(Biology and Ecology of Marine Life) David, Sheba Rani Nakka - Ramesh, Santhanam - Santhanam, Ramasamy - Biology and Ecology of Pharmaceutical Marine Life - echinoderms-CRC Press (20 PDF

Biology and Ecology of
Pharmaceutical Marine Life:

Echinoderms
Biology and Ecology of Marine Life
Series Editor
Ramasamy Santhanam
Former Dean, Tamil Nadu Veterinary and Animal Sciences University
Thoothukudi, India
Biology and Ecology of Edible Marine Bivalve Molluscs

Ramasamy Santhanam
Biology and Ecology of Edible Marine Gastropod Molluscs

Ramasamy Santhanam
Biology and Ecology of Pharmaceutical Marine Cnidarians

Ramasamy Santhanam, Santhanam Ramesh, Gubbiyappa Shivakumar
Biology and Ecology of Pharmaceutical Marine Life: Echinoderms

Ramasamy Santhanam, Santhanam Ramesh, Sheba Rani Nakka David
Biology and Ecology of Pharmaceutical Marine Mollusks

Ramasamy Santhanam, Manavalan Gobinath, Santhanam Ramesh
Biology and Ecology of Pharmaceutical Marine Plants

Ramasamy Santhanam, Santhanam Ramesh, Hafiz Ansar Rasul Suleria
Biology and Ecology of Pharmaceutical Marine Sponges

Ramasamy Santhanam, Santhanam Ramesh, Anbu Jeba Sunilson
Biology and Culture of Portunid Crabs of World Seas

Ramasamy Santhanam
Biology and Ecology of Toxic Pufferfish

Ramasamy Santhanam
Biology and Ecology of Venomous Marine Snails

Ramasamy Santhanam
Biology and Ecology of Venomous Stingrays

Ramasamy Santhanam
For more information about this series, please visit:

https://www.crcpress.com/Biology-and-Ecology-of-Marine-Life/book-series/AAPBIOECOMAR
Biology and Ecology of
Pharmaceutical Marine Life:
Echinoderms
Ramasamy Santhanam
Santhanam Ramesh
Sheba Rani Nakka David
CRC Press
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Contents
Preface...............................................................................................................................................ix
Authors...............................................................................................................................................xi
Chapter 1 Introduction...................................................................................................................1
1.1 Echinoderms—Overview...................................................................................1
1.2 Ecology of Echinoderms....................................................................................2
1.3 Biology of Echinoderms.....................................................................................2
1.3.1 Anatomy.............................................................................................. 2
1.3.2 Food and Feeding Habits....................................................................2
1.3.3 Digestive System................................................................................. 3
1.3.4 Nervous System and Senses................................................................ 3
1.3.5 Circulatory System..............................................................................3
1.3.6 Respiratory System.............................................................................3
1.3.7 Excretory System................................................................................3
1.3.8 Reproductive System...........................................................................3
1.3.9 Regeneration Ability........................................................................... 4
1.3.10 Predators..............................................................................................4
1.3.11 Venomous/Poisonous Echinoderms....................................................4
1.4 Characteristics of Echinoderms.........................................................................5
1.4.1 Basic Features of Echinoderms...........................................................5
1.4.2 Characteristics of Different Classes of Echinoderms......................... 5
1.4.2.1 Class Asteroidea...................................................................5
1.4.2.2 Class Ophiuroidea................................................................ 6
1.4.2.3 Class Echinoidea.................................................................. 7
1.4.2.4 Class Holothuroidea.............................................................9
1.4.2.5 Class Crinoidea.................................................................... 9
1.5 Conservation Measures of Echinoderms.......................................................... 11
1.6 Importance of Echinoderms............................................................................. 11
1.6.1 Echinoderms in Commercial Fisheries............................................. 12
1.6.1.1 Sea Urchin Species with a Major Contribution to the
Modern Fishery.................................................................. 13
1.6.1.2 Sea Cucumber Species that Contribute to the Modern
Fishery................................................................................ 14
1.6.2 Echinoderms in Aquaculture (Farming)........................................... 15
1.6.3 Echinoderms as Food........................................................................ 15
1.6.4 Echinoderms in the Aquarium and Souvenir Trades........................ 16
1.6.4.1 Potential Echinoderms in the Ornament Industry............. 16
1.6.5 Echinoderms in Scientific Research (Experimental Model
Systems)............................................................................................. 16
1.6.6 Echinoderms in Bionics.................................................................... 17
1.6.7 Echinoderms in Bioindicators of Environmental Quality................ 17
1.6.8 Ecological Role of Echinoderms....................................................... 17
1.6.9 Echinoderms as Medicine................................................................. 18
v
vi Contents
1.6.10 Echinoderms as Sources of Pharmaceutical (Bioactive)

