 733 x
A. H. Clark, 1917
Ophiophyllum 717 x
marginatum A. H. Clark,
1916
(continued)
Appendix
(m)
Appendix
Ophioplinthus divisa 2075– x

1899)
Ophioplinthus inflata 1484 x
(Koehler, 1897)
Ophioplinthus inornata 1485– x x
(Lyman, 1878) 2418
Ophioplinthus nexila x
(Kyte, 1987)
Ophiotypa simplex 3652– x
Koehler, 1897 3811
Ophiura 405- mb,sb x x x x x x x x
(Ophiuroglypha) 5869
irrorata irrorata
(Lyman, 1878)
Ophiura flagellata 128– mb x x
(Lyman, 1878) 2014
Ophiura leptoctenia 27–3239 sb,rub x
H. L. Clark, 1911
Ophiura luetkenii 0–1097 mb,rb,sb x x
(Lyman, 1860)
Ophiura lymani 30–800 rb,sb,kf x
(Ljungman, 1871)
Ophiura nana (Lütken & 1650 x
Mortensen, 1899)
Ophiura plana (Lütken 2070– x x x x x x
& Mortensen, 1899) 3241
(continued)
575
576
(m)
Ophiura stenobrachia 3667 x x
H. L. Clark, 1917
Theodoria madseni x
Tommasi, 1976
Class Echinoidea
Order Cidaroida
Family Cidaridae
Austrocidaris 1–845 rb, sb x
canaliculata (A. Agassiz,
1863)
Austrocidaris spinulosa 13–641 x
Mortensen, 1910
Centrocidaris 87–550 sb x x x x
doederleini (A. Agassiz,
1898)
Eucidaris thouarsii 0–150 cr,mb,rb,sb x x x x x x x x x x x
(Valenciennes, 1846)
Eucidaris thouarsii 0–150 rb x x
galapagensis
Doederlein, 1887
Hesperocidaris 2–183 cr,mb,rb,sb x x x x x x
asteriscus H. L. Clark,
1948
Hesperocidaris dubia 55–600 x x x x
(H. L. Clark, 1907)
Hesperocidaris 45 x
houstonianus H. Clark,
1939
Hesperocidaris 48–274 x x x x
panamensis (A. Agassiz,
1898)
Appendix
(continued)
(m)
Appendix
Hesperocidaris perplexa 10–1500 rb,mb,sb x x x x x x

(H. L. Clark, 1907)
Family Ctenocidaridae
Aporocidaris milleri 300– mb,rb,sb x x x x x
(A. Agassiz, 1898) 3937
Family Histocidaridae
Histocidaris cobosi (A. 702 x
Agassiz, 1898)
Order Echinothurioida
Family Echinothuriidae
Araeosoma eurypatum 1227 x
A. Agassiz & H.
L. Clark, 1909
Araeosoma leptaleum 740– rb x x
A. Agassiz & H. 1046
L. Clark, 1909
Tromikosoma hispidum 1820– rb,sb x x x x x x x
(A. Agassiz, 1898) 3375
Tromikosoma 2054– mb,rb x x x x
panamense (A. Agassiz, 3374
1898)
Family
Kamptosomatidae
Kamptosoma asterias 1020– mb x
(A. Agassiz, 1881) 1278
(continued)
577
578
(m)
Order Diadematoida
Family
Aspidodiadematidae
Plesiodiadema 2830– mb x x x
globulosum (A. Agassiz, 3900
1898)
Plesiodiadema horridum 1625– x x x x x
(A. Agassiz, 1898) 3381
Family Diadematidae
Astropyga pulvinata 0–90 cr,mb,rb,sb x x x x x x x x x x
(Lamarck, 1816)
Centrostephanus 0–125 cr,rb,sb x x x x x x x x
coronatus (Verrill, 1867)
Centrostephanus x
rodgersii (A. Agassiz,
1863)
Centrostephanus sylviae x
Fell, 1975
Diadema mexicanum 0–113 mb,cr,rb,sb x x x x x x x x x x x x
A. Agassiz, 1863
Diadema paucispinum x
(A. Agassiz, 1863)
Diadema savignyi 1–3 rb, sb x
Michelin, 1845
Echinothrix calamaris 0–46 cr,rb x
(Pallas, 1774)
Echinothrix diadema 3–5 cr x
(Linnaeus, 1758)
Order Pedinoida
Family Pedinidae
(continued)
Appendix
(m)
Appendix
Caenopedina diomedeae 723–850 x

Mortensen, 1939
Order Salenioida
Family Saleniidae
Salenocidaris miliaris 1159– x x x x x
(A. Agassiz, 1898) 3376
Order Arbacioida
Family Arbaciidae
Arbacia dufresnii 0–315 rb x
(Blainville, 1825)
Arbacia spatuligera 0–50 sb,rb,mb x x
(Valenciennes, 1846)
Arbacia stellata 0–92 cr,mb,rb, sb x x x x x x x x x x
(Blainville, 1825)
Dialithocidaris 3193– x x
gemmifera A. Agassiz, 3279
1898
Tetrapygus niger 0–25 rb,sb x x
(Molina, 1872)
Order Camarodonta
Family Parechinidae
Loxechinus albus 0–340 rb x x
(Molina, 1782)
Order Temnopleuroida
Family
Temnopleuridae
Pseudechinus 0–820 rb, kf x x
magellanicus (Philippi,
1857)
(continued)
579
580
(m)
Family Toxopneustidae
Lytechinus anamesus H. 1–207 cr,mb,rb,sb x
L. Clark, 1912
Lytechinus panamensis x
Mortensen, 1921
Lytechinus pictus 0–300 cr,rb,mb,sb x x x x x
(Verrill, 1867)
Lytechinus 0–134 rb,cr,sb x x x
semituberculatus
(Valenciennes in
L. Agassiz, 1846)
Tripneustes depressus 0–75 cr,rb,sb x x x x x x x x x x x x
A. Agassiz, 1863
Tripneustes gratilla 1–6 rb x x
(Linnaeus, 1758)
Toxopneustes roseus 0–55 cr,rb,sb,mb x x x x x x x x x x x
(A. Agassiz, 1863)
Family
Strongylocentrotidae
Allocentrotus fragilis 50–1200 mb,rb,sb x
Jackson, 1912
Strongylocentrotus 4–84 rb,sb x
franciscanus
(A. Agassiz, 1863)
Strongylocentrotus 0–161 mb,rb x
purpuratus (Stimpson,
1857)
Order Echinoida
Family Echinidae
Family Echinometridae
(continued)
Appendix
(m)
Appendix
Caenocentrotus gibbosus 0–35 rb x x x

(L. Agassiz & Desor,
1846)
Echinometra insularis H. 0–50 rb,sb x x x
L. Clark, 1912
Echinometra mathaei x
(Blainville, 1825)
Echinometra oblonga 0–34 cr,rb x x x x x
(Blainville, 1825)
Echinometra vanbrunti 0–106 cr,rb,sb x x x x x x x x x x x x x x
A. Agassiz, 1863
Echinostrephus x
aciculatus A. Agassiz,
1863
Order Clypeasteroida
Family Clypeasteridae
Clypeaster elongatus H. 10 sb x
L. Clark, 1948
Clypeaster europacificus 0–402 mb,sb,rb x x x x x x x
H. L. Clark, 1914
Clypeaster ochrus 0–162 rb,sb x x x x x x
H. L. Clark, 1914
Clypeaster reticulatus x
Linnaeus, 1758
Clypeaster rotundus 0–92 rb,sb,mb x x x x x x x x x
(A. Agassiz, 1863)
Clypeaster speciosus 0–128 rb,sb x x x x
Verrill, 1870
Family Dendrasteridae
(continued)
581
582
(m)
Dendraster excentricus 0–232 mb,rb,sb x
(Eschscholtz, 1831)
Dentraster laevis H. 13–22 sb x
L. Clark, 1948
Dendraster mexicanus 0–30 sb x
H. L. Clark, 1948
Dendraster vizcainoensis 0–30 sb x
Grant & Hertlein, 1938
Family Mellitidae
Encope cocosi 4–15 sb x
H. L. Clark, 1948
Encope ecuadorensis 1–3 sb x
H. L. Clark, 1948
Encope grandis 0–120 sb x x x x x x
L. Agassiz, 1841
Encope insularis 1–10 sb x
H. L. Clark, 1948
Encope irregularis 0–36.6 sb x x
H. L. Clark, 1948
Encope laevis 0–7 sb x x x x
H. L. Clark, 1948
Encope michelini 3–90 sb x
L. Agassiz, 1841
Encope micropora 0–82 mb,sb x x x x x x x x x x
L. Agassiz, 1841
Encope micropora 1–5 sb x x
ecuadorensis
H. L. Clark, 1948
Encope micropora 1–10 sb x
insularis H. L. Clark,
1948
Appendix
(continued)
(m)
Appendix
Encope micropora 1 sb x
irregularis H. L. Clark,
1948
Encope micropora 3 sb x
tetrapora L. Agassiz,
1841
Encope pacifica (Verrill, 0–40 sb x
1867)
Encope perspectiva 8–27 mb,sb x x x x
L. Agassiz, 1841
Encope wetmorei 1–45 sb x x x x
A. H. Clark, 1946
Mellita grantii 0–10 sb x x
Mortensen, 1948
Mellita kanakoffi 0–102.9 sb x x x
Durham, 1961
Mellita longifissa 0–60 sb x x x x x x x x x x
Michelin, 1858
Mellita notabilis 0.5–0.9 sb x x x x
H. L. Clark, 1947
Mellitella stokesii 0–10 sb, mb x x x x x
(L. Agassiz, 1841)
Order Cassiduloida
Family Cassidulidae
Rhyncholampas pacificus 6–134 rb,sb x x x x
(A. Agassiz, 1863)
Order Holasteroida
Family Pourtalesiidae
Cystocrepis setigera 3182 x
(A. Agassiz, 1898)
(continued)
583
584
(m)
Pourtalesia tanneri x
A. Agassiz, 1898
Familly Urechinidae
Cystechinus loveni x
A. Agassiz, 1898
Pilematechinus rathbuni 3374 x x
(A. Agassiz, 1898)
Urechinus naresianus 755– x
A. Agassiz, 1879 4400
Order Spatangoida
Family Aeropsidae
Aeropsis fulva 1455– mb x x x x
(A. Agassiz, 1898) 5200
Family Brissidae
Brissopsis columbaris 589– mb,sb x x x
A. Agassiz, 1898 3279
Brissopsis pacifica 9–237 sb,mb,rb x x x x x x
(A. Agassiz, 1898)
Brissus agassizii x
Doederlein, 1885
Brissus obesus Verrill, 0–240 sb,cr,rb x x x x x x x
1867
Brissus latecarinatus x
(Leske, 1778)
Meoma frangibilis 96 x
Chesher, 1970
Meoma ventricosa 0–200 sb x x x x x x
grandis Gray, 1851
Metalia nobilis Verrill, 0–18 sb x x
1867
(continued)
Appendix
(m)
Appendix
Metalia spatagus x
(Linnaeus, 1758)
Rhabdobrissus pacificus 6–137 sb,mb,sb x x x
H. L. Clark, 1940
Family Hemiasteridae
Hemiaster tenuis 980– x x x x
(A. Agassiz, 1898) 4027
Family Loveniidae
Homolampas hastata 1785– x x x x
A. Agassiz, 1898 3376
Lovenia cordiformis 0–210 mb,sb x x x x x x
A. Agassiz, 1872
Family
Macropneustidae
Argopatagus aculeata 1952 x
(A. Agassiz, 1898)
Family Maretiidae
Nacospatangus oblonga x
(Mortensen, 1950)
Family Palaeotropidae
Scrippsechinus fisheri 270–460 x
Allison, Durham &
Mintz, 1967
Family Prenasteridae
Tripylus excavatus 0–130 x
Philippi, 1845
Family Schizasteridae
Abatus cavernosus 0–760 x
(Philippi, 1845)
(continued)
585
586
(m)
Abatus philippii Lovén, 27–804 x
1871
Aceste ovata A. Agassiz x
& H. L. Clark, 1907
Agassizia scrobiculata 0–209 mb,rb,sb x x x x x x x x
Valenciennes, 1846
Brisaster latifrons 9–2817 mb,sb x x x
(A. Agassiz, 1898)
Brisaster moseleyi 124–730 x
(A. Agassiz, 1881)
Brisaster townsendi x
(A. Agassiz, 1898)
Moira atropos clotho 10–40 mb,sb x x x
Michelin, 1855
Schizaster rotundatus x
(Doederlein, 1906)
Tripylaster philippii 0–100 sb x
(Gray, 1851)
Family Spatangidae
Spatangus californicus 10–644 mb,rb,sb x
H. L. Clark, 1917
Class Holothuroidea
Order Dendrochirotida
Family Cucumariidae
Abyssocucumis 3241– x x x x
abyssorum (Théel, 1886) 4000
Abyssocucumis 1585– x
albatrossi Cherbonnier, 5690
1947
(continued)
Appendix
(m)
Appendix
Cucumaria chilensis 32–162 x

Ludwig, 1875
Cucumaria flamma 0–15 rb x x x x x x
Solís-Marín & Laguarda-
Figueras, 1999
Cladodactyla crocea 0–4300 sb x
(Lesson, 1830)
Heterocucumis 0–379 rb,sb x x
godeffroyi (Semper,
1868)
Hemioedema spectabilis 14642 sb x
(Ludwig, 1882)
Leptopentacta nina x
Deichmann, 1941
Leptopentacta nova x
Deichmann, 1941
Leptopentacta panamica x
Deichmann, 1941
Neocucumis panamensis x
Heding & Panning, 1954
Neocucumis veleronis 0–22 rb x x x
(Deichmann, 1941)
Pattalus mollis Selenka, 0–45 rb x x
1868
Pentacta panamensis x
Verrill, 1867
Pseudocnus californicus 0–190 cr,rb,sb x x x x x x x x
(Semper, 1868)
Pseudocnus dubiosus 0–300 rb,sb x
(Semper, 1868)
(continued)
587
588
(m)
Pseudocnus dubiosus 0–300 rb,sb x x
leoninus (Semper, 1868)
Pseudocnus perrieri 0–197 rb,sb x
(Eckman, 1927)
Staurocucumis 1587– mb,rb x x x
abyssorum (Théel, 1886) 4088
Stereoderma laevigata x
Verrill, 1876
Thyonella mexicana 10–35 sb,rub x x
(Deichmann, 1941)
Trachythyone lechleri 5–238 rb x
(Lampert, 1885)
Trachythyone peruana 5–42 rb x x
(Semper, 1868)
Family Phyllophoridae
Allothyone mexicana x x
(Deichmann, 1946)
Athyonidium chilensis 0–13 sb,rb,kf x x
(Semper, 1868)
Euthyonidium ovulum x
Deichmann, 1938
Euthyonidium veleronis 0.3 x x
Deichmann, 1937
Pentamera beebei 73 x
Deichmann, 1938
Pentamera chierchia 0–78 cr,sb,rb x x x x x x
(Ludwig, 1887)
Pentamera chiloensis 6–124 sb x
(Ludwig, 1887)
(continued)
Appendix
(m)
Appendix
Pentamera zacae 24 mb,rb,sb x x x