Compounds and in the Development of New Drugs....................... 18
1.6.10.1 Present Status of New Natural Products (NP) as
Pharmaceutical Compounds from Echinodermata........ 19
1.6.10.2 Pharmaceutical (Bioactive) Compounds from Sea
Stars................................................................................ 19
1.6.10.3 Pharmaceutical (Bioactive) Compounds from
Brittle Stars.....................................................................20
Urchins...........................................................................20
Cucumbers......................................................................20
Lilies............................................................................... 21
Chapter 2 Biology and Ecology of Pharmaceutical Sea Stars (Class: Asteroidea)...................... 23

2.1 Family: Asteriidae (Class: Asteroidea; Order: Forcipulatida)........................ 23
2.2 Family: Heliasteridae (Order: Forcipulatida).................................................66
2.3 Family: Stichasteridae (Order: Forcipulatida)................................................ 67
2.4 Family: Zoroasteridae (Order: Forcipulatida)................................................ 70
2.5 Family: Astropectinidae (Order: Paxillosida)................................................ 72
2.6 Family: Ctenodiscidae (Order: Paxillosida)...................................................84
2.7 Family: Luidiidae (Order: Paxillosida).......................................................... 85
2.8 Family: Porcellanasteridae (Order: Paxillosida)............................................92
2.9 Family: Echinasteridae (Order: Spinulosida)................................................. 93
2.10 Family: Pterasteridae (Order: Spinulosida).................................................. 103
2.11 Family: Acanthasteridae (Order: Valvatida)................................................ 106
2.12 Family: Archasteridae (Order: Valvatida).................................................... 111
2.13 Family: Asterinidae (Order: Valvatida)........................................................ 113
2.14 Family: Asteropseidae (Order: Valvatida).................................................... 116
2.15 Family: Goniasteridae (Order: Valvatida).................................................... 119
2.16 Family: Mithrodiidae (Order: Valvatida)..................................................... 127
2.17 Family: Ophidiasteridae (Order: Valvatida)................................................. 132
2.18 Family: Oreasteridae (Order: Valvatida)...................................................... 150
2.19 Family: Asterinidae (Order: Valvatida)........................................................ 168
2.20 Family: Solasteridae (Order: Valvatida)....................................................... 169
Chapter 3 Biology and Ecology of Pharmaceutical Brittle Stars and Basket Stars
(Class: Ophiuroidea).................................................................................................. 171
3.1 Family: Ophiolepididae (Class: Ophiuroidea; Order: Amphilepidida)........ 171
3.2 Family: Ophiopholidae (Order: Amphilepidida).......................................... 172
3.3 Family: Ophiotrichidae (Order: Amphilepidida)......................................... 175
3.4 Family: Gorgonocephalidae (Order: Euryalida).......................................... 177
3.5 Family: Ophiocomidae (Order: Ophiacanthida).......................................... 179
3.6 Family: Ophiomyxidae (Order: Ophiacanthida).......................................... 186
3.7 Family: Ophiodermatidae (Order: Ophiacanthida)...................................... 186
3.8 Family: Hemieuryalidae (Order: Ophiurida)............................................... 188
3.9 Family: Ophiuridae (Order: Ophiurida)....................................................... 190
3.10 Family: Ophiopyrgidae (Order: Ophiurida)................................................. 195
Contents vii
Chapter 4 Biology and Ecology of Pharmaceutical Sea Urchins and Sand Dollars
(Class: Echinoidea).................................................................................................... 197
4.1 Family: Arbaciidae (Order: Arbacioida)...................................................... 197
4.2 Family: Echinidae (Order: Camarodonta)................................................... 198
4.3 Family: Echinometridae (Order: Camarodonta).......................................... 199
4.4 Family: Parechinidae (Order: Camarodonta)...............................................208
4.5 Family: Strongylocentrotidae (Order: Camarodonta).................................. 210
4.6 Family: Temnopleuridae (Order: Camarodonta).......................................... 216
4.7 Family: Toxopneustidae (Order: Camarodonta)........................................... 218
4.8 Family: Scutellidae (Order: Clypeasteroida)................................................226
4.9 Family: Diadematidae (Order: Diadematoida).............................................229
4.10 Family: Loveniidae (Order: Spatangoida)....................................................234
4.11 Family: Schizasteridae (Order: Spatangoida)...............................................236
4.12 Family: Glyptocidaridae (Order: Stomopneustoida)....................................237
4.13 Family: Stomopneustidae (Order: Stomopneustoida)..................................239
Chapter 5 Biology and Ecology of Pharmaceutical Sea Cucumbers (Class: Holothuroidea)...... 243
5.1 Family: Synaptidae (Class: Holothuroidea; Order: Apodida)......................243
5.2 Family: Cucumariidae (Order: Dendrochirotida).........................................244
5.3 Family: Phyllophoridae (Order: Dendrochirotida).......................................276
5.4 Family: Psolidae (Order: Dendrochirotida)..................................................281
5.5 Family: Sclerodactylidae (Order: Dendrochirotida).....................................285
5.6 Family: Elpidiidae (Order: Elasipodida)......................................................292
5.7 Family: Holothuriidae (Order: Holothuriida)...............................................294
5.8 Family: Caudinidae (Order: Molpadida)......................................................349
5.9 Family: Stichopodidae (Order: Synallictida)...............................................352
5.10 Family: Synallactidae (Order: Synallictida).................................................370
5.11 Miscellaneous...............................................................................................372
Chapter 6 Biology and Ecology of Pharmaceutical Sea Lilies or Feather Stars

(Class: Crinoidea)......................................................................................................373
6.1 Family: Antedonidae (Class Crinoidea; Order: Comatulida).......................373
6.2 Family: Comatulidae (Order: Comatulida)..................................................375
6.3 Family: Himerometridae (Order: Comatulida).............................................377
6.4 Family: Mariametridae (Order: Comatulida)...............................................379
6.5 Family: Hemicrinidae (Order: Cyrtocnida)..................................................380
6.6 Family: Holopodidae (Order: Cyrtocnida)...................................................382
6.7 Family: Proisocrinidae (Order: Isocrinida)..................................................383
References...................................................................................................................................... 385
Compounds....................................................................................................................................405
Index............................................................................................................................................... 413
Preface
Nature is an ancient pharmacy, rich in bioactive compounds with several biological properties.
Throughout the ages, natural products have always been the mainstay of disease therapy and are
still considered to play an important role in modern medicine. In this regard, marine organisms are
considered as treasures that remain a relatively unexplored source for novel bioactive compounds.
The marine environment, which is extremely complex with immense biodiversity, is an enormous
source of marine natural products (bioactive compounds or secondary metabolites) with potential
therapeutic properties. The upsurge of interest during the last decade in the ocean as a potential
source of bioactive compounds and new drugs has stimulated a flurry of activity in the research lab-
oratories and clinics throughout the world. Approximately 16,000 marine natural products have so
far been isolated from marine organisms (25% from algae, 33% from sponges, 18% from cnidarians
[coelenterates], and 24% from other invertebrate phyla such as ascidians, molluscs, echinoderms,
and bryozoans).
Among the marine invertebrates, the importance of echinoderms as a promising source of bioac-
tive compounds for development of pharmaceuticals and potential therapeutic applications has been
growing rapidly. Numerous new natural compounds have now been isolated from these echinoderms
with interesting pharmaceutical activities. The bioactive compounds derived from echinoderms are
compounds of interest showing an extensive application in the treatment of many diseases. These
compounds, which can be subdivided into six major chemical classes, namely, polyketides, terpenes,
peptides, alkaloids, shikimates, and sugars have shown several biological properties, such as anti-
bacterial, anticoagulant, antifungal, antimalarial, antiprotozoal, antituberculosis, anti-inflammatory,
antitumour, anti-HIV, and antiviral activities. A new drug, histochrome, isolated from the pigment
cells of sea urchins, presents cardiological and ophthalmological activity. Further, the sea urchin
Arbacia lixula is a new source of the active stereoisomer of astaxanthin, which is known to have
particular bioactivity for the prevention of neurodegenerative diseases. The sea cucumber is also val-
ued as a disease preventive and longevity tonic. It has been listed as a medicinal agent in the Bencao
Congxin (New Compilation of Materia Medica).
Though a few books are presently available on marine natural products, a comprehensive book
on the biology and ecology of pharmaceutical echinoderms has not so far been published. This
publication, the first of its kind, with contributions from scientists in both the marine biology and
pharmacy disciplines, fills this long-felt need. The taxonomy, common name, global distribution,
habitat, diagnostic features and pharmaceutical compounds (along with their activities) of 215 spe-
cies of echinoderms are given with suitable illustrations. It is hoped that the present publication,
when brought out, will be of great use as a standard reference for researchers, teachers and students
of various disciplines such as marine biology, pharmacy, fisheries science, biochemistry and bio-
technology; libraries of colleges, universities and institutions; and as a valuable guide for pharma-
ceutical companies involved in the development of new drugs from echinoderms.
We are highly indebted to Dr. K. Venkataramanujam, former Dean, Fisheries College and
Research Institute, Tamil Nadu Veterinary and Animal Sciences University, Thoothukudi, India,
for his valued comments and suggestions on the manuscript. We sincerely thank all our interna-
tional friends who were kind enough to collect and send certain species of echinoderms for the
present purpose. The services, such as photography and secretarial assistance, rendered by Mrs.
Albin Panimalar Ramesh, are also gratefully acknowledged.