Deichmann, 1938
Phyrella aculeata x
(Ludwig, 1894)
Thyone bidentata 2–30 cr,rb x x x x
Deichmann, 1941
Thyone neofusus x
Deichmann, 1941
Thyone parafusus 25–35 mb,sb x
Deichmann, 1941
Thyone strangeri 0–12 rb x
Deichmann, 1941
Family Psolidae
Lissothuria hancocki 1–301 rb x
(Deichmann, 1941)
Lissothuria mortenseni x
Pawson, 1967
Lissothuria ornata 0–37 cr,rb x x x x x x
Verrill, 1867
Lissothuria veleronis x
(Deichmann, 1941)
Neopsolidium 15 kf x
convergens (Herouard,
1901)
Psolidium disciformis 8–448 rb x
(Théel, 1886)
Psolidium dorsipes 11–451 cr,mb,rb,sb x x x x
Ludwig, 1886
Psolidium ekmani x
Deichmann, 1941
(continued)
589
590
(m)
Psolidium eubullatum x
Deichmann, 1941
Psolidium gracile 2323 x
Ludwig, 1894
Psolidium panamense 2323 x
Ludwig, 1894
Psolidium planum 16–110 x
Deichmann, 1941
Psolus antarcticus 6–1080 rb x
Philippi, 1857
Psolus chitonoides 0–247 rb x
H. L. Clark, 1901
Psolus digitatus Ludwig, 1271 x
1894
Psolus diomedeae 13–302 rb,sb x x x x
Ludwig, 1894
Psolus paradubiosus 10–567 rb x
Carriol & Féral, 1985
Psolus patagonicus 0–110 rb,kf x
Ekman, 1925
Psolus squamatus 7–1087 rb x x
(Koren, 1844)
Psolus squamatus 7–207 rb x
segregatus Perrier, 1905
Family
Sclerodactylidae
Afrocucumis ovulum 0–7 rb,cr x x x x
(Selenka, 1867)
Apentamera lepra 55–91 sb x x
Deichmann, 1941
(continued)
Appendix
(m)
Appendix
Athyone glasselli 0–6 sb x

(Deichmann, 1936)
Euthyonidiella zacae 17–25 rb x x x
(Deichmann, 1938)
Neopentamera anexigua 8–22 cr x x
Deichmann, 1941
Neothyone gibber 0–50 cr,rb x x x x x x x
(Selenka, 1867)
Neothyone gibbosa 0–14 rb x x x x
Deichmann, 1941
Neothyone panamensis 0–8.6 rb x x
(Ludwig, 1887)
Pachythyone lugubris cr,rb x
(Deichmann, 1939)
Pachythyone 12 cr,rb x
pseudolugubris
Deichmann, 1941
Order Dactylochirotida
Family Ypsilothuriidae
Ypsilothuria 225– mb,sb x x x x x x x x
bitentaculata (Ludwig, 4082
1893)
Order Aspidochirotida
Family Holothuriidae
Actinopyga mauritiana x
(Quoy & Gaimard, 1833)
Holothuria (Cystipus) 45–100 sb x x
casoae Laguarda-
Figueras & Solís-Marín,
2009
(continued)
591
592
(m)
Holothuria (Cystipus) 1–203 cr,rb,sb x x x x x x x
inhabilis Selenka, 1867
Holothuria (Cystipus) 0–22 mb,sb,rb x x x x x
rigida (Selenka, 1867)
Holothuria (Halodeima) 5–26 mb,cr,rb,sb x x x x x
atra (Jaeger, 1833)
Holothuria (Halodeima) x
chilensis Semper, 1868
Holothuria (Halodeima) 0–18 rb x x x x x
inornata Semper, 1868
Holothuria (Halodeima) 0–27 rb,sb,cr x x x x x x x x x x x
kefersteini (Selenka,
1867)
Holothuria 0–306 cr,rb,mb,sb x x x x x x x x x x
(Lessonothuria) pardalis
Selenka, 1867
Holothuria 0.3–52 cr,rb,sb x x x x x x x x
(Mertensiothuria) hilla
Lesson, 1830
Holothuria 1–17 rb,sb x x x x x x x
(Mertensiothuria)
leucospilota (Brandt,
1835)
Holothuria 0–100 mb,rb,sb,cr x x x x x x x x x x x
(Platyperona) difficilis
Semper, 1868
Holothuria 1–4 x x
(Platyperona) parvula
(Selenka, 1867)
(continued)
Appendix
(m)
Appendix
Holothuria 0–6 rb x x
(Selenkothuria) carere
Honey-Escandón &
Solís-Marín, 2011
Holothuria 0–55 cr,rb,sb,mb x x x x x x x x
(Selenkothuria) lubrica
Selenka, 1867
Holothuria 0–59 cr,rb x x x x x x x
(Selenkothuria)
portovallartensis Caso,
1954
Holothuria 0–55 rb x x x x x
(Selenkothuria) theeli
Deichmann, 1938
Holothuria x
(Semperothuria)
cinerascens (Brandt,
1835)
Holothuria 0–22 rb,sb,cr x x x x x x x x x x
(Semperothuria) imitans
Ludwig, 1875
Holothuria 0–91 rb,mb,sb x x x x
(Semperothuria)
languens Selenka, 1867
Holothuria (Stauropora) 0–130 rb,cr,sb x x x x x x x
pluricuriosa
Deichmann, 1937
Holothuria 0–64 cr,rb,sb x x x
(Theelothuria)
paraprinceps
Deichmann, 1937
593
(continued)
594
(m)
Holothuria 1–121 rb,sb,cr x x x x x x x x x
(Thymiosycia) arenicola
Semper, 1868
Holothuria 0–67 cr,rb,sb x x x x x x x x x x
(Thymiosycia) impatiens
(Forskaal, 1775)
Holothuria 77–250 mb,rb,sb x x x x x x x
(Vaneyothuria) zacae
Deichmann, 1937
Labidodemas 0.5–16 rb,cr,sb x x x x x x x
americanum Deichmann,
1938
Labidodemas 1–18 rb,sb,cr x x x x x
maccullochi
(Deichmann, 1958)
Family Stichopodidae
Apostichopus 0–36 mb,rb,sb x
parvimensis (H. L. Clark,
1913)
Isostichopus fuscus 0–61 cr,rb,mb,sb x x x x x x x x x x x x x x
(Ludwig, 1875)
Parastichopus 1–180 mb,rb x
californicus (Stimpson,
1857)
Stichopus horrens 0–20 cr,rb,sb x x x x
Selenka, 1867
Stichopus x
monotuberculatus (Quoy
& Gaimard, 1833)
Family Synallactidae
(continued)
Appendix
(m)
Appendix
Bathyplotes moseleyi 24–690 mb,rb x x

(Théel, 1886)
Bathyplotes natans 1644 x x
(M. Sars, 1868)
Östergren, 1896
Bathyplotes patagiatus 1644 x
Fisher, 1907
Capheira sulcata 2877– mb x x
Ludwig, 1893 4334
Meseres macdonaldi 1644 x x
Ludwig, 1894
Meseres torvus (Théel, x
1886)
Mesothuria (Mesothuria) 725– mb x x x x
multipes Ludwig, 1894 4064
Mesothuria (Zygothuria) x
lactea (Théel, 1886)
Molpadiodemas x
atlanticus (R. Perrier,
1898)
Molpadiodemas 2487– x
neovillosus O’Loughlin 3667
& Ahearn, 2005
Molpadiodemas x
ustulatus O’Loughlin &
Ahearn, 2005
Molpadiodemas villosus 3667 x
(Théel, 1886)
Molpadiodemas x
violaceus (Théel, 1886)
(continued)
595
596
(m)
Paelopatides confundens 1571– mb x x
Théel, 1886 4070
Paelopatides suspecta 2323 x
Ludwig, 1894
Pseudostichopus 2149 x
macdonaldi (Ludwig,
1894)
Pseudostichopus mollis 100– mb,sb x x x x x x
Théel, 1886 5203
Pseudostichopus 1158– x x x
peripatus (Sluiter, 1901) 3667
Synallactes aenigma 2418– mb,sb x x x x x
Ludwig, 1894 4334
Synallactes alexandri 589– x x
Ludwig, 1894 1008
Synallactes triplax 549 x
A. H. Clark, 1920
Synallactes virgulasolida 1030 mb x
Massin & Hendrickx,
2010
Order Elasipodida
Family Deimatidae
Deima validum 1618– mb,sb x x x x
pacificum Ludwig, 1894 2487
Oneirophanta mutabilis 3241 x x x
affinis Ludwig, 1894
Oneirophanta mutabilis 3241– mb x x x
mutabilis Théel, 1879 3670
Oneirophanta setigera 3667– x
(Ludwig, 1893) 4088
(continued)
Appendix
(m)
Appendix
Family Elpidiidae
Achlyonice ecalcarea 1608– mb,sb x
Théel, 1879 3587
Amperima naresi (Théel, 2010– mb x
1882) 7130
Amperima vitjazi x
Gebruk, 1988
Elpidia atakama 7720 mb x
Belyaev, 1971
Elpidia chilensis 2710– mb x
Belyaev, 1971 4600
Peniagone anamesa x
(A. H. Clark, 1920)
Peniagone diaphana 4140– mb x
(Théel, 1882) 4160
Peniagone elongata 4140– mb x
(Théel, 1879) 4160
Peniagone gracilis 2487– mb x x
(Ludwig, 1894) 4160
Peniagone intermedia 2418– mb x x
Ludwig, 1893 3667
Peniagone papillata x
Hansen, 1975
Peniagone vitrea Théel, 1160– mb x x x x x x
1882 4507
Scotoplanes clarki 3570– mb x x
Hansen, 1975 5107
Scotoplanes globosa 545– mb x
Théel, 1879 6770
(continued)
597
598
(m)
Family Laetmogonidae
Laetmogone theeli 2418 x
Ludwig, 1893
Laetmogone violacea 466– x
Théel, 1879 5083
Pannychia moseleyi 199– mb,sb x x x x
Théel, 1882 2599
Family Pelagothuridae
Enypniastes eximia 4140– pelagic x
Théel, 1882 4160
Pelagothuria natatrix 0–4505 pelagic x x x x x x x
Ludwig, 1893
Family Psychropotidae
Benthodytes incerta 2418 x
Ludwig, 1893
Benthodytes 978– mb x x x x x x
sanguinolenta Théel, 2323
1882
Benthodytes typica 1158– mb x x x
Théel, 1882 4700
Psychronaetes hanseni 3852– mb x
Pawson, 1983 4189
Psychropotes depressa x
(Théel, 1882)
Psychropotes longicauda 3334– mb x x x x x
Théel, 1882 4160
Psychropotes verrucosa 2418– mb x x x x
(Ludwig, 1894) 4160
Order Molpadiida
(continued)
Appendix
(m)
Appendix
Family Caudinidae
Ceraplectana 5203 mb x
trachyderma H. L. Clark,
1908
Hedingia californica 85–2850 sb,mb x x
(Ludwig, 1894)
Paracaudina chilensis 0–990 mb,sb x x
chilensis (Müller, 1850)
Family Molpadiidae
Molpadia antarctica 80–1218 mb,sb x
(Théel, 1886)
Molpadia intermedia 53–2972 mb x x x
(Ludwig, 1894)
Molpadia granulata 2690– mb x x
(Ludwig, 1894) 5869
Molpadia musculus 37–6134 mb,sb x x x x x x x x x
Risso, 1826
Molpadia spinosa 3279 x
(Ludwig, 1893)
Order Apodida
Family Chiridotidae
Chiridota aponocrita A. 9–137 cr,rb,sb x x
H. Clark, 1920
Chiridota pacifica x
Heding, 1928
Chiridota pisanii 0–228 mb,sb x
Ludwig, 1886
Chiridota rigida Semper, 2–9 cr,rb,sb x
1868
(continued)
599
600
(m)
Taeniogyrus contortus 0–560 mb,sb,kf x
(Ludwig, 1875)
Trochodota purpurea 0–36 rb,sb x
(Lesson, 1830)
Family Myriotrochidae
Myriotrochus giganteus 3667 mb x x
A. H. Clark, 1920
Myriotrochus 3667 mb x
(Oligotrochus) bathybius
A. H. Clark, 1920
Family Synaptidae
Anapta fallax Lampert, 0–350 sb x
1889
Epitomapta tabogae 0–10 x x
Heding, 1928
Euapta godeffroyi 0–79 cr,rb,sb x x x x x
(Semper, 1868)
Polyplectana oculata x
Heding, 1928
Protankyra abyssicola x
(Théel, 1886)
Protankyra brychia 3900– x x x x
(Verrill, 1885) 4000
Protankyra pacifica 870– mb x x x
(Ludwig, 1894) 4990
Habitat classification: cr Coral Reefs, m mangroves, mb muddy bottom, rb rocky bottom, sb sandy bottom, sg seagrass, rub rubble bottom, kf kelp forest. Countries
abbreviation: MEX Mexico, REV Revillagigedo Archipelago, GUA Guatemala, SAL El Salvador, HON Honduras, NIC Nicaragua, CRC Costa Rica, COC Cocos Island,
PAN Panama, COL Colombia, MAL Malpelo Island, GAL Galapagos Archipelago, ECU Ecuador, PER Perú, CHL Chile, PAS Eastern Islands
Appendix
Table A.2 Taxonomic list of the Echinoderms of the Atlantic Ocean and Caribbean Sea of Latin America and the Canary Islands
Depth Habitat MEX BEL GUA HON NIC CRC PAN COL VENc VENa BRA URG ARG MAV CUB HAI RDO PRI CAN
(m)
Appendix
Class Crinoidea
Order Comatulida
Family Antedonidae
Antedon bifida (Pennant, 1777) 0–400 rb,rub x x
Antedon nuttingi (A. H. Clark, 364–429 rb x
1936)
Caryometra alope A. H. Clark, 475–640 rb x
1940
Caryometra atlantidis A. 366–530 mb x x
H. Clark, 1940
Caryometra lisa A. H. Clark, 702 rb x
1940
Caryometra monilicirra x
A. H. Clark, 1940
Caryometra spinosa x
A. H. Clark, 1940
Caryometra tenuipes 386 rb x x
(A. H. Clark, 1908)
Coccometra guttata 288–480 mb,rb,rub x x x
A. H. Clark, 1918
Coccometra hagenii (Pourtalès, 14–1046 rb,rub x x
1867)
Coccometra nigrolineata 40–987 cr,rb,rub x x x x
A. H. Clark, 1918
Comatonia cristata (Hartlaub, 50–396 cr,rb x
1912)
Ctenantedon kinziei Meyer, Sep-49 cr x x x
1972
Hypalometra defecta 46–400 cr,mb,rb x x x x x x
(Carpenter, 1888)
Isometra vivipara Mortensen, 79–350 mb,sb,rub x x
1917
Leptometra celtica (M’Andrew x
& Barrett, 1858)
Phrixometra nutrix (Mortensen, 137–150 mb,rb,sb x x
1918)
(continued)
601
602
(m)
Poliometra prolixa (Sladen, 219 x
1881)
Trichometra cubensis 210– mb,rb x x x x
(Pourtalès, 1869) 4829
Zenometra columnaris 309– mb,rb x x
(Carpenter, 1881) 1033
Family Pentametrocrinidae
Pentametrocrinus atlanticus 860– mb,sb x
(Perrier, 1883) 1674
Superfamily Comasteracea
Family Comasteridae
Comactinia echinoptera 2–1033 cr,rb,sb,sg,rub x x x x x x x x x x x x x x
(Müller, 1840)
Comactinia meridionalis 3–1033 cr,rb,sb x x x x x x x x x x x
meridionalis (L. Agassiz, 1865)
Comactinia meridionalis 58–373 cr,rb x x x
hartlaubi Messing, 1978
Comissia venustus 24–479 cr,mb,rb,rub x x x x x
(A. H. Clark, 1909)
Davidaster discoideus 0–640 cr,br,rub x x x x x x x x x x x
(Carpenter, 1888)
Davidaster rubiginosus 0–334 cr,rb x x x x x x x x x x x x
(Pourtalès, 1869)
Leptonemaster venustus 55–777 mb x x x
A. H. Clark, 1909
Nemaster grandis A. H. Clark, 3–124 cr,rb x x x x x x
1909
Neocomatella alata (Pourtalès, 10–560 cr,br x x x x
1878)
Neocomatella europaea 466–960 rb,rub x
A. H. Clark, 1913
Neocomatella pulchella 3–695 cr,br x x x x x x
(Pourtalès, 1878)
Family Colobometridae
Analcidometra armata 3–155 cr x x x x x x x x x x
(Pourtalès, 1869)
(continued)
Appendix
(m)
Appendix
Analcidometra caribbea 23–26 x

(A. H. Clark, 1908)
Family Charitometridae
Crinometra brevipinna 69–795 cr,rb x x x x x x x
brevipinna (Pourtalès, 1867)
Crinometra brevipinna 201–367 rb x
concinna A. H. Clark, 1909
diadema (Hartlaub, 1912)
granulifera (Pourtalès, 1878)
margaritacea A. H. Clark, 1909
Crinometra brevipinna pulchra 484–567 rb x
A. H. Clark, 1909
Family Thalassometridae
Horaeometra duplex 159–567 rb x x x
(Carpenter, 1888)
Stylometra spinifera 69–658 cr,rb,sb,rub x x x x x x x x
(Carpenter, 1881)
Family Tropiometridae
Tropiometra carinata 0–84 cr,rb,rub x x x x
(Lamarck, 1816)
Family Atelecrinidae
Atelecrinus balanoides 512– mb,rb,sb x x x x x
Carpenter, 1881 2890
Family Bathycrinidae
Bathycrinus gracilis Wyville- x
Thomson, 1877
Monachocrinus caribbeus 684– mb,rb x x x
(A. H. Clark, 1908) 1574
Family Bourgueticrinidae
Conocrinus lofotensis (Sars, x
1868)
Democrinus conifer 155– cr,mb,rb x x x x x x x
(A. H. Clark, 1909) 1750
603
(continued)
604
(m)
Democrinus brevis 374– mb x x
(A. H. Clark, 1909) 1574
Democrinus rawsoni 70–650 mb,rb x x x x x
(Pourtalès, 1874)
Family Septocrinidae
Rouxicrinus vestitus Mironov & 421–887 mb x
Pawson, 2010
Order Isocrinida
Family Isocrinidae
Endoxocrinus (Diplocrinus) x
wyvillethomsoni (Jeffreys,
1870)
Endoxocrinus parrae (Gervais, 154–971 cr,mb,rb x x x x x x
1835)
Endoxocrinus prionodes 80.7 rb x
(H. L. Clark, 1941)
Isocrinus blakei (Carpenter, 220– cr,rb,rub x x
1884) 1200
Neocrinus decorus Thomson, 154– mb,rb,rub x x x x x x
1864 1219
Family Isselicrinidae
Cenocrinus asterius (Linnaeus, 170–412 cr,mb,rb x x x
1767)
Order Cyrtocrinida
Family Holopodidae
Holopus rangii Orbigny, 1837 9–700 rb x x x
Class Asteroidea
Order Paxillosida
Family Astropectinidae
Astropecten acutiradiatus 35–66 sb x x
Tortonese, 1956
Astropecten alligator Perrier, 22–576 cr,mb,sb,rub x x x x x x x
1881
Astropecten americanus 110–641 mb,sb,rub x x x x
Verrill, 1880
(continued)
Appendix
(m)
Appendix
Astropecten antillensis Lütken, 3–278 mb,sb,rub x x x x x

1860
Astropecten aranciacus 1–180 sb,rub x
(Linnaeus, 1758)
Astropecten articulatus Say, 0–256 mb,sb x x x x x x x x x x x x x x
1825
Astropecten brasiliensis Müller 2–60 sb x x x x
& Troschel, 1842
Astropecten brasiliensis riensis 18–66 sb x
Doederlein, 1917
Astropecten 51.3 rub x
caribemexicanensis Caso, 1990
Astropecten cingulatus Sladen, 16–1350 mb,sb,rub x x x x x x x x x
1883
Astropecten comptus Verrill, 18–130 sb,rub x x
1915
Astropecten duplicatus Gray, 0–550 mb,sb,sg,rub x x x x x x x x x x
1840
Astropecten hermatophilus 15–1500 x
Sladen, 1883
Astropecten irregularis 10–1000 mb,sb,rub x
(Pennant, 1777)
Astropecten marginatus Gray, 0.5–130 mb,sb,rub x x x x x x x x
1840
Astropecten nitidus Verrill, 11–686 mb,sb,rub x x x x x
1915
Astropecten spiniphorus 45 x
Madsen, 1950
Astropectinides mesactus 80–165 x
(Sladen, 1883)
Bathybiaster loripes Sladen, 80–500 mb,sb x x x
1889
Blakiaster conicus Perrier, 55–366 sb x x x x
1881
Dipsacaster antillensis 113– cr x
Halpern, 1968 3627
(continued)
605
606
(m)
Dytaster grandis nobilis 3305– x
Sladen, 1889 4846
Dytaster insignis Perrier, 1884 2209– sb x
3654
Leptychaster kerguelenensis 27–183 x x
E. A. Smith, 1876
Lonchotaster tartareus Sladen, 4390 x
1889
Persephonaster echinulatus 196–720 cr,mb,sb x x x x x x x x
H. L. Clark, 1941
Persephonaster leptactis 2928– x
H. L. Clark, 1941 3294
Persephonaster patagiatus 733– mb,sb x x x x x x x
(Sladen, 1889) 2165
Plutonaster agassizi agassizi 70–3110 cr,mb,sb x x x
Verrill, 1880
Plutonaster bifrons (Wyville- 800– x x
Thomson, 1873) 2970
Plutonaster efflorescens 666– mb,sb x x x
(Perrier, 1884) 2061
Psilaster andromeda florae 1409– mb,sb x
(Verrill, 1878) 1629
Psilaster cassiope Sladen, 1889 151– mb,sb x x x x x
1680
Psilaster herwigi (Bernasconi, 100–800 x x
1972)
Tethyaster grandis (Verrill, 24–139 mb,sb x x x x
1899)
Tethyaster subinermis (Philippi, 50–1400 mb x
1837)
Tethyaster vestitus vestitus 5–306 mb,sb x x x x x x
(Say, 1825)
Family Ctenodiscidae
Ctenodiscus australis Lütken, 70–4605 sb,rub x x x
1871
(continued)
Appendix
(m)
Appendix
Ctenodiscus crispatus (Retzius, 1415– x

1805) 1865
Family Goniopectinidae
Goniopecten demonstrans 300– mb,sb x x x x x
Perrier, 1881 1250
Prionaster elegans Verrill, 208–535 mb,sb x x x x x
1899
Family Luidiidae
Luidia alternata alternata (Say, 0–1500 m,mb,sb x x x x x x x x x x x x
1825)
Luidia bardabensis Perrier, 60–430 mb,rb,sb x x x x x x
1881
Luidia ciliaris (Philippi, 1837) 1–400 sb x
Luidia clathrata (Say, 1825) 0–175 mb,rb,sb,rub x x x x x x x x x x x x
Luidia heterozona barimae 16–90 mb,sb x x x x x
John & A. M. Clark, 1954
Luidia lawrencei Hopkins & 0–175 mb x
Knott, 2010
Luidia ludwigi scotti Bell, 1917 33–157 mb,rb,sb x x x x
Luidia patriae Bernasconi, 100–126 mb,sb x
1941
Luidia sagamina aciculata 71–83 sb x
Mortensen, 1933
Luidia sarsi Düben & Koren, x
1845
Luidia sarsi elegans Perrier, 60–365 mb,sb x x
1875
Luidia senegalensis (Lamarck, 0–73 mb,sb x x x x x x x x x x x x x
1816)
Family Porcellanasteridae
Eremicaster vicinus Ludwig, 3950– mb,rub x
1907 7250
Styracaster horridus Sladen, 3410– sb x
1883 5062
Thoracaster cylindratus Sladen, 2540– mb,sb x x x
1883 5990
607
(continued)
608
(m)
Family Pseudarchasteridae
Pseudarchaster discus Sladen, 140–283 x x
1889
Pseudarchaster gracilis 170– mb,sb x x x x x x x x x
gracilis (Sladen, 1889) 2940
Pseudarchaster parelii (Düben 265–869 x
& Koven, 1846 )
Order Notomyotida
Family Benthopectinidae
Benthopecten simplex simplex 1175– mb,sb,rb x x
(Perrier, 1881) 3713
Benthopecten spinosus Verrill, 1857– mb x
1884 1958
Cheiraster (Barbadosaster) 130– cr,mb,sb,rb x x x x x x x x
echinulatus (Perrier, 1875) 5062
Cheiraster (Cheiraster) ludwigi 1266 sb,rb x
Fisher, 1913
Cheiraster (Cheiraster) planus 226– mb,sb,rb x x x x
Verrill, 1915 1339
Cheiraster (Cheiraster) sepitus 485– mb,sb x x x x
(Verrill, 1885) 5062
Cheiraster (Christopheraster) 250– mb,sb,rb x x x x x x
blakei A. M. Clark, 1981 1958
Cheiraster (Christopheraster) 380– mb,sb,rb x x x x x
mirabilis (Perrier, 1881) 1470
Cheiraster (Luidiaster) planeta 370–500 mb x x
(Sladen, 1889)
Gaussaster antarcticus (Sladen, 3305 mb x
1889)
Pectinaster gracilis Verrill, 576 x
1915
Order Valvatida
Family Asterinidae
Allopatiria ocellifera (Gray, 30–200 mb,rub x
1847)
(continued)
Appendix
(m)
Appendix
Asterina fimbriata Perrier, 1875 0–250 mb,sb,rub x x

Asterina gibbosa (Pennant, 0–130 rb,rub x
1777)
Asterina stellifera (Moebius, 0–50 mb,sb,rub x x x
1859)
Asterinides folium (Lütken, 0–256 cr,rb,sb,rub x x x x x x x
1860)
Asterinides hartmeyeri 0–1 rb x x x
(Doederlein, 1910)
Asterinides pompom 3–6 rb x x
(A. M. Clark, 1983)
Asterinopsis pilosa (Perrier, 11–256 mb,sb x x
1881)
Tremaster mirabilis Verrill, 150– rb,rub x x x
1880 1060
Stegnaster wesseli (Perrier, 0–183 cr,rb,sb,rub x x x x
1875)
Family Asteropseidae
Poraniella echinulata (Perrier, 3–339 cr,rb x x x x x
1881)
Family Chaetasteridae
Chaetaster nodosus Perrier, 30–1140 mb,rb,sb x x x x x
1875
Family Ganeriidae
Cycethra verrucosa (Philippi, 0–500 sb,rub x x x
1857)
Ganeria falklandica Gray, 1847 0–135 sb,rub x x
Perknaster sladeni (Perrier, 120–500 x x
1891)
Vemaster sudatlanticus 5055– x
Bernasconi, 1965 5208
Family Goniasteridae
Anthenoides piercei Perrier, 20–844 mb,sb x x x x x x x x x
1881
Apollonaster yucatanensis 1097– mb,sb x x
Halpern, 1970 1175
609
(continued)
610
(m)
Astroceramus brachyactis 420– mb,sb x x x
H. L. Clark, 1941 1067
Ceramaster grenadensis 70–3109 mb,sb x x x x x x
(Perrier, 1881)
Ceramaster grenadensis 200– mb x x x
grenadensis (Perrier, 1881) 3109
Ceramaster grenadensis 106–192 x x
patagonicus (Sladen, 1889)
Circeaster americanus 500– mb,sb x x x x
(A. H. Clark, 1916) 1450
Cladaster rudis Verrill, 1899 150–900 mb,sb x
Diplasiaster productus 78–567 mb,sb x
(A. H. Clark, 1917)
Floriaster maya Downey, 1980 933– mb,sb x
1024
Goniaster tessellatus (Lamarck, 16–155 cr,mb,sb,rb x x x x
1816)
Hippasteria falklandica Fisher, 251–225 x x
1940
Hippasteria phrygiana 108–162 x
argentinensis Bernasconi, 1961
Litonotaster intermedius 1958– mb,rb,sb x x x x x
(Perrier, 1884) 3530
Mediaster bairdi (Verrill, 1882) 640– sb,rb x x
1590
Mediaster pedicellaris (Perrier, 197–580 mb,rb,sb x x x
1881)
Nymphaster arenatus (Perrier, 60–3000 mb,sb x x x x x x x x x x x
1881)
Paragonaster grandis H. 257–540 sb x x
L. Clark, 1941
Paragonaster subtilis (Perrier, 1845– rb,sb x
1881) 4700
Pawsonaster parvus (Perrier, 30–600 mb,rb,sb x x x x
1881)
(continued)
Appendix
(m)
Appendix
Peltaster placenta (Müller & 10–1107 mb,sb x x x x x

Trochel, 1842)
Plinthaster dentatus (Perrier, 60–2910 mb,rb,sb x x x x x x x x
1884)
Rosaster alexandri (Perrier, 60–2940 cr,mb,sb x x x x x x
1881)
Tessellaster notabilis 329–575 mb,sb x x
H. L. Clark, 1941
Tosia parva (Perrier, 1881) 100–600 x x x
Family Leilasteridae
Leilaster radians (Perrier, 102–293 sb,rub x x x
1881)
Family Mithrodiidae
Mithrodia clavigera (Lamarck, 0–157 cr,rb,sb x x x x
1816)
Familia Odontasteridae
Acodontaster elongatus 40–840 sb,rub x x
granuliferus (Koehler, 1912)
Diplodontias singularis (Müller 0–84 rb,sb x
& Troschel, 1843)
Odontaster penicillatus 8–350 mb,sb,rub x x
(Philippi, 1870)
Odontaster hispidus Verrill, 50–1160 mb,sb x x
1880
Family Ophidiasteridae
Copidaster cavernicola Solís- 13–18 mb x
Marín & Laguarda-Figueras,
2010
Copidaster lymani A. H. Clark, 0–34 cr,mb,rb x x x x
1948
Hacelia attenuata (Gray, 1840) 1–150 sb,rub x
Hacelia superba H. L. Clark, 91–200 rb x x
1921
Linckia bouvieri Perrier, 1875 0–380 cr,rb x x
Linckia guildingi Gray, 1840 0–298 cr,sg,rb,sb x x x x x x x x x x x x
(continued)
611
612
(m)
Linckia nodosa Perrier, 1875 0–475 cr,rb,sb x x x x x
Narcissia canariensis 37–155 rb x x
(d’Orbigny, 1839)
Narcissia trigonaria Sladen, 37–750 cr,rb,sb x x x x x x
1889
Ophidiaster alexandri Verrill, 52–585 x x
1915
Ophidiaster bayeri A. H. Clark, intertidal rub x
1948
Ophidiaster guildingii Gray, 0–330 cr,rb x x x x x x x
1840
Ophidiaster ophidianus 0–105 rb x
(Lamarck, 1816)
Tamaria floridae (Perrier, 50–600 rb,sb x x
1881)
Tamaria halperni Downey, 180–510 cr,rb,sb x x x
1971
Family Oreasteridae
Oreaster reticulatus (Linnaeus, 0–1500 cr,m,rb,sb,sg,rub x x x x x x x x x x x x x x x
1758)
Family Poraniidae
Marginaster pectinatus Perrier, 166–450 cr,mb,rb,rub x x x x
1881
Porania (Porania) antarctica 18–320 sb,rub x x
magellanica Studer, 1876
Porania (Porania) pulvillus 35–680 rb x
insignis Verrill, 1895
Poraniopsis echinaster Perrier, 30–420 sb,rub x
1891
Poraniopsis mira (de Loriol, 0–500 x x
1904)
Family Solasteridae
Laetmaster spectabilis (Perrier, 1521 rb x
1881)
Lophaster verrilli A. H. Clark, 6–1100 mb,rb,sb,rub x x x x
1938
(continued)
Appendix
(m)
Appendix
Solaster caribbaeus Verrill, 64–869 mb,rb x x