ix
Authors
Ramasamy Santhanam is the former dean of Fisheries College and Research Institute, Tamil
Nadu Veterinary and Animal Sciences University, India. His fields of specialization are fisher-
ies environment and marine biology. He presently serves as a fisheries expert for various gov-
ernment and nongovernment organizations in India. Dr. Santhanam has published twenty-five
books on various aspects of fisheries science and seventy research papers. He was a member of
the American Fisheries Society, United States; World Aquaculture Society, United States; Global
Fisheries Ecosystem Management Network (GFEMN), United States; and IUCN’s Commission on
Ecosystem Management, Switzerland.
Santhanam Ramesh, Senior Lecturer, Faculty of Pharmacy, SEGi University, Selangor, Malaysia,
obtained a PhD from Jawaharlal Nehru Technological University, Hyderabad, India, in 2011. His
fields of specialization are biopharmaceutical products, pharmaceutical nanotechnology and novel
drug delivery systems. Dr. Ramesh has ten years of teaching experience and three years of over-
seas experience. He has authored six books, Marine Pharmaceutical Compounds; Freshwater
Phytopharmaceutical Compounds; Biology and Ecology of Pharmaceutical Marine Plants;
Biology and Ecology of Pharmaceutical Marine Sponges; Biology and Ecology of Pharmaceutical
Marine Molluscs and A Text Book of Novel Drug Delivery Systems. Ten of his research papers have
been published in nationally and internationally reputed journals. He presently serves as an associ-
ate member of the Academy Pharmacy Group, Royal Pharmaceutical Society, Great Britain, and as
a fellow of the Institution of Chemists, India.
Sheba Rani Nakka David, Senior Assistant Professor, PAPRSB Institute of Health Sciences (IHS),
Universiti Brunei Darussalam (UBD), Brunei Darussalam, earned a PhD in biomedical engineer-
ing from the School of Bioscience and Engineering, Jadavpur University, Kolkata, India. She has
thirteen years of experience in academic research and four years in industry. Her research inter-
ests include marine natural products and their screening, biomaterials and transdermal formula-
tions. She has thirty-six research publications in peer-reviewed journals to her credit, and she is
an editorial board member of internationally reputed journals, including Pharmacognosy and the
International Journal of Pharmacy and Natural Medicines. She is also a member of professional
organizations such as the Biomedical Engineering Society of India, Pharmacy Council of India and
Society for Biomaterials & Artificial Organs of India.
xi
1 Introduction
1.1 ECHINODERMS—OVERVIEW
The echinoderms (phylum Echinodermata)—the name comes from the Greek word for “spiny
skin”—are a globally distributed and morphologically diverse group of invertebrates whose his-
tory dates back 500 million years. Approximately 13,000 echinoderms species are known of which
7,000 living species fall into five well-defined classes of this phylum, including sea stars (starfish)
(Asteroidea, 1,500 species); brittle stars (Ophiuroidea, 2,000 species); sea urchins, heart urchins,
and sand dollars (Echinoidea, 1,700 species); sea cucumbers (Holothuroidea, 110 species); and sea
lilies and feather stars (Crinoidea, 600 species). Interestingly, the echinoderms are a group of almost
exclusively marine organisms with the few exceptions living in brackish water. Almost all of the
echinoderms are benthic, i.e., they live on or in the substrate. While the echinoderms are a diverse
group, they share certain unique features—viz. pentaradial symmetry, an endoskeleton made up
of ossicles, a water vascular system, and mutable collagenous tissue. While larval echinoderms
are bilaterally symmetrical, the adults are pentaradially symmetrical. The water vascular system
is a complex network of canals and reservoirs. It uses hydraulic pressure and the action of muscles
to operate the various podia (tube feet and tentacles). While tube feet are used in locomotion and
attachment, tentacles are used in feeding. The podia also function in gas exchange and sensory
reception. The water vascular system is usually open at one end of the animal. This opening is
called the madreporite, where seawater enters the system. While most asteroids and echinoids have
madreporites, the holothuroids have an internal madreporite. The madreporite is absent in the cri-
noids. Sea urchins and some sea stars have pedicellariae (jaw-like structures) on the aboral side.
These pedicellariae are used to keep debris off the organism, to hold and capture small prey and
in defence. The echinoderms, like some other groups of animals, have the ability to regenerate lost
body parts. The crinoids are particularly adept at regenerating body parts and can regenerate both
their arms and visceral mass. The crinoids also use regeneration to grow additional arms (i.e., they
autotomize one arm and re-grow two arms). The brittle star (ohiuroid) splits across its central disc
and each half regenerates into a new individual.
1
2 Biology and Ecology of Pharmaceutical Marine Life: Echinoderms
1.2 ECOLOGY OF ECHINODERMS