1915
Order Velatida
Family Caymanostellidae
Caymanostella spinimarginata 3109– mb x
Belyaev, 1978 3493
Family Korethrasteridae
Peribolaster folliculatus 81–133 x x
Sladen, 1889
Remaster gourdoni Koehler, 10–540 x x
1912
Remaster palmatus (Perrier, 296–585 mb,rb,sb x x x x
1881)
Family Myxasteridae
Pythonaster murrayi Sladen, 3477 x
1889
Family Pterasteridae
Calyptraster coa Sladen, 1882 260–930 x
Calyptraster personatus 2151– mb,rb x x x
(Perrier, 1885) 6560
Diplopteraster clarki 82–177 x
Bernasconi, 1937
Diplopteraster verrucosus 74–270 x
(Sladen, 1882)
Hymenaster anomalus Sladen, 1984– rb x
1882 2606
Hymenaster pellucidus 1784– sb x x
Thomson, 1873 3294
Hymenaster pergamentaceus 4846 sb x x
Sladen, 1882
Hymenaster regalis Verrill, 1857 x
1895
Hymenaster rex Perrier, 1885 1139– rb x x
2285
Pteraster abyssorum (Verrill, 576– rb x x
1895) 3740
613
(continued)
614
(m)
Pteraster acicula (Downey, 196– cr,mb x x x
1970) 3713
Pteraster affinis lebruni Perrier, 74–341 x x
1891
Pteraster caribbaeus Perrier, 275–825 mb,sb,rub x x
1881
Pteraster gibber (Sladen, 1882) 27–500 x x
Pteraster militarioides 271–466 mb x x
militarioides H. L. Clark, 1941
Pteraster militarioides stoibe 777 mb x
H. L. Clark, 1941
Pteraster militaris (Müller, 549–594 x
1776)
Pteraster personatus Sladen, 480– mb x
1891 1780
Pteraster rugosus H. L. Clark, 91–466 mb,sb x x x
1941
Pteraster stellifer Sladen, 1882 79–2804 x x
Order Spinulosida
Family Echinasteridae
Echinaster (Echinaster) 60–475 mb,rb,sb x x x x x x x
modestus Perrier, 1881
Echinaster (Echinaster) 2–250 rb,sg,rub
sepositus (Retzius, 1783)
Echinaster (Othilia) 0–360 mb,rb,sb,sg x x x x x x
brasiliensis Müller & Troschel,
1842
Echinaster (Othilia) 0–55 cr,m,mb,rb,sb,rub x x x x x x x x
echinophorus (Lamarck, 1816)
Echinaster (Othilia) guyanensis 13–119 mb,sb x x x x x x x x x x
A. M. Clark, 1987
Echinaster (Othilia) sentus 0–175 cr,rb,sb,sg,rub x x x x x x x x
(Say, 1825)
Echinaster (Othilia) 10–200 cr,mb,sb x x
serpentarius Müller &
Troschel, 1842
(continued)
Appendix
(m)
Appendix
Echinaster (Othilia) spinulosus 1–238 cr,mb,sb,rb x x x x x x

Verrill, 1869
Henricia antillarum (Perrier, 192– sb,rub x x x x x x
1881) 1390
Henricia downeyae A. 342– sb,rub x x
M. Clark, 1987 1037
Henricia obesa (Sladen, 1889) 22–210 sb,rub x x
Henricia sanguinolenta (O. 0–200 x
F. Müller, 1776)
Henricia sexradiata (Perrier, 29–366 sb,rub x x
1881)
Henricia studeri (Perrier, 1891) 74–430 sb,rub x x
Order Forcipulatida
Family Asteriidae
Anasterias antarctica (Lütken, 1–183 rub,rb x x
1857)
Anasterias minuta Perrier, 1875 0–100 x
Anasterias pedicellaris 0–120 x x
(Koehler, 1923)
Anasterias spirabilis (Bell, 34–54 x
1881)
Asterias forbesi (Desor, 1848) 0–619 rb,rub x x x
Coscinasterias linearis (Perrier, 366 rb x
1881)
Coscinasterias tenuispina 0–165 rb,sb,rub x x x x
(Lamarck, 1816)
Cosmasterias lurida (Philippi, 0–650 mb,sb x x
1858)
Diplasterias brandti (Bell, 0–450 sb,mb,rub x x
1881)
Lethasterias australis Fisher, 81–155 x x
1923
Lysasterias perrieri (Studer, 0–320 x
1885)
Marthasterias glacialis 0–180 rb x
(Linnaeus, 1758)
615
(continued)
616
(m)
Neosmilaster steineni (Studer, x x
1885)
Perissasterias polyacantha H. 96–760 mb,sb x
L. Clark, 1923
Psalidaster mordax Fisher, 80–600 x x
1940
Sclerasterias contorta (Perrier, 384–607 x x
1881)
Sclerasterias tanneri (Verrill, x
1895)
Stephanasterias albula 33–2300 cr,sg x x x
(Stimpson, 1853)
Coronaster briareus (Verrill, 50–700 cr,rb x x x x x
1882)
Family Heliasteridae
Labidiaster radiosus Lütken, 5–200 sb,rub x x
1871
Family Pedicellasteridae
Ampheraster alaminos 256– sb x
Downey, 1971 3089
Pedicellaster pourtalesii 338–466 mb x x x
Perrier, 1881
Family Stichasteridae
Allostichaster hartti (Rathbun, 147–380 x
1879)
Smilasterias triremis Sladen, 0–2707 x x
1889
Family Zoroasteridae
Cnemidaster sigsbeei (Perrier, 365–735 mb,sb x x
1894)
Doraster constellatus Downey, 345–914 mb,rb,sb x x x x x x
1970
Mammaster sigsbeei (Perrier, 430–613 mb x x
1880)
Zoroaster fulgens Thomson, 220– mb,rb,sb x x x x x x x
1873 3000
(continued)
Appendix
(m)
Appendix
Order Brisingida
Family Brisingidae
Brisinga costata Verrill, 1884 630– cr,mb,rb x x x
1903
Hymenodiscus coronata 100– mb x
(G. O. Sars, 1872) 2904
Hymenodiscus verticillata 640 rb x
(Sladen, 1889)
Midgardia xandaros Downey, 366–460 sb x x
1972
Novodinia americana (Verrill, 408–576 mb x
1880)
Novodinia antillensis 366– mb,sb x x x
(A. H. Clark, 1934) 2700
Novodinia pandina (Sladen, 54–990 rb x x
1889)
Stegnobrisinga splendens 402–933 x x
H. L. Clark, 1926
Family Freyellidae
Colpaster scutigerula Sladen, 930– mb x x
1889 2790
Freyastera mexicana 2683– rb x
(A. H. Clark, 1939) 5110
Freyastera tuberculata (Sladen, 3360– x
1889) 5620
Freyella elegans (Verrill, 1884) 2928– x
3294
Freyella microspina Verrill, 1848 x
1894
Class Ophiuroidea
Order Euryalida
Family Asteronychidae
Asteronyx loveni Müller & 265– cr,rb,sb x x x x
Troschel, 1842 2499
Astrodia tenuispina (Verrill, 512–935 x x
1884)
617
(continued)
618
(m)
Family Asteroschematidae
Asteroschema arenosum 677– x x
Lyman, 1878 1555
Asteroschema brachiatum 108–783 cr x x
Lyman, 1879
Asteroschema elongatum 42–708 cr,rb,sb x x
Koehler, 1914
Asteroschema intectum Lyman, 238–475 cr,sb x x
1878
Asteroschema laeve (Lyman, 42–539 cr x x x x
1875)
Asteroschema oligactes (Pallas, 124–521 cr x x
1788)
Asteroschema tenue Lyman, 66–180 x
1875
Ophiocreas lumbricus Lyman, 15–600 x x
1869
Ophiocreas oedipus Lyman, 1061– cr x
1879 2228
Ophiocreas spinulosus Lyman, 227–576 cr x x
1883
Family Gorgonocephalidae
Asteroporpa annulata Lütken, 15–397 cr,rb,sb,rub x x x x x x x x
1856
Asteroporpa pulchra H. 320–475 x
L. Clark, 1915
Astracme mucronatus (Lyman, 70–521 cr x x
1869)
Astrocaneum herrerai (H. 0–25 cr x
L. Clark, 1918)
Astrochele lymani Verrill, 1878 0–300 x
Astrocyclus caecilia (Lütken, 20–677 cr,rb,sb x x x
1856)
Astrocnida isidis (Duchassaing, 20–180 cr x x
1850)
(continued)
Appendix
(m)
Appendix
Astrogomphus vallatus Lyman, 60–800 cr,rb x x x

1869
Astropartus mediterraneus 32–265 mb,rb,sb x
(Risso, 1826)
Astrophyton muricatum 0–508 cr,rb,sb,sg,rub x x x x x x x x x x x x x x x x
(Lamarck, 1816)
Astrotoma agassizii Lyman, 74–1000 x x x
1875
Gorgonocephalus chilensis 0–500 x x x x
(Philippi, 1858)
Schizostella bifurcata 12–46 cr x x
A. H. Clark, 1952
Order Ophiurida
Family Ophiomyxidae
Ophioblenna antillensis 1–24 cr,rb x x x x x
Lütken, 1859
Ophiobrachion uncinatus 457 mb x
Lyman, 1883
Ophiobyrsa perrieri Lyman, 527 x
1883
Ophiobyrsa serpens Lyman, 51–126 cr,rb x x x
1883
Astrogeron supinus (Lyman, 530– x
1883) 1143
Ophioleptoplax brasiliana 15–520 x
Tommasi & Abreu, 1974
Ophiomyxa brevicauda Verrill, 23–360 mb x
1899
Ophiomyxa flaccida (Say, 0–1500 cr,rb,sb,sg x x x x x x x x x x x x
1825)
Ophiomyxa pentagona 0–1060 mb,rb,sb,sg,rub x
(Lamarck, 1816)
Ophiomyxa stimpsonii (Lyman, 108–472 mb x
1875)
Ophiomyxa tumida Lyman, 23–601 cr,mb,rb,sb x x x x
1883
(continued)
619
620
(m)
Ophiomyxa vivipara Studer, 0–183 x x x
1876
Ophiophrixus quadrispinosus 265–510 x
(Koehler, 1914)
Ophioprium cervicornis x
(Lyman, 1883)
Ophiosciasma attenuatum 130–540 x
Lyman, 1878
Ophioscolex disacanthus 342–549 rb, mb x x
H. L. Clark, 1915
Ophioscolex glacialis Müller & 50–2727 x
Troschel, 1842
Ophioscolex nutrix (Mortensen, x x x
1936)
Ophioscolex serratus 166 sb x
H. L. Clark, 1900
Ophiosyzygus disacanthus H. 127–278 cr,mb x
L. Clark, 1911
Family Amphilepididae
Amphilepis norvegica 100– mb x
(Ljungman, 1865) 2900
Amphilepis sanmatiensis 1–145 x x
Bernasconi & D’Agostino,
1975
Amphilepis teodorae Tommasi x
& Abreu, 1974
Family Amphiuridae
Amphilimna mirabilis 200–550 mb x x x
(H. L. Clark, 1941)
Amphilimna olivacea (Lyman, 15–600 mb,rb,sb x x x x x x
1869)
Amphiodia atra (Stimpson, 1.5–100 sb x x x
1852)
Amphiodia guillermosoberoni 0.9–3.4 sb x
Caso, 1979
(continued)
Appendix
(m)
Appendix
Amphiodia habilis 34 x
Albuquerque, Campos-Creasey
& Guille, 2001
Amphiodia planispina (von 0–578 mb,sb,sg,rub x x x x x x x
Martens, 1867)
Amphiodia pulchella (Lyman, 0–370 cr,mb,rb,sb,sg x x x x x x x x
1869)
Amphiodia trychna H. L. Clark, 1–160 m,sb,sg,rub x x x x x x x
1918
Amphiodia violacea (Lütken, x
1856)
Amphioplus (Amphioplus) 6–587 mb,sb x x x x
abditus (Verrill, 1871)
Amphioplus (Amphioplus) x
brasiliensis Tommasi, 1970
Amphioplus (Amphioplus) 0–1207 cr,mb,rb,sg x x x
coniortodes H. L. Clark, 1918
Amphioplus (Amphioplus) 0–82 cr,mb,rb,sg x x
sepultus Hendler, 1995
Amphioplus (Amphioplus) 0.3–0.6 mb,sg x
thrombodes H. L. Clark, 1918
Amphioplus (Unioplus) incisus 1639 x x
(Lyman, 1883)
Amphioplus albidus 1–500 mb,rb,sb x x x
(Ljungman, 1867)
Amphioplus daleus (Lyman, x
1879)
Amphioplus lucyae Tommasi, 5–600 sb x x
1971
Amphioplus mathildae 0–120 x
Amphioplus peregrinator x
(Koehler, 1912)
Amphioplus tumidus (Lyman, 70–578 mb x
1878)
(continued)
621
622
(m)
Amphipholis gracillima 0–63 mb,sb x x x
(Stimpson, 1852)
Amphipholis januarii 1–311 cr,mb,rb,sg,rub x x x x x
Ljungman, 1867
Amphipholis pachybactera x
H. L. Clark, 1918
Amphipholis squamata (Delle 0–1962 cr,m,rb,sb,sg,rub x x x x x x x x x x x x x
Chiaje, 1828)
Amphipholis subtilis 0–1330 x
(Ljungman, 1867)
Amphiura (Amphiura) callida 50 x
Albuquerque, Campos-Creasey
& Guille, 2001
Amphiura (Amphiura) correcta x
Koehler, 1907
Amphiura (Amphiura) x
grandisquama Lyman, 1869
Amphiura (Amphiura) 0–400 x x x
magellanica Ljungman, 1867
Amphiura (Amphiura) rosea x
Tommasi & Oliveira, 1976
Amphiura (Ophionema) 1–600 x
intricata Lütken, 1869
Amphiura algida Koehler, 1911 x
Amphiura belgicae Koehler, x x
1900
Amphiura chiajei Forbes, 1843 5–1200 x
Amphiura complanata 0–810 x x
Ljungman, 1867
Amphiura crassipes Ljungman, 0–60 mb,sb x x x
1867
Amphiura deichmanni x
Tommasi, 1965
Amphiura diducta Koehler, x
1914
(continued)
Appendix
(m)
Appendix
Amphiura eugeniae Ljungman, 0–800 mb x x x

1867
Amphiura fibulata Koehler, 2–810 cr,rb,sb,sg,rub x x x x
1913
Amphiura flexuosa Ljungman, 0–50 rub x x
1867
Amphiura iraciae Tommasi & x
Oliveira, 1976
Amphiura joubini Koehler, 5–3834 rub x x x
1912
Amphiura kinbergi Ljungman, 3–300 x
1872
Amphiura latispina Ljungman, 10–50 x x
1867
Amphiura muelleri Marktanner- 134–600 x
Turneretscher, 1887
Amphiura otteri Ljungman, 198– mb,sb x x x x x
1872 3200
Amphiura palmeri Lyman, 5–479 cr,rb x x x x
1882
Amphiura princeps Koehler, 0–107 mb,sb x x x x
1907
Amphiura rathbuni Koehler, 29–502 mb x x
1914
Amphiura scabriuscula 0–26 x x
(Lütken, 1859)
Amphiura semiermis Lyman, 82–1448 mb x x
1869
Amphiura stimpsonii Lütken, 0–2844 cr,rb,sb,rub x x x x x x
1859
Microphiopholis atra 1–38 mb x x x
(Stimpson, 1852)
Microphiopholis gracillima 0–26 mb x x x x
(Stimpson, 1854)
Microphiopholis subtilis x
(Ljungman, 1867)
(continued)
623
624
(m)
Nudamphiura carvalhoi 15–117 x
Tommasi, 1965
Ophiocnida loveni (Ljungman, 7–48 x
1867)
Ophiocnida scabra Lyman, 1100 x
1879
Ophiocnida scabriuscula 0–70 cr,rb,sg x x x
(Lütken, 1859)
Ophionephthys limicola 1–12 cr,m,sb,mb x x x x
Lütken, 1869
Ophiophragmus chilensis 0–112 x x
(Müller & Troschel, 1843)
Ophiophragmus cubanus 1–36 m,mb,sg x x x
(A. H. Clark, 1917)
Ophiophragmus filograneus 0–3 x x
(Lyman, 1875)
Ophiophragmus luetkeni 0–50 x
(Ljungman, 1872)
Ophiophragmus moorei 1–2 rb x
Thomas, 1965
Ophiophragmus pulcher 0.5–33 cr,m,mb,sg,rub x x x x x x x
H. L. Clark, 1918
Ophiophragmus riisei (Lütken, 1–311 cr,mb,sb x x x x x x
1859)
Ophiophragmus septus 0.3–116 cr,m,mb,sb x x x x x x x
(Lütken, 1859)
Ophiophragmus wurdemani 2–11 cr,sb x x x
(Lyman, 1860)
Ophiostigma isocanthum (Say, 0–223 cr,mb,sg,sb,rub x x x x x x x x x
1825)
Ophiostigma siva Hendler, 0.5–99 cr,rb x x x x
1995
Family Hemieuryalidae
Hemieuryale pustulata Von 128–148 x x x
Martens, 1867
(continued)
Appendix
(m)
Appendix
Ophiochondrella squamosa 604 x

(Lyman, 1883)
Ophiochondrus convolutus 165–720 cr x x
Lyman, 1869
Ophiochondrus crassispinus 420 x
Lyman, 1883
Ophiochondrus gracilis Verrill, 198–468 x
1899
Ophiochondrus stelliger 144– x x
Lyman, 1879 1080
Sigsbeia conifera Koehler, 4–450 mb x x x x
1914
Sigsbeia murrhina Lyman, 16–706 cr,mb x x x x
1878
Family Ophiacanthidae
Subfamily Ophiacanthinae
Ophiacantha abyssicola 35–351 mb,rub x
G. O. Sars, 1871
Ophiacantha affinis Koehler, 366–420 x
1914
Ophiacantha aristata Koehler, 822– x
1895 1700
Ophiacantha aspera Lyman, 320 x
1878
Ophiacantha bidentata 32–4730 cr,mb,rb x x x
(Retzius, 1805)
Ophiacantha brasiliensis 145–380 x
Ophiacantha cervicornis 366–539 x
Lyman, 1883
Ophiacantha crassidens Verrill, 980– mb x
1885 3120
Ophiacantha densispina x
Mortensen, 1936
Ophiacantha echinulata 304– mb,rb,sb x x x
Lyman, 1878 1958
(continued)
625
626
(m)
Ophiacantha ensifera (Verrill, 198–468 x
1899)
Ophiacantha mesembria 252– mb x x
H. L. Clark, 1915 1125
Ophiacantha metallacta 295–763 mb x x
H. L. Clark, 1915
Ophiacantha pentacrinus 320– x x x
Lütken, 1869 1249
Ophiacantha rosea Lyman, 402– mb,rb x
1878 1538
Ophiacantha scutata Lyman, 521–549 x
1878
Ophiacantha sertata (Lyman, 183–618 cr,rb x
1869)
Ophiacantha stellata Lyman, 183 x
1875
Ophiacantha setosa (Retzius, 40–1480 sb x
1805)
Ophiacantha vivipara 0–1097 x x x x
Ljungman, 1870
Ophiacanthella troscheli 185–238 cr,rb,sb x
(Lyman, 1878)
Ophialcaea nuttingii Verrill, 366 x
1899
Ophiotreta lineolata (Lyman, 347–521 x
1883)
Ophiotreta mixta (Lyman, 338–442 x
1878)
Ophiotreta sertata (Lyman, 293–567 mb x x
1869)
Ophiotreta valenciennesi 200– x
valenciennesi (Lyman, 1879) 1440
Ophiotreta valenciennesi 155– mb x
rufescens (Koehler, 1896) 1440
(continued)
Appendix
(m)
Appendix
Subfamily Ophiohelinae
Ophiomyces frutectosus 151–410 mb x
Lyman, 1869
Ophiomyces mirabilis Lyman, 270–560 rb x
1869
Subfamily Ophioplinthacinae
Ophiocamax austera Verrill, 198– x
1899 1045
Ophiocamax fasciculata 208–979 mb,rb,sb x x x x x x x
Lyman, 1883
Ophiocamax hystrix Lyman, 171–695 mb x x
1878
Ophiomitra ornata Verrill, 201 x
1899
Ophiomitra robusta Koehler, 395 x
1914
Ophiomitra valida Lyman, 131–608 cr x x
1869
Ophiomitrella clavigera 250– x
(Ljungman, 1865) 1500
Ophiomitrella conferta x x
(Koehler, 1922)
Ophiomitrella cordifera x
Koehler, 1909
Ophiomitrella glabra (H. cr x x
L. Clark, 1901)
Ophiomitrella ingrata Koehler, x
1908
Ophiomitrella laevipellis 155–507 cr x x
(Lyman, 1883)
Ophioplinthaca carduus 796 mb x
(Lyman, 1878)
Ophioplinthaca chelys 795– x x
(Wyville-Thomson, 1878) 3305
Ophioplinthaca dipsacos 777 x
(Lyman, 1878)
(continued)
627
628
(m)
Ophiothamnus exigua (Lyman, 439 x
1878)
Subfamily Ophiotominae
Ophiolimna littoralis Koehler, 0–63 x
1913
Ophiopristis hirsuta (Lyman, 147–830 cr x
1875)
Ophiotoma gracilis (Koehler, 490– mb x
1914) 1236
Family Ophiactidae x
Hemipholis cordifera Bosc, 0–520 cr,mb,rb,sb,rub x x x x x x
1802
Histampica duplicata (Lyman, 125– cr,mb x x x
1875) 2870
Histampica rugosa (H. 1125– x
L. Clark, 1941) 1281
Ophiactis abyssicola (M. Sars, 125– mb,rb x
1861) 4000
Ophiactis algicola H. L. Clark, 0–24 cr x x x
1933
Ophiactis asperula (Philippi, 0–310 x x x
1858)
Ophiactis balli (W. Thompson, 30–1765 rb x
1840)
Ophiactis brasiliensis Manso, 1.5–163 x
1988
Ophiactis dispar Verrill, 1899 62 x
Ophiactis ljungmani x
Marktanner-Turneretscher,
1887
Ophiactis loricata Lyman, 198 mb x
1869
Ophiactis lymani Ljungman, 0–600 rb x
1872
Ophiactis muelleri Lütken, 14–67 sb x x x x
1856
(continued)
Appendix
(m)
Appendix
Ophiactis quinqueradia 0–640 cr,mb,rb,sb,sg x x x x x x x x

Ljungman, 1871
Ophiactis rubropoda 2–32 cr x
Singletary, 1973
Ophiactis savignyi (Müller & 0–1500 cr,m,rb,sb,sg,rub x x x x x x x x x x x x x
Troschel, 1842)
Ophiactis virens (M. Sars, 0–90 rb x
1857)
Family Ophiochitonidae
Ophiochiton ternispinus 377– mb,rb x x x x
Lyman, 1883 3550
Ophioplax clarimundae x
Tommasi, 1970
Ophioplax ljungmani Lyman, 22–471 cr x x x
1875
Ophioplax pardalis H. L. Clark, 347 x
1941
Ophioplax spinulifera H. 274–420 x
L. Clark, 1941
Family Ophiocomidae
Ophiocoma echinata (Lamarck, 0–183 cr,m,rb,sb,sg,rub x x x x x x x x x x x x x x
1816)
Ophiocoma paucigranulata 0–455 cr,rb x x x
Devaney, 1974
Ophiocoma pumila Lütken, 0–368 cr,rb,sg x x x x x x x x x x x x
1859
Ophiocoma riisei Lütken, 1859 1–2 cr x x
Ophiocoma wendtii Müller & 0–384 cr,m,rb,sg,rub x x x x x x x x x x x
Troschel, 1842
Ophiocomella ophiactoides (H. 0–70 cr,rb,sb,sg,rub x x x x x x x
L. Clark, 1901)
Ophiocomella sexradia x
(Duncan, 1887)
Ophiocomina nigra 0–400 sb,rub x
(Abildgaard, in O.F. Müller,
1789)
(continued)
629
630
(m)
Ophiopsila aranea Forbes, 0–185 sg,rub x
1843
Ophiopsila fulva Lyman, 1878 29–320 cr x x
Ophiopsila guineensis Koehler, 18–110 x
1914
Ophiopsila hartmeyeri Koehler, 1–183 cr,rb,sb x x x x x x x
1913
Ophiopsila maculata (Verrill, 41–3000 x x
1899)
Ophiopsila riisei Lütken, 1859 0–366 cr,m,rb,sg x x x x x x x x x
Ophiopsila vittata H. L. Clark, 4–15 cr,rb x x x
1918
Family Ophiodermatidae
Bathypectinura heros (Lyman, 276– mb,sb x x x x x x x
1879) 3150
Ophiorachnella angulata x
(Lyman, 1883)
Ophiarachnella petersi 320–475 x
(Lyman, 1878)
Ophioconis forbesi (Heller, 20–200 rb x
1863)
Ophioconis miliaria Lyman, 444–759 x
1878
Ophioderma anitae Hotchkiss, cr,sb x
1982
Ophioderma appressa (Say, 0–580 cr,rb,sg,rub x x x x x x x x x x x x x
1825)
Ophioderma besnardi 0–600 x
Tommasi, 1970
Ophioderma brevicaudum 0–64 cr,rb,sb,sg x x x x x x x x x
Lütken, 1856
Ophioderma brevispinum (Say, 1–223 cr,m,mb,rb,sb,sg,rub x x x x x x x x x x x
1825)
Ophioderma cinereum Müller 0–1719 cr,m,sb,sg x x x x x x x x x x x x
& Troschel, 1842
(continued)
Appendix
(m)
Appendix
Ophioderma divae Tommasi, x

1971
Ophioderma ensiferum Hendler 10–30 cr,rb x x x
& Miller, 1984
Ophioderma guttatum Lütken, 0–30 cr,rb,rub x x x x
1859
Ophioderma januarii Lütken, 0–1500 cr x x
1856
Ophioderma longicaudum 0–200 sg,rub x
(Retzius, 1805)
Ophioderma phoenium H. 1–14 cr,rb,sb x x x x x
L. Clark, 1918
Ophioderma rubicundum 0–360 cr,sg,rb,sb,rub x x x x x x x x
Lütken, 1856
Ophioderma squamosissimum 3–85 cr,rb x x x x
Lütken, 1856
Ophiopaepale goesiana 101–436 cr,mb,sb x x x
Ljungman, 1872
Ophiurochaeta littoralis 1–110 cr,sb x
(Koehler, 1913)
Family Ophionereididae
Ophionereis dolabriformis John 10–93 cr,mb x x
& A. M. Clark, 1954
Ophionereis olivacea H. 0–500 cr,m,sb x x x x x x x x
L. Clark, 1901
Ophionereis reticulata (Say, 0–1500 cr,m,rb,sb,sg,rub x x x x x x x x x x x x
1825)
Ophionereis sexradia 18–130 x
Mortensen, 1936
Ophionereis squamulosa 0.5–110 cr,sb,rb,sg,rub x x x x x x x
Koehler, 1913
Ophionereis vittata Hendler, 10–126 cr x x x x
1995
Family Ophiotrichidae
Ophiothela danae Verrill, 1869 x
(continued)
631
632
(m)
Ophiotrichoides lymani x
Ludwig, 1882
Ophiothrix (Ophiothrix) ailsae x
Tommasi, 1970
Ophiothrix (Ophiothrix) x
trindadensis Tommasi, 1970
Ophiothrix angulata (Say, 0.5–540 cr,m,mb,rb,sb,sg,rub x x x x x x x x x x x
1825)
Ophiothrix angulata violacea x
(Müller & Troschel, 1842)
Ophiothrix brachyactis 1–6 cr,rb,rub x x x x x
H. L. Clark, 1915
Ophiothrix cimar Hendler, 0–10 cr,sg x x
2005
Ophiothrix fragilis 0–1250 rb,sg,rub x
(Abildgaard, 1789)
Ophiothrix lineata Lyman, 0–74 cr,rb x x x x x x
1860
Ophiothrix luetkeni Wyville- 50–500 rb,sb,rub x
Thomson, 1873
Ophiothrix maculata 114–410 x
Ljungman, 1872
Ophiothrix oerstedii Lütken, 0–31 cr,sb,mb,sg,rb,rub x x x x x x x x x
1856
Ophiothrix pallida Ljungman, 183 x
1871
Ophiothrix platyactis 41 rb x
H. L. Clark, 1939
Ophiothrix rathbuni Ludwig, 8–600 x
1882
Ophiothrix stri Hendler, 2005 0–10 cr x x
Ophiothrix suensonii Lütken, 0–1000 cr,m,mb,rb,sb x x x x x x x x x x x
1856
Ophiothrix synoecina Schoppe, 0–5 rb x
1996
(continued)
Appendix
(m)
Appendix
Family Ophiuridae
Subfamily Ophiolepidinae
Amphipholizona delicata H. 15–600 mb x x
L. Clark, 1915
Ophiolepis ailsae Hendler & 156–353 x
Turner, 1987
Ophiolepis elegans Lütkens, 1–329 cr,m,mb,rb,sb,sg,rub x x x x x x x x
1859
Ophiolepis gemma Hendler & 2–139 cr,rb,sb x x x
Turner, 1987
Ophiolepis impressa Lütken, 0–1500 cr,rb,sb,sg,rub x x x x x x x x x x x
1859
Ophiolepis kieri Hendler, 1979 2–8 sg x x x
Ophiolepis paucispina (Say, 0–37 cr,m,rb,sb,sg,rub x x x x x x x x x
1825)
Ophiolepis pawsoni Hendler, 24.4 x
1988
Ophiolipus agassizii Lyman, 146–310 rb x
1878
Ophiomidas dubius (Lyman, 158–272 x
1878)
Ophiomusium acuferum 76–575 cr,mb,rb x x x x
Lyman, 1875
Ophiomusium anaelisae 180–260 x
Ophiomusium constrictum x
Mortensen, 1936
Ophiomusium eburneum 35–3477 cr,mb,sb x x x x
Lyman, 1869
Ophiomusium leptobrachium 1249– sb x x
H. L. Clark, 1941 1518
Ophiomusium lütkeni Lyman, 190–278 rb,sb x
1878
Ophiomusium lymani Thomson, 62–4700 mb,rb,sb,rub x x x x
1873
(continued)
633
634
(m)
Ophiomusium microporum H. 347–530 x
L. Clark, 1941
Ophiomusium moniliforme H. 1125 x
L. Clark, 1941
Ophiomusium planum Lyman, 368– rb,sb x
1879 4062
Ophiomusium sculptum Verrill, 475 x
1899
Ophiomusium serratum Lyman, 520– rb,sb x x
1878 3103
Ophiomusium stellatum Verrill, 356–521 x
1899
Ophiomusium testudo Lyman, 60–914 cr,rb,sb x x x
1875
Ophiomusium validum 108– cr,mb x x x
Ljungman, 1872 2732
Ophioplocus januarii (Lütken, 0–180 rb,sb,rub x x x
1856)
Ophiosphalma armigerum 262– rb,sb x x x
(Lyman, 1878) 4024
Ophiosphalma dyscritum 512–713 x
H. L. Clark, 1941
Ophiosphalma monoplax 376–746 x
H. L. Clark, 1915
Ophiothyreus goesii Ljungman, 144–540 cr x x x
1872
Ophiozonella clypeata (Lyman, 158–272 x
1883)
Ophiozonella falklandica x x
Mortensen, 1936
Ophiozonella granulifera 1006– rb,sb x x
H. L. Clark, 1941 1098
Ophiozonella marmorea 484– x
Lyman, 1883 1385
Ophiozonella molesta (Koehler, 2115 x
1904)
(continued)
Appendix
(m)
Appendix
Ophiozonella nivea (Lyman, 70–1249 rb,sb x x x