The echinoderms, which live exclusively in the marine habitat, are found distributed in almost all
depths (from shallow, intertidal coastal waters to over 3,000 m deep in ocean trenches) and lati-
tudes, as well as reef environments or shores. They are benthic, stenohaline marine invertebrates
and their limited osmoregulatory ability restricts the majority of species to the marine environment.
As such, this phylum has become a symbol of marine life (Mondol et al., 2017).
1.3 BIOLOGY OF ECHINODERMS

1.3.1 Anatomy
Echinoderms are characterized by their radial symmetry, many having five or multiples of
five arms. They have a shell, made mainly of calcium carbonate, which is covered by skin.
During larval development, an echinoderm has a distinct bilateral symmetry that is lost during
metamorphosis and the radial symmetry appears only after the formation of the mesoderm.
The adult echinoderms have a water vascular system with external tube feet, used mainly in
locomotion, and a calcareous endoskeleton consisting of ossicles connected by a mesh of collagen
fibres. The skeletal system is one of the most characteristic features of the echinoderms, which
varies both in the arrangement details, as well as in their development extent, between the five
different classes. Spines are frequently associated with these plates.
1.3.2 Food and Feeding Habits

Sea stars are carnivores and scavengers. They feed on all sorts of other invertebrates, particularly
bivalves, snails, crustaceans, marine worms, other echinoderms and even fish. Sea stars use suc-
tion in the tube feet for movement and feeding. They wrap their bodies around quahogs and other
bivalves, using the suction from their tube feet to pull shells apart. When the prey is opened, the sea
star pushes its stomach out of its body and into the bivalve, secreting enzymes that digest the prey’s
soft body tissues. The liquefied bivalve is then absorbed into the stomach. Sea stars are attracted
to the bodies of dead animals on the seafloor. Some are very specific in their eating habits and will
only eat sea cucumbers for instance, while others are more adaptable and will eat a wide variety of
prey. Crinoids and some brittle stars are passive filter feeders, absorbing suspended particles from
passing water. Sea urchins, which are grazing herbivores, have a highly developed feeding structure
called Aristotle’s lantern. This is a five-sided pyramidal shaped structure with teeth at the point that
faces downwards from the middle of the sea urchin’s body. It can be extended to some degree and
swung from side to side to some extent too. Sea urchins are mostly grazers of algae that covers rocks
Introduction 3
or of larger marine plants such as grasses and kelp though some will feed on corals while others
feed on minute food particles in sand. Sea cucumbers are deposit feeders that remove food particles
from sand or mud.
1.3.3 Digestive System
Echinoderms have a simple digestive system with a mouth, stomachs, intestine, and anus. In many,
the mouth is on the underside and the anus on the top surface of the animal. Sea stars can push their
stomachs outside of their body and insert it into its prey, allowing them to digest the food externally.
This ability allows sea stars to hunt prey that are much larger than their mouths would otherwise
allow.
1.3.4 Nervous System and Senses