1875)
Ophiozonella tessellata 320–475 x
(Lyman, 1879)
Subfamily Ophioleucinae
Ophiostriatus striatus 270– x
(Mortensen, 1933) 3500
Ophiernus adspersus adspersus 68–3650 mb,rb x x x
Lyman, 1883
Ophioleuce depressum (Lyman, 585 x
1869)
Ophiopyren longispinus 329 x
Lyman, 1878
Ophiostriatus atlanticus 296–396 mb x
(Mortensen, 1933)
Subfamily Ophiurinae
Amphiophiura metabula 274– mb,sb x x x
H. L. Clark, 1915 1830
Amphiophiura oedignatha 284–944 mb x x x
H. L. Clark, 1915
Amphiophiura ornata (Lyman, x
1878)
Anthophiura ingolfi Fasmer, 21–1200 cr x
1930
Ophiambix devaneyi Paterson, 146–494 mb x
1985
Ophiocten amitinum Lyman, 46–545 x x x
1878
Ophiomastus meridionalis x
(Lyman, 1879)
Ophiomastus satelitae 115–600 x
Ophiomastus secundus Lyman, 108– mb,sb x x x
1878 2035
Ophiomisidium pulchellum 11–3061 cr,rb,sb x x x
(Wyville-Thomson, 1878)
(continued)
635
636
(m)
Ophiomisidium speciosum 880– mb x x
Koehler, 1914 1636
Ophiomisidium tommasii 250–808 x
Borges, Monteiro & Amaral,
2006
Ophiopleura inermis (Lyman, 1786 x
1878)
Ophioplinthus abyssorum 338–366 x
(Lyman, 1883)
Ophioplinthus inornata 54–3384 x x
(Lyman, 1878)
Ophioplinthus martensi x
(Studer, 1885)
Ophiophycis mirabilis Koehler, x
1901
Ophiura acervata (Lyman, 20–1900 cr,mb,rb,sb x x x x x x
1869)
Ophiura carinifera (Koehler, 400 x
1901)
Ophiura carnea Lütken, 1858 40–1260 mb,sb x
Ophiura clemens (Koehler, x
1904)
Ophiura falcifera (Lyman, 73–1037 mb x x
1869)
Ophiura fallax Cherbonnier, 20–842 mb,sb x x
1959
Ophiura grubei Heller, 1863 10–350 mb,sb,rub x
Ophiura (Ophiuroglypha) 600– mb,sb x
irrorata concreta (Koehler, 4315
1901)
Ophiura (Ophiuroglypha) x
irrorata irrorata (Lyman,
1878)
Ophiura ljungmani (Lyman, 46–6398 mb,rb,sb x x x
1878)
Ophiura lymani (Ljungman, 0–463 x x x
1871)
Appendix
(continued)
(m)
Appendix
Ophiura ophiura (Linnaeus, 0–300 mb,sb x

1758)
Ophiura sarsi (Lütken, 1855) 82–827 mb x
Ophiura scomba Paterson, 1985 358– rb,sb x
3943
Ophiura tenera (Lyman, 1883) 158–500 mb x
Ophiuraster perissus H.L. x
Clark, 1939
Class Echinoidea
Order Cidaroida
Family Cidaridae
Austrocidaris canaliculata 1–424 x x
(A. Agassiz, 1863)
Austrocidaris lorioli 160– x
(Mortensen, 1903) 1081
Austrocidaris spinulosa 124–641 x x
Mortensen, 1910
Calocidaris micans 33–624 mb,rb,sb x x
(Mortensen, 1903)
Cidaris abyssicola (A. Agassiz, 225–375 mb,rb,sb x
1869)
Cidaris blakei (A. Agassiz, 270–720 mb,rb x x
1878)
Cidaris cidaris (Linnaeus, 50–2000 mb x x
1758)
Cidaris rugosa (H. L. Clark, 40–540 mb,rb,sb x x x x
1907)
Eucidaris tribuloides 0–1500 cr,rb,sb,sg,rub x x x x x x x x x x x x x x
tribuloides (Lamarck, 1816)
Histocidaris nuttingi 225–740 mb,rb,sb x x
Mortensen, 1926
Histocidaris purpurata 300– x
(Wyville-Thomson, 1872) 1800
Histocidaris sharreri 200 – rb,sb x x
(A. Agassiz, 1880) 740
(continued)
637
638
(m)
Stereocidaris ingolfiana 300– mb,rb,sb x x
Mortensen, 1903 1745
Stylocidaris affinis (Philippi, 22–1000 cr,mb,rb,sb,rub x x x x x x x x
1845)
Stylocidaris lineata Mortensen, 20–1717 cr,mb,rb,sb,rub x x x x x x x x
1910
Tretocidaris bartletti 48–1089 mb,rb,sb x x x x x x x
(A. Agassiz, 1880)
Family Rhabdicidaridae
Porocidaris purpurata 640–782 rb,sb x x
(Wyville-Thomson, 1872)
Order Echinothurioida
Family Echinothuriidae
Araeosoma belli Mortensen, 130– mb,sb x x x x x x
1903 1020
Araeosoma fenestratum 16–1180 mb,rb,sb x x x x x
(Thomson, 1872)
Calveriosoma hystrix (Wyville- 360– mb,sb x x x
Thomson, 1872) 2545
Hygrosoma petersii 200– mb,sb x x x
(A. Agassiz, 1880) 3700
Phormosoma placenta placenta 50–3700 mb,sb x x x x x x x x
Thomson, 1872
Phormosoma placenta sigsbei 200– mb x x
A. Agassiz, 1880 1800
Sperosoma grimaldii Koehler, 300– x
1897 2300
Order Diadematoida
Family Aspidodiadematidae
Aspidodiadema jacobyi 170–720 cr,mb,rb x x x x x x
A. Agassiz, 1880
Aspidodiadema tonsum 180–925 x
A. Agassiz, 1879
Plesiodiadema antillarum 684– mb,rb,sb x x x x x x
(A. Agassiz, 1880) 3111
Family Diadematidae
Astropyga magnifica 5–89 cr,rb,sb,sg,rub x x x x x x x
Appendix
A. H. Clark, 1934
(continued)
(m)
Appendix
Astropyga nuptialis Tommasi, 1–20 x

1958
Centrostephanus besnardi x
Bernasconi, 1955
Centrostephanus longispinus 10–1000 cr,mb,rb,sb,rub x x x
(Philippi, 1845)
Centrostephanus longispinus 33–842 sb,rub x x x
rubicingulus H. L. Clark, 1921
Diadema antillarum antillarum 0–800 cr,m,rb,sb,sg,rub x x x x x x x x x x x x x x x
(Philippi, 1845)
Diadema ascensionis 1–400 x
Mortensen, 1909
Order Pedinoida
Family Pedinidae
Caenopedina cubensis 220– mb,rb,sb x x x x
A. Agassiz, 1869 1200
Order Salenioida
Family Saleniidae
Salenia goesiana Lovén, 1874 90–842 rb x x
Salenocidaris profundi 200– mb,rb,sb x x x
(Duncan, 1877) 3700
Salenocidaris varispina 250– mb,rb,sb x x x x x x
A. Agassiz, 1869 3015
Order Arbacioida
Family Arbaciidae
Arbacia dufresnii (Blainville, 0–340 rb,rub x x x
1825)
Arbacia lixula (Linnaeus, 1758) 0–50 rb,sb x x
Arbacia punctulata (Lamarck, 0–225 cr,rb,sb,sg,rub x x x x x x x
1816)
Coelopleurus floridanus 60–2380 cr,mb,rb,sb x x x x x x x x
A. Agassiz, 1871
Habrocidaris scutata 920– x
(A. Agassiz, 1880) 1400
Podocidaris sculpta 8–780 cr,mb,rb,sb x x
A. Agassiz, 1869
639
(continued)
640
(m)
Order Camarodonta
Family Parechinidae
Loxechinus albus (Molina, 30–340 rb,rub x x
1782)
Paracentrotus gaimardi 1–5 x
(Blainville, 1825)
Paracentrotus lividus 0–80 rb,sg x
(Lamarck, 1816)
Psammechinus x
microtuberculatus (Blainville,
1825)
Order Temnopleuroida
Family Temnopleuridae
Genocidaris maculata 12–500 cr,mb,rb,sb x x x x
A. Agassiz, 1869
Pseudechinus magellanicus 1–361 rb,rub x x x
(Philippi, 1857)
Trigonocidaris albida 70–720 cr,rb,sb x x x
A. Agassiz, 1869
Family Toxopneustidae
Lytechinus callipeplus 22–357 rb,sb x x
H. L. Clark, 1912
Lytechinus euerces 40–777 mb,rb,sb x x x x
H. L. Clark, 1912
Lytechinus variegatus carolinus 0–250 rb,sb,rub,sg x
A. Agassiz, 1863
Lytechinus variegatus 0–580 cr,mb,rb,sb,sg,rub x x x x x x x x x x x x x
variegatus (Lamarck, 1816)
Lytechinus williamsi Chesher, 3–256 cr,rb,sb,sg,rub x x x x x x x
1968
Pseudoboletia occidentalis 69–155 x
H. L. Clark, 1921
Sphaerechinus granularis 3–100 rb x
(Lamarck, 1816)
Toxopneustes pileolus 0–35 cr,mb,rb,sb x
(Lamarck, 1816)
(continued)
Appendix
(m)
Appendix
Tripneustes ventricosus 0–842 cr,rb,sb,sg,rub x x x x x x x x x x x x

(Lamarck, 1816)
Order Echinoida
Family Echinidae
Echinus esculentus Linneaeus, 10–289 rb x
1758
Echinus tylodes H. L. Clark, 350–760 sb x
1912
Gracilechinus gracilis 69–457 mb,rb,sb x x x
(A. Agassiz, 1869)
Sterechinus agassizii 24–470 x x
Mortensen, 1910
Sterechinus neumayeri 13–250 x
(Meissner, 1900)
Family Echinometridae
Echinometra lucunter lucunter 0–45 cr,rb,sg,rub x x x x x x x x x x x x x x
(Linnaeus, 1758)
Echinometra viridis A. Agassiz, 0–40 cr,m,rb,sb,sg,rub x x x x x x x x x x x x
1863
Order Holectypoida
Family Echinoneidae
Echinoneus cyclostomus Leske, 0.5–585 cr,mb,rb,sb,sg,rub x x x x x x x x x x
1778
Order Clypeasteroida
Family Clypeasteridae
Clypeaster aloysioi (Brito, x
1959)
Clypeaster chesheri Serafy, 18–101 cr,mb,sb x x x x x
1970
Clypeaster cyclopilus 69–460 sb x x
H. L. Clark, 1941
Clypeaster euclastus 36–530 cr,mb,sb x x x x x x
H. L. Clark, 1941
Clypeaster lamprus 78–450 cr,mb,sb x x x x x x
H. L. Clark, 1914
(continued)
641
642
(m)
Clypeaster luetkeni Mortensen, 9–55 sb x x
1948
Clypeaster oliveirai Krau, 1952 50 sb x
Clypeaster prostratus Ravenel, 15–75 sb x x x x
1848
Clypeaster ravenelii 20–278 mb,sb x x x
A. Agassiz, 1869
Clypeaster rosaceus (Linnaeus, 0–285 sb,sg x x x x x x x x x x x
1758)
Clypeaster speciosus Verrill, x
1870
Clypeaster subdepressus (Gray, 1–210 mb,rb,sb,sg,rub x x x x x x x x
1825)
Family Echinocyamidae
Echinocyamus grandiporus 99–2500 mb,sb x x x x
Mortensen, 1907
Echinocyamus macrostomus 170– sb x x
Mortensen, 1907 2286
Echinocyamus pusillus (Müller, 0–1250 sb,rub x
1776)
Family Mellitidae
Encope aberrans Martens, 13–32 sb x
1867
Encope emarginata (Leske, 0–60 mb,sb x x x x x x x x
1778)
Encope michelini L. Agassiz, 0–240 sb,sg x x x
1841
Leodia sexiesperforata (Leske, 0–180 sb,sg,rub x x x x x x x x x x x x
1778)
Mellita quinquiesperforata 0–180 mb,sb,rub x x x x x x x x x x x
(Leske, 1778)
Mellita quinquiesperforata 0–10 sb x
latiambulacra (H. L. Clark,
1840)
Mellita tenuis H. L. Clark, 1940 0–3 sb,rub x
Order Cassiduloida
Family Cassidulidae
Appendix
(continued)
(m)
Appendix
Cassidulus caribaearum 0.5–197 sb x x x x

Lamarck, 1801
Cassidulus infidus Mortensen, 3.5 x
1948
Cassidulus mitis Krau, 1954 1–45 x
Eurhodia relicta Mooi, 1990 69–155 x
Family Echinolampadidae
Conolampas sigsbei 130–800 mb,rb,sb x x x x x
(A. Agassiz, 1878)
Echinolampas depressa Gray, 37–366 sb x x x
1851
Order Holasteroida
Family Pourtalesiidae
Pourtalesia miranda 450– x
A. Agassiz, 1869 5850
Order Spatangoida
Family Asterostomatidae
Archaeopneustes hystrix 22–1610 mb x x x x
(A. Agassiz, 1880)
Family Brissidae
Anabrissus damesi (A. Agassiz, 640 x
1881)
Brissopsis alta Mortensen, 181–329 mb,sb x
1907
Brissopsis atlantica Mortensen, 18–641 mb,sb x x x x x x x x x
1907
Brissopsis atlantica 37–3200 mb x x x x
mediterranea Mortensen, 1913
Brissopsis elongata elongata 3–270 mb,sb x x x x x x x x
Mortensen, 1907
Brissus unicolor (Leske, 1778) 0–250 mb,rb,sb,rub x x x x x x x x x x x x
Linopneustes longispinus 55–720 mb x x x x x
(A. Agassiz, 1878)
Meoma ventricosa ventricosa 0.3–293 cr,mb,sb,rb,sg,rub x x x x x x x x x x x x x x
(Lamarck, 1816)
Neopneustes micrasteroides 148–330 x
(A. Agassiz, 1878)
643
(continued)
644
(m)
Palaeobrissus hilgardi 150– mb x x
A. Agassiz, 1880 1025
Paleopneustes cristatus 69–805 cr,mb,rb,sb x x x x x
A. Agassiz, 1873
Paleopneustes josephinae 350–436 x
Lovén, 1872
Paleopneustes tholoformis 90–525 cr,mb x x
Chesher, 1968
Plagiobrissus grandis (Gmelin, 1–210 sb,sg x x x x x x x x x
1788)
Plethotaenia angularis 567–570 mb,sb x
Chesher, 1968
Plethotaenia spatangoides 117–619 mb,sb x x x
(A. Agassiz, 1883)
Rhabdobrissus costae (Gasco, 25–200 mb,sb x
1876)
Rhynobrissus cuneus Cooke, 0–10 sb x
1957
Family Hemiasteridae
Holanthus expergitus (Lovén, 380– mb,sb x x x
1874) 4833
Family Loveniidae
Echinocardium cordatum 0–230 sb x x
(Pennant, 1777)
Family Maretiidae
Homolampas fragilis 350– mb,sb x x x x x
(A. Agassiz, 1869) 3550
Homolampas lovenioides 910 x
Mortensen, 1948
Family Prenasteridae
Tripylus excavatus Philippi, 50–113 x x
1845
Family Schizasteridae
Abatus agassizii (Pfeffer, 1889) 75–970 x
Abatus cavernosus (Philippi, 1–676 mb,sb x x
1845)
Abatus philippii Lovén, 1871 71–225 x x
Appendix
(continued)
(m)
Appendix
Aceste bellidifera Thomson, 116– sb x x x

1877 5220
Agassizia excentrica 27–900 cr,mb,sb x x x x x x x x x x x
A. Agassiz, 1869
Brisaster fragilis (Düben & x
Koren, 1846)
Brisaster moseleyi (A. Agassiz, 401– x
1881) 1100
Hypselaster limicolus 27–340 mb,sb x x x
(A. Agassiz, 1878)
Moira atropos clotho Michelin, 0–445 mb,rb,sb x x x x x x x x x x
1855
Ova canaliferus (Lamarck, 9–100 mb,sb x
1816)
Tripylaster philippii (Gray, 13–595 x x
1851)
Schizaster doederleini 9–220 mb,sb x x x x x x x x
(Chesher, 1972)
Schizaster floridiensis 12–65 sb x x x x x x x
(Chesher, 1972)
Schizaster orbignyanus 22–500 mb,sb x x x x
A. Agassiz, 1880
Family Spatangidae
Spatangus purpureus (Müller, 15–900 mb,sb,rub x
1776)
Class Holothuroidea
Order Dendrochirotida
Family Cucumariidae
Aslia lefevrei (Barrois, 1882) 6–25 rb x
Cladodactyla crocea (Lesson, 0–4300 mb,rb,sb,rub x x
1830)
surinamensis (Semper, 0–5 rb,sg x x x x x x x
1868)
Cucumaria manoelina x
Tommasi, 1971
(continued)
645
646
(m)
Duasmodactyla seguroensis 0–50 cr,rb,sb,sg x x x x
(Deichmann, 1930)
Euthyonacta solida 1.5–183 rb,sb x x
(Deichmann, 1930)
Hemioedema spectabilis 2–160 mb,sb x
(Ludwig, 1882)
Heterocucumis godeffroyi 0–379 sb,rub x
(Semper, 1868)
Heterocucumis steineni 0–400 rb x x
(Ludwig, 1898)
Leptopentacta deichmannae 17–37 mb,sb,rub x x
Domantay, 1958
Neocnus incubans Cherbonnier, 0–5 rb x
1972
Ocnus braziliensis (Verrill, 0–2 x
1868)
Ocnus pygmaeus (Théel, 1886) 0–72 cr,rb,sb,rub x x x x
Ocnus suspectus (Ludwig, 0–1874 rb,sg x x x x x x x
1874)
Paracolochirus mysticus 69–155 x
(Deichmann, 1930)
Pawsonia saxicola (Brady & 4–130 rb x
Robertson, 1871)
Pentacta pygmaea (Théel, 0.6–1.7 x x
1886)
Pseudocnus dubiosus leoninus 0–340 mb,rb,sb x x
(Semper, 1868)
Pseudocnus cornutus 135–189 x x
(Cherbonnier, 1941)
Pseudocnus perrieri (Ekman, 0–197 mb,rb,sb x x x
1927)
Pseudrotasfer microincubator 271–290 x
Bohn, 2007
Stereoderma unisemita 31.1– x
(Stimpson, 1851) 40.2
Thyonella gemmata (Pourtalès, 0–64 cr,rb,sb,mb,sg x x x
1851)
Appendix
(continued)
(m)
Appendix
Thyonella pervicax (Théel, 9–67 rb,sb x x x

1886)
Thyonella sabanillaensis 2–30 sb x x x
(Deichmann, 1930)
Trachythyone crassipeda x
Cherbonnier, 1961
Trachythyone lechleri 0–30 x
(Lampert, 1885)
Trachythyone parva (Ludwig, 0–180 mb,rb,sb x x
1874)
Trachythyone peruana 90–159 x x
(Semper, 1868)
Family Phyllophoridae
Havelockia inermis (Heller, 0.5–60 x x
1868)
Neothyonidium parvum 0–3 sg x x x x
(Ludwig, 1881)
Pentamera chiloensis (Ludwig, 0–111 sb,rub x
1887)
Pentamera pulcherrima Ayres, 0–60 mb x x x
1854
Phyllophorus (Urodemella) 0–99 cr,rb,sg x x x x x x
occidentalis (Ludwig, 1875)
Stolus cognatus (Lampert, 0–38 mb,rb,sb,sg x x x x x x
1885)
Thyone cognata (Lampert, x x
1885)
Thyone deichmannae Madsen, 6–366 sg x
1941
Thyone fusus (Müller, 1776) 5–615 mb,sb,rub x
Thyone montoucheti Tommasi, x
1971
Thyone pawsoni Tommasi, 6–51 sb x x x
1972
Thyone pseudofusus 0.5–46 rb,sb,sg,rub x x x x x x
Deichmann, 1930
Thyone tanyspeira Pawson & 51–170 mb x x
Miller, 1988
647
Family Psolidae
(continued)
648
(m)
Lissothuria antillensis Pawson, 1–17 cr,rb x x x
1967
Lissothuria braziliensis (Théel, 0–2 x x
1886)
Neopsolidium convergens 0–20 rb x x
(Herouard, 1901)
Psolidium dorsipes Ludwig, 0–483 rb x x
1886
Psolus antarcticus Philippi, 35–1080 rb x
1857
Psolus marcusi Tommasi, 1971 95 rb x
Psolus patagonicus Ekman, 0–430 rb x x x
1925
Psolus squamatus (Koren, 7–207 rb,rub x x
1844)
Psolus tuberculosus Théel, 73–243 x x
1886
Psolus victoriae Tommasi, x
1971
Family Sclerodactylidae
Euthyonidiella dentata x x
Cherbonnier, 1961
Euthyonidiella destichada 0–7 mb,rb,sb,sg x x x x x
(Deichmann, 1930)
Euthyonidiella trita (Sluiter, 0–100 rb x x x
1910)
Pseudothyone belli (Ludwig, 0–37 mb,rb,sb,sg x x x x x x
1886)
Sclerodactyla briareus 15 x
(Lesueur, 1824)
Order Dactylochirotida
Family Ypsilothuriidae
Echinocucumis asperrima 43–723 x
(Théel, 1886)
Echinocucumis hispida (Barret, 50–1000 x
1856)
(continued)
Appendix
(m)
Appendix
Ypsilothuria talismani 270– mb x x

talismani Perrier, 1886 2684
Order Aspidochirotida
Family Holothuriidae
Actinopyga agassizii (Selenka, 0–54 cr,rb,sg,rub x x x x x x x x x x x
1867)
Holothuria (Cystipus) cubana 0–8 cr,mb,rb,sb x x x x x x x x
Ludwig, 1875
Holothuria (Cystipus) 0–457 mb,sb x x x x x x x
occidentalis Ludwig, 1875
Holothuria (Cystipus) 3–370 cr,mb,sb,rub x x x x x x x
pseudofossor Deichmann, 1930
Holothuria (Halodeima) 0–10 cr,m,rb,sb,sg,rub x x x x x x x x x
floridana Pourtalès, 1851
Holothuria (Halodeima) grisea 0–30 cr,mb,rb,sb,sg x x x x x x x x x x
Selenka, 1867
Holothuria (Halodeima) 0–183 cr,m,mb,rb,sb,sg x x x x x x x x x x x x x
mexicana Ludwig, 1875
Holothuria (Holothuria) 0–15 mb,sb x
dakarensis Panning, 1939
Holothuria (Holothuria) helleri 15 sb,rub x
Marenzeller, 1878
Holothuria (Holothuria) 1–77 rb x
mammata Grube, 1840
Holothuria (Holothuria) 0–30 mb,sb,sg x
tubulosa Gmelin, 1788
Holothuria (Platyperona) 0–8 cr,m,mb,rb,sb,rub x x x x x x
parvula (Selenka, 1867)
Holothuria (Platyperona) rowei x
Pawson & Gust, 1981
Holothuria (Platyperona) 0–30 rb x
sanctori Delle Chiaje, 1823
Holothuria (Panningothuria) 3–193 rb x
forskali Delle Chiaje, 1823
Holothuria (Roweothuria) 0–30 rb x
arguinensis Koehler & Vaney,
1906
Holothuria (Roweothuria) polii 0–250 mb,sb,sg x
649
Delle Chiaje, 1823
(continued)
650
(m)
Holothuria (Semperothuria) 0–42 cr,m,mb,rb,sb,sg,rub x x x x x x x x
surinamensis Ludwig, 1875
Holothuria (Selenkothuria) 0–42 mb,rb,sb x x x x x x x x x
glaberrima Selenka, 1867
Holothuria (Theelothuria) 0–402 cr,mb,rb,sb,sg x x x x x x
princeps Selenka, 1867
Holothuria (Thymiosycia) 0–121 cr,m,mb,sb,sg x x x x x x x x x x x
arenicola Semper, 1868
Holothuria (Thymiosycia) 0–27.4 cr,mb,rb,sb,sg x x x x x x x
impatiens (Forskaal, 1775)
Holothuria (Thymiosycia) x
rathbunii Lampert, 1885
Holothuria (Thymiosycia) 0–30 cr x x x x x x x x x
thomasi Pawson & Caycedo,
1980
Holothuria (Vaneyothuria) 20–466 mb,rb,sb x x x x
lentiginosa von Marenzeller,
1892
Holothuria (Vaneyothuria) 69–466 cr,mb x
lentiginosa enodis Miller &
Pawson, 1979
Family Stichopodidae
Astichopus multifidus (Sluiter, 1–162 cr,rb,sb,sg x x x x x x
1910)
Eostichopus arnesoni Cutress 36 sb x x
& Miller, 1982
Eostichopus regalis (Cuvier, 5–800 mb,sb,sg,rub x x
1817)
Isostichopus badionotus 0–274 cr,mb,rb,sb,sg x x x x x x x x x x x
(Selenka, 1867)
Isostichopus macroparentheses 0–18 rb x x
(H. L. Clark, 1922)
Parastichopus tremulus 20–1960 mb,sb x
(Gunnerus, 1767)
Family Synallactidae
Amphigymnas bahamensis 439–900 mb x x x
Deichman, 1930
Appendix
(continued)
(m)
Appendix
Bathyplotes bigelowi 402–585 mb x

Deichmann, 1940
Bathyplotes natans (M. Sars, 210– cr,mb x x
1868) Östergren, 1896 1600
Bathyplotes pourtalesi (Théel, 134– mb x x x
1886) 2919
Hansenothuria benti Miller & 548–903 cr,sb x
Pawson, 1989
Meseres atlanticus (R. Perrier, x
1902)
Meseres occultatus (von 232– mb x
Marenzeller, 1893) 2180
Mesothuria connectens 1975– x
(Perrier, 1898) 2515
Mesothuria gargantua 180–720 mb x
Deichmann, 1930
Mesothuria intestinalis 18–1445 mb x x
(Ascanius, 1805)
Mesothuria (Zygothuria) lactea 484– mb x x x x
(Théel, 1886) 5100
Mesothuria maroccana Perrier, 914– mb x x
1902 5062
Mesothuria (Mesothuria) 151– mb x
rugosa Herouard, 1912 3890
Mesothuria (Penichrothuria) 699– mb x x x x
verrilli (Théel, 1886) 4060
Molpadiodemas depressus 1353– x
(Herouard, 1902) 5690
Paelopatides gigantea (Verrill, 1100– x
1884) 4060
Pseudostichopus occultatus 232 mb,sb x
Merenzeller, 1893
Pseudostichopus peripatus 3411– x
(Sluiter, 1901) 3459
Scotothuria herringi Hansen, x
1978
Synallactes crucifera Perrier, 1000– mb x x
1898 3476
651
(continued)
652
(m)
Order Elasipodida
Family Deimatidae
Deima validum validum Théel, 914– mb x x x
1879 2780
Orphnurgus asper Théel, 1882 600– mb x
1049
Family Elpidiidae
Peniagone purpurea (Théel, 2800– x
1882) 3000
Family Pelagothuridae
Enypniastes eximia Théel, 1882 461–689 x
Family Psychropotidae
Benthodytes lingua Perrier, 540– mb x x
1896 2200
Benthodytes sanguinolenta 914– mb x x x
Théel, 1882 3100
Benthodytes typica Théel, 1882 315– mb x x x x
5046
Benthodytes valdiviae Hansen, x
1975
Psychropotes depressa (Théel, 955– mb x x
1882) 4060
Psychropotes longicauda 2210– x
Théel, 1882 5000
Psychropotes scotiae (Vaney, x
1908)
Psychropotes semperiana 1143– x x
Théel, 1882 5600
Order Molpadiida
Family Caudinidae
Acaudina molpadioides 3545– mb x
(Semper, 1867) 3635
Paracaudina chilensis chilensis 0–990 mb,sb,rub x x x
(Müller, 1850)
Family Molpadiidae
(continued)
Appendix
(m)
Appendix
Molpadia barbouri Deichman, 440– mb x x