Echinoderms do not have brains; they have nerves running from the mouth into each arm or along
the body. They have tiny eyespots at the end of each arm that only detect light or dark. Some of
their tube feet are also sensitive to chemicals and this allows them to find the source of smells, such
as food.
1.3.5 Circulatory System

Echinoderms have a network of fluid-filled canals that function in gas exchange, feeding, and in
movement. The network contains a central ring and areas that contain the tube feet, which stretch
along the body or arms. The tube feet poke through holes in the skeleton and can be extended or
contracted. Echinoderms do not have a true heart and the blood often lacks any respiratory pigment
(pike haemoglobin).
1.3.6 Respiratory System
Echinoderms have a poorly developed respiratory system. They use simple gills and their tube feet
to take in oxygen and pass out carbon dioxide.
1.3.7 Excretory System
Echinoderms have a simple excretory system with no kidneys and use diffusion to rid their bodies
of nitrogenous waste, which is mainly ammonia gas.
1.3.8 Reproductive System
Echinoderms are either male or female and become sexually mature after about 2–3 years. During
sexual reproduction, the eggs and sperm cells are typically released into open water, where fertiliza-
tion takes place. A female can release 100 million eggs at once. Larvae develop, which eventually
settle on the seafloor in their adult form. Usually, the echinoderms are nearly all gonochoric, though
a few species are hermaphroditic. Asexual reproduction by transverse fission has also been observed
in adult sea cucumbers (e.g., Holothuria parvula). Some sea stars and brittle stars have the ability to
reproduce asexually by dividing in two halves while they are small juveniles.
1.3.9 Regeneration Ability
The echinoderms are also known for their regeneration ability. If an arm breaks off of some echi-
noderms, a new arm or even a new echinoderm can regrow. When threatened or during asexual
reproduction, most sea cucumbers, starfishes, and sea lilies often intentionally lose parts of their
arms, which they can later regenerate. Sea urchins are constantly replacing spines lost by damage.
In most of these species, asexual reproduction is by transverse fission with the disc splitting in two.
Although in most species at least part of the disc is needed for complete regeneration, in a few spe-
cies of starfishes, such as Sclerasterias euplecta and Linckia columbiae, a single severed arm can
grow into a complete individual over a period of several months. Thus, an individual may have arms
of varying lengths.
1.3.10 Predators
Crabs, sharks, eels and other fish, sea birds, octopuses, and larger starfish are predators of echi-
noderms. Sea otters are keen on sea urchins, as are fish, starfish, and people. Sea urchin eggs are
eaten in many parts of the world, especially Japan. Starfish are eaten by fish such as some rays
and sharks. They are also eaten by other larger starfish and some predatory molluscs such as large
snails. Brittle stars can regenerate lost arms or arm segments and use this ability to escape preda-
tors, such as some gastropods, some fish, crabs, shrimps, and other echinoderms such as starfish.
1.3.11 Venomous/Poisonous Echinoderms
It is reported that about 2,000 species of the marine fauna are either venomous or poisonous and are
responsible for injuries to humans. Among the echinoderms, the sea urchins assume greater signifi-
cance as far as venomology is concerned. The venoms of some sea urchins have toxins and bioactive
molecules that produce toxicity effects on their victims by a variety of mechanisms. Thermolabile
toxins are reputed to be responsible. Immersion in hot water provides pain relief and promotes
resolution of symptoms. Comprehensive studies that led to the identification of pure toxins from the
crude venoms of these sea urchins are few and unfinished; therefore, it is important to do further
intensive studies on this field in the future. About 30 species of echinoderms as have been reported
to be harmful (James, 2010; Venkataraman et al., 2012).
Starfishes
Crown of thorns, Acanthaster planci
Starfish, Pentaeaster regulus
Plain sand star, Astropecten indicus
Biscuit sea star, Goniodiscaster scaber
Mosaic sea star, Plectaster decanus
Brittle stars
Chain-link brittle star, Ophiomastix annulosa
Sea urchins
Long-spined sea urchin, Diadema savigni
Long-spined black sea urchin, Diadema setosum
Collector urchin, Tripneustes gratilla
Flower urchin, Toxopneustes pileolus
Black sea urchin or long-spined sea urchin, Stomopneustes variolaris
Green sea urchin or variegated sea urchin, Lythechinus variegates
Burrowing sea urchin, Echinometra locunter
Long-spined urchin, Diadema paucispinum
Long-spined urchin, Diadema setosum
Introduction 5
Purple sea urchin or stony sea urchin, Paracentrotus lividus