1940 1529
Molpadia blakei (Théel, 1886) 3482– mb x x
3518
Molpadia cubana Deichmann, 24–1464 mb x x x
1940
Molpadia eltaninae Pawson, 20–855 x x
1977
Molpadia musculus Risso, 1826 35–5205 mb x x x x x x x
Molpadia oolitica (Pourtalès, 42–1440 mb x
1851)
Molpadia parva (Théel, 1886) 125– mb x x x x
2844
Order Apodida
Family Chiridotidae
Chiridota marenzelleri R. 40–159 mb,rub x x
Perrier, 1904
Chiridota rotifera (Pourtalès, 0–360 cr,rb,sb,sg x x x x x x x
1851)
Taeniogyrus contortus 82–200 mb,sb x x
(Ludwig, 1875)
Trochodota purpurea (Lesson, 0–100 sb,rub x x
1830)
Family Synaptidae
Anapta fallax Lampert, 1889 0–350 sb,rub x x
Epitomapta roseola (Verrill, 1.5–40 rb,sb x x x
1874)
Eupatinapta acanthia (A. 10.7 x
H. Clark, 1899)
Euapta lappa (Müller, 1850) 0–256 cr,rb,sb,sg x x x x x x x x x x x
Leptosynapta brasiliensis 1–4 x
Freire & Grohmann, 1989
Leptosynapta imswe Pawson, 0–2 x
1976
Leptosynapta inhaerens 0–173 mb x
(Müller, 1776)
653
(continued)
654
(m)
Leptosynapta multigranula H. 0.5–2.9 sb x x
L. Clark, 1924
Leptosynapta nannoplax x
Pawson, 1976
Leptosynapta parvipatina H. 9–12 x
L. Clark, 1924
Leptosynapta roseogradia 0.5 x
Pawson, 1976
Leptosynapta tenuis (Ayres, 0.5–167 x x
1851)
Protankyra benedeni (Ludwig, 5–100 x x
1881)
Protankyra brychia (Verrill, x
1885)
Protankyra ramiurna Heding, 0–18 mb,rb,sb,sg x x
1928
Synaptula hydriformis 0–10 cr,m,sb,sg x x x x x x x x x x x
(Lesueur, 1824)
Synaptula secreta Ancona- 1–10 x
Lopez, 1957
Habitat classification: cr Coral Reefs, m mangroves, mb muddy bottom, rb rocky bottom, sb sandy bottom, sg seagrass, rub rubble bottom. Countries abbreviation: MEX Mexico, BEL Belize, GUA
Guatemala, HON Honduras, NIC Nicaragua, CRC Costa Rica, PAN Panama, COL Colombia, VENc Venezuela Caribbean, VENa Venezuela Atlantic, BRA Brazil, URG Uruguay, ARG Argentina, MAV
Malvinas Islands, CUB Cuba, HAI Haiti, RDO Dominican Republic, PRI Puerto Rico, CAN Canary Islands
Appendix
Index
A B
Abatus, 375, 379 Barren, 483–485, 504
Abundance, 6, 18, 28, 30, 372 Belize, 87, 88
Acanthasther planci, 28 Bernasconi, Irene, 5, 360
Albatross, 15, 70, 110, 132, 136, 157, 189, Biodiversity, 12, 24, 45, 145, 191
365, 406, 442 Bioerosion, 30, 88
Allan Hancock, 15, 71, 111, 190 Biomass, 33, 43, 172, 208
Allometric scaling, 202, 205 Bocas del Toro, 18, 70, 73, 108
Allostichaster, 367, 369, 382 Boulder, 84, 169, 199
Amphiura, 26, 28, 167, 235, 247, 319–321, Brazil, 2, 6, 126, 305
368, 374 Brazilian Province, 351, 496
Amplaster, 353 Brazilian Current, 345, 351
Anasterias, 367, 369, 371, 382, 384, 385
Anchialine, 27
Ancon, 278 C
Aquaculture, 3, 6, 48, 99, 132, 173 Cabo Verde, 481, 482, 495
Arbacia, 19, 25, 71, 82, 131, 240, 243, 256, Caenocentrotus gibbosus, 195, 206, 289
288, 289, 309, 326, 368, 375, 388, 451, Cahuita, 81
494, 497, 501Astropecten articulatus Canary Islands, 6
Archipelago, 92, 108, 149 Caño Island, 88
Argentina, 2, 4, 346, 365 CARICOMP, 81, 82
Argentine Continental Shelf, 360 Caso, Ma. Elena, 5, 6, 16
Argentine Province, 367 Central America, 80
Argentine Sea, 360, 364, 366 Centrostephanus coronatus, 19, 29, 170, 195,
Astropecten, 18, 23, 25, 26, 36, 39, 71, 93, 98, 206, 259, 285
167, 172, 173, 192, 241, 242, 263, 265, Cherbonier, 4, 5, 442
291, 315–317, 324, 369, 373, 381, 458, Chile, 4, 187, 281, 387
492, 495 Clypeaster rosaceus, 82, 122, 255, 256,
Astropecten articulatus, 93, 265, 413, 458 413, 416
Astropecten duplicates, 265, 413, 458 Cóbanos, Los, 84
Athyonidium chilensis, 285, 294, 297 Cocos Island, 88
Atlantis, 408, 414 Coelomocyte types, 208
Azores, 479, 481 Colombian Museum of Marine Natural History
Aztecs, 4, 39, 40 (MHNMC), 156

in Latin America, DOI: 10.1007/978-3-642-20051-9,
656 Index
Cosmasterias, 369, 373, 381, 382, 388 F

Conservation, 45, 46, 75, 221 Ferrol, 278
Coral Reefs, 6, 25, 77, 110 Fisheries, 6
Costa Rica, 68, 71, 79, 93, 515 Fission, 197–199, 369, 371
Culebra Island, 120, 439, 458 Florida Museum of Natural
Ctenantedon kinziei, 75, 154, 161 History (FLMNH)
Fossil, 80
French Guiana, 4, 156, 516
D
Deep azooxanthellate coral
communities, 164, 167 G
Diadema antillarum, 26, 32, 80, 93, 113, 126, Galapagos, 18
136, 145, 154, 162, 173, 240, 262, 409, Glycosides, 382
413, 414, 442, 446, 448, 460, 464, Gorgona, 131, 49, 153, 157
483, 530 Gran Canaria, 475, 482
Diadema aff. antillarum, 82 Gulf of California, 13, 15, 18
Diadema mexicanum, 29, 31, 33, 83, 159, 192, Gulf of Chiriqui, 109, 110, 117, 130
195, 215, 528 Gulf of Fonseca, 68, 71, 84
Disease, 34, 134, 210, 262 Gulf of Mexico, 32
Distribution, 6, 18, 32 Gulf of Nicoya, 68, 71, 80
Diversity, 18 Gulf of Panama, 69, 109, 112, 120, 131
Dominican Republic, 6 Guyana, 4, 156, 442
E H
Earthquake, 262 Haiti, 6, 87
Echinometra lucunter, 26, 32, 79, 83, 90, 93, Heavy metals, 96, 263, 413
113, 122, 162, 240, 256, 308, 413, 444, Heliaster helianthus, 285, 288, 384
459, 464, 465 Hispabiota Marina Project, 431
Echinometra vanbrunti, 19, 24, 29, 31, 36, 39, Hispaniola, 2, 427, 440
83, 84, 93, 159, 170, 172, 183, 191, Holothuria, 15, 19, 24–26, 31, 35, 36, 44, 82,
195, 206, 208, 223, 289, 528 84, 86, 90, 93, 97, 119, 131, 132, 154,
Echinometra viridis, 80, 87, 88, 113, 154, 240, 155, 168, 170–172, 175, 191, 192, 198,
444, 528 199, 210, 217, 218, 242, 289, 291, 294,
Ecoregion, 37, 38 309, 322, 323
Eco-regionalization, 416 Holothuria atra, 154, 195, 199
Ecozone, 13, 416–418, 421 Holothuria glaberrima, 173, 245, 450, 451
Ecuador, 132, 184, 186, 203 Holothuria theeli, 86, 191, 198, 199,
El Hierro, 474, 475, 501 217, 294
El Niño, 88 Holothuria (Thymiosycia) thomasi
El Salvador, 2, 67, 71, 84, 93 Honduras, 80, 82, 87
Encope grandis, 26, 128 Huatulco, 154, 462
Encope micropora, 24, 183, 205, 206, 212, 291 Hydro-physical stress, 413
Endemism, 26, 38, 92, 284, 515
Equatorial front, 185
Eucidaris thouarsii, 29, 31, 84, 170, 191, I
289, 528 Immune response, 208, 223, 310, 455
Eucidaris galapagensis, 183, 192, 195, 200, INVEMAR, 148, 153, 164
201, 213–215, 221, 223 Isostichopus badionotus, 44, 86, 119, 132, 172,
Eucidaris tribuloides, 25, 26, 80, 93, 154, 254, 242–246, 324
324, 411, 461, 465 Isostichopus fuscus, 19, 24, 31, 33, 35, 36, 42,
Eulimids, 214 93, 97, 172, 192, 196–199, 210, 217,
Euapta lappa, 26, 413, 416, 462, 530 219, 223, 292, 297
Extinction, 44, 125, 221 Isthmus, 68, 123, 129, 533
Index 657
J O
Johnson-Smithsonian Deep-Sea Ophiocoma, 24, 26, 79, 90
Expedition, 435 Ophioplocus, 320, 374
Ophiothrix, 348, 492, 494
Oreaster reticulatus, 4, 433, 434, 494
L
La Gomera, 474, 475
La Niña, 208 P
La Parguera, 440, 447, 449 Paita, 278, 281, 284, 288
Larval abundance, 444, 447, 463 Paleontology, 252, 95
Larval biology, 198 Panama, 2, 5, 6, 38, 67, 68, 70, 73, 79, 81, 82,
Leptopentacta deichmannae, 235, 247 87, 92, 93, 98, 107–110, 112–114,
Loxechinus albus, 288, 294, 296, 2978, 375, 117–120, 122–133, 135–137, 147, 149,
377, 378 156, 157, 257, 465, 515, 516, 533, 536
Lytechinus variegatus, 25, 32, 80, 86, 113, Paracas, 278, 280, 288, 289, 294
154, 164, 234, 235, 240, 244, 258, 309, Parasitism, 210, 214
411, 414, 444, 445, 461, 464 Paraster doederleini, 82
Lytechinus semituberculatus, 192, 195, 201, Parastichopus parvimensis, 33, 35, 46
202, 216, 288 Parish-Smithsonian, 429
Luidia, 18, 23, 25, 26, 32, 71, 98, 172, 173, Patagonian, 351, 364, 371, 374, 380–382, 387,
194, 207, 243, 263, 288, 291, 305, 316, 388, 516
324, 360, 369, 373, 382, 384, 492, Pattalus mollis, 285, 297
493, 528 Pearl Islands, 109, 112, 128
Luidia senegalensis, 413, 415, 305 Peru, 2, 4, 6, 92, 132, 157, 277, 278, 280-284,
288, 289, 291, 292, 294, 296–298,
515–517, 524, 92
M Pharia pyramidata, 170, 191
Macaronesian, 481, 483 Phataria unifascialis, 28, 29
Madeira, 125, 481, 483 Phylogenetic, 89, 120, 202
Magellanic Province, 364 Phylogeography, 126
Magueyes, 443, 448 Physiology, 90, 120, 244, 450
Malpelo, 149, 153, 157, 160 Phytoplankton, 16, 406
Malvinas (Falkland) Current, 303 Platasterias, 26, 207, 544
Mangrove, 12, 77, 119 Predation, 183, 210, 213
Marine Biodiversity Information System Programa EcoMar, 431
(SIBM), 147, 156 Pseudechinus, 308, 375, 381
Marine Science Collection of Department of Pseudocnus, 288
Marine Sciences (MSC), 443 Psolidium, 292
Mexico, 6, 8 Puerto Rico, 2, 6, 427, 431, 439, 441–444,
Molecular evolution, 455 447, 449, 450, 456–462, 464–466
Monophoraster, 352, 353
Mortality, 87
Museum of Comparative Zoology (MCZ), 430 R
Museum of Natural Sciences Recruitment, 115, 444
of Berlin (ZMB), 430 Regeneration, 452, 453
Reproduction, 35, 89, 120, 243, 373, 375,
443, 446
N Reproductive cycles, 120
National Museum of Natural History of Santo Río de la Plata, 345, 346, 350, 351, 360, 362,
Domingo (MNHNSD), 430 364, 515
Nicaragua, 2, 67, 68, 70, 71, 73, 81–83, 86, 93, Rosario Corals, and San Bernardo Islands
97, 98 National Natural Park (PNNCRSB),
Nidorellia armata, 172, 192, 291 154, 161, 167
Norcross-Bartlett, 429 RVBlake, 442
658 Index
RV John Elliot Pillsbury, 429, 430 Tethyaster vestitus vestitus, 32, 80, 82
RV Oregon, 73, 112 Tetrapygus niger, 4
RV Stranger, 429 Thalassia testudinum, 433
Toxopneustes roseus, 114, 129
Trench, 68, 238, 248, 251, 441
S Tripneustes depressus, 130, 414, 433
Salvajes, 481 Tripneustes ventricosus, 26, 82, 240, 444
Samanco, 278, 288 Trophic cascade, 213, 500
San Andrés and Providencia, 147–149, 164
San Blas, 70, 108, 113, 114, 119, 133, 134, 202
Seagrass, 163 U
Sechura, 278, 288 UNAM, 15, 16, 47, 48, 410
Settlement, 444, 446, 448 Upwelling, 148, 187, 189, 235, 236, 238, 248,
Speciation, 127 251, 281, 304
Spawning, 41, 369, 372, 378, 379, 444 Uruguay, 2, 6, 317, 345–351, 353, 354, 362,
Spicule, 86, 219 365, 516
Stichaster striatus, 297 Urracá, 71, 112
Stichopus horrens, 198
Surinam, 4
Sustainable production, 4, 156 V
Vieques island, 439, 458
T
Tayrona National Natural Park, 154, 161 Z
Tenerife, 474, 475, 481, 483, 492, 501, 502 Zaca, 111
Te Vega, 72

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Echinoderm Research and Diversity

Echinoderm Research and

ISBN 978-3-642-20050-2 ISBN 978-3-642-20051-9 (eBook)

Library of Congress Control Number: 2012941234

Ó Springer-Verlag Berlin Heidelberg 2013

Printed on acid-free paper

Springer is part of Springer Science+Business Media (www.springer.com)

This is a remarkable book. First and foremost, it compiles an immense amount of

Florida, USA, March 2012 John M. Lawrence

San José, Costa Rica, March 2012 Jorge Cortés

The Iberoamerican Echinoderms Network (Red Iberoamericana de Equinodermos)

Juan José Alvarado

1 Echinoderm Research and Diversity in Latin America. . . . . . . . . 1

2 The Echinoderms of Mexico: Biodiversity, Distribution

3 Central America Echinoderms: Diversity, Ecology

4 Echinoderm Diversity in Panama: 144 Years of Research

5 Echinoderms of Colombia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

6 Echinoderms of Ecuador . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

7 Echinoderms from Venezuela: Scientific Recount,

8 Echinoderms of Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277

9 Echinoderm from Brazil: Historical Research and the Current

10 Echinoderm Research in Uruguay . . . . . . . . . . . . . . . . . . . . . . . 345

11 Echinoderms from Argentina: Biodiversity, Distribution

12 Echinoderms of the Cuban Archipelago. . . . . . . . . . . . . . . . . . . 403

13 Recent Echinoderms from Hispaniola . . . . . . . . . . . . . . . . . . . . 425

14 Review of Echinoderm Research in Puerto Rico,

15 Echinoderms of the Canary Islands, Spain. . . . . . . . . . . . . . . . . 471

16 Latin America Echinoderm Biodiversity and Biogeography:

Juan José Alvarado and Francisco Alonso Solís-Marín

The phylum Echinodermata consists of approximately 7,000 living species and

J. J. Alvarado and F. A. Solís-Marín (eds.), Echinoderm Research and Diversity 1

21,069,501 km2, and according to the Sea around Us project. It possesses

1.2 Echinoderm Research in LA: An Overview

Latin American echinoderm studies started in the 13th century. Bernardino de

Lyman (1833–1897), an U.S. zoologist who studied under the supervision of

1.3 General Results

Most Latin American countries have a common beginning in echinoderm research,

Museum of Comparative Zoology at Harvard University, California Academy of

Insituto de Ciencias del Mar y Limnología de la Universidad Nacional Autónoma

Cherbonier G (1959a) Echinodermes de la Guyane Francaise (Crinoides, Astérides, Ophiurides,

Francisco A. Solís-Marín, Magali B. I. Honey-Escandón,

F. A. Solís-Marín (&) M. B. I. Honey-Escandón A. Durán-González

J. J. Alvarado and F. A. Solís-Marín (eds.), Echinoderm Research and Diversity 11

2.1 Oceanography and Marine Environments in Mexico

2.2 Echinoderm Fauna of Mexico: An Inventory and Analysis

2.2.1 Historical Review

Studies on Mexican echinoderms started in the nineteenth century, when some

Fig. 2.2 Maria Elena Caso

Fig. 2.3 Echinoderm

Gonzalez-Azcarraga (2009), Bribiesca-Contreras (2011), Estrada-Rodriguez

2.2.2 Biodiversity and Distribution of the Mexican Echinoderms

2.2.3 Gulf of California

Fig. 2.4 Number of

2.2.4 Mexican Pacific Ocean

genus is Amphiodia, with seven species. Ophiactis savignyi is the ophiuroid

2.2.5 Gulf of Mexico

The delimitation of endemism in marine species is difficult due to dispersion of

guillermosoberoni is the only Mexican ophiuroid species known to inhabit low

2.2.6 Mexican Fossil Echinoderms

2.3.1 Ecology and Reproduction Studies on Mexican Echinoderms

Ecological studies of the Echinodermata in Mexico have contributed to the

Salcedo-Martinez et al. (1988) generated a benthic inventory of echinoderms in

Caso et al. (1996) performed qualitative and quantitative analyses of the