Black Sea urchin, Arbacia lixula
Purple sea urchin or violet sea urchin, Sphaerechinus granularis
Hatpin urchin, Centrostepbanus longispinus
Banded sea urchin or double-spined urchin, Echinothrix calamaris
Deep-sea sea urchin, Araeosoma owstoni
Red Sea fire urchin, Asthenosoma marisrubri
Sea urchin, Phormosoma sp.
Sea lilies
Black and white sea lily, Tropimetra carinata
Sea cucumbers
Leopard sea cucumber, Bohadschia argus
Holothurian, Actinocucumis typicus
Brown sandfish or chalky cucumber, Bohadschia marmorata
Black sea cucumber or lollyfish, Holothuria atra
Sandfish or golden sandfish, Holothuria scabra
Brown sandfish, Holothuria spinifera
1.4 CHARACTERISTICS OF ECHINODERMS

1.4.1 Basic Features of Echinoderms
1. Pentametry is 5-rayed symmetry.
2. There is a water vascular system, a network of water-filled channels that often ends in suck-
ered tube feet (podia in large numbers) that enable them to move about and bring objects
to the mouth.
3. The endoskeleton is made of many calcareous (bony) pieces or ossicles.
4. There are no brain, heart, or eyes.
5. Reproduction can be sexual or asexual.
1.4.2 Characteristics of Different Classes of Echinoderms

There are five classes of echinoderms (extant), viz. Asteroidea, Ophiuroidea, Echinoidea,
Holothuroidea, and Crinoidea.
1.4.2.1 Class Asteroidea
1.4.2.1.1 Habitats
This class of echinoderms includes sea stars, which were previously known as starfish. There are
approximately 1,500 extant species in this class of echinoderms. The sea stars are found in the
ocean and at different depths. They can live in the coral reefs and on sand or rocks.
1.4.2.1.2 Anatomy
These free-moving organisms are composed of a central disc from which usually five arms radiate,
although some species may have more (up to 40). They show a bilateral symmetry during the larval
phase, which is lost during metamorphosis, developing radial symmetry, typically pentamerism in
adults. A pore called “madreporite” is located in the starfish body and this pore is responsible for
the entry of water in a hydraulic system, named “water vascular system,” which is made up of a
network of fluid-filled canals. This system helps in locomotion, adhesion, food manipulation, and
gas exchange. The mouth and anus are close together in the centre of the disc on the underside of
the body, together with the water intake pore (madreporite).
1.4.2.1.3 Food and Feeding

The majority of starfishes are carnivorous and feed on sponges, bryozoans, ascidians, mollusks,
bivalves, and snails. Others feed on detritus, eating decomposed organic material and fecal matter.
In the starfish feeding, their stomach is everted through the mouth opening over the prey, thus sur-
rounding the prey with the digestive organs. Digestive juices are secreted and the prey is liquefied.
The food is digested and together with the stomach is again sucked through the mouth opening into
the body.
1.4.2.1.4 Reproduction
The sea stars are able to reproduce by sexual or asexual reproduction. In the sexual stage, the sea
stars are simultaneous hermaphrodites, producing at the same time eggs and sperm. The eggs and
sperm are released into the water and the embryos and larvae lead a planktonic life or are housed in
rocks. In the asexual stage, the starfish may be able to reproduce by fission of their central discs or
by fission of one or more of their arms.
1.4.2.1.5 Regenerative Ability

The sea stars are well known for their regenerative ability. They are able to regenerate all the lost
arms or part of the central disc. The loss of parts of the body in sea stars can also occur as a protec-
tive function, as losing a body part may be to escape a predator (self-amputation) or occur during
asexual reproduction.
1.4.2.2 Class Ophiuroidea

1.4.2.2.1 Habitats
This class of echinoderms includes brittle stars and basket stars. It is the largest echinoderm class
with 2,100 species. Ophiuroids are dominant in many parts of the deep sea, where in certain regions
the bottom may swarm with brittle stars.
1.4.2.2.2 Anatomy
The brittle stars or serpent stars have highly flexible arms radiating from a central circular or pentag-
onal disc. The body outline is similar to the starfish, but the central disc is sharply marked off from
the arms and contains all the internal organs responsible for digestion and reproduction. The under-
side of the disc contains the mouth, with five jaws formed from skeletal plates. The madreporite is
Introduction 7
located within one of the jaw plates and not on the upper side of the animal, as it is in starfishes.
Through writhing movements of the arms, the brittle stars produce locomotion.

The ophiuroids are scavengers or detritivores and small organic particles, small crustaceans, and
worms are moved into the mouth by the tube feet. The digestive system is confined to the disc and
lacks an anus.
In most species the sexual individuals are separate, although a few are hermaphroditic. The gonads
are located in the disc, and the gametes are shed into the surrounding water.
1.4.2.2.5 Regenerative Ability

Brittle stars can easily regenerate lost arms or arm segments unless all arms are lost. Discarded
arms do not have the ability to regenerate. The ophiuroids use this capacity to escape predators
or reproduction. Some brittle stars, such as Ophiactis savignyi and Ophiocomella ophiactoides,
exhibit fissiparity with the disc splitting in half.
1.4.2.3 Class Echinoidea

1.4.2.3.1 Habitats
This class of echinoderms includes sea urchins and sand dollars, and there are about 1,000 species
in this group. These echinoids are distributed worldwide in marine habitats from the intertidal zone
to 5,000 m deep.
1.4.2.3.2 Anatomy
These echinoderms are usually globular, hemispherical, or disc-shaped. The skeletal plates, named
ambulacral areas, are arranged in meridional bands, which bear openings through which the ambu-
lacral feet protrude. The tube feet are moved by a water vascular system, allowing the sea urchin
to pump water in and out of the tube feet, enabling it to move. As sea urchins move slowly, they
feed mostly on algae. Surrounding the mouth, there is a circular opening where the skeletal plates
are replaced by a membrane termed the peristome. Normally, the anus is in the pole opposite to
the mouth in a region called the periproct. Around the periproct, the genital plates alternate with
the ocular plates, and one of the genital plates is modified to serve as a madreporite. Five teeth are
visible in the centre of the peristome, and the entire chewing organ is known as Aristotle’s lantern.
At the top of the lantern, a short esophagus is open, which leads into the stomach. The intestine
bends backwards in the opposite direction to that of the course of the stomach, and in the case of the
sea urchin this leads to a median dorsal anus, while in the sand dollars it passes along the posterior
interambulacrum to an anal opening either on or close to the margin of the disc.

Echinoids graze on algae, bryozoans, and dead animals.
The sexes are separate. In some species, gametogenesis is regulated by photoperiod so that spawning of
most or all members of a population occurs during the same time. Some female urchins brood their young
externally, within the protection of their spines or tube feet. In species with indirect development, an
echinopluteus larva is produced. Such a larva is bilaterally symmetrical and undergoes metamorphosis to
attain the pentaradial symmetry of the adult. The female’s eggs float freely in the sea and are fertilized by
free-floating sperm released by males. The fertilized eggs develop into a free-swimming blastula embryo
in as few as 12 hours, but several months are needed for the larva to complete its full development, which
begins with the formation of the test plates around the mouth and anus.
1.4.2.3.5 Predators
Members of this class are food for crabs, sea stars, fish, birds, otters, and other mammals.
1.4.2.3.6 Economic Importance

Echinoids of economic importance for the US are the red (Strongylocentrotus franciscanus), purple
(S. purpuratus), and green (S. droebachiensis) sea urchins. These urchins are harvested for their roe
and are exported to Japan.
1.4.2.3.7 Research Use

Probably the single most important contribution of these animals to scientific knowledge is their
embryological development. Researchers investigate the development of deuterostomes using sea
urchin eggs.
Introduction 9
1.4.2.4 Class Holothuroidea

1.4.2.4.1 Habitats
There are about 1,100 species of sea cucumbers. About 70 species of sea cucumbers have been
exploited worldwide, out of which 11 species have been found to be commercially important. Sea
cucumbers are found in nearly every marine environment but are most diverse on tropical shallow-
water coral reefs. Their habitats range from the intertidal, where they may be exposed briefly at low
tide, to the floor of the deepest oceanic trenches.
1.4.2.4.2 Anatomy
The sea cucumbers are elongated echinoderms without a definite skeleton and pentaradial symme-
try, with a mouth at one extremity surrounded by a circle of branched tentacles and an anus at the
opposite extremity. Typically, the body is five sided and on each side bears a double row of tube feet,
used in locomotion. The body wall is highly muscular. The alternate use of longitudinal and circular
muscles enables the cucumber to creep like a worm. Although there is no continuous skeleton, the
body wall is rather firm, and this is in large measure due to the presence of microscopic calcare-
ous plates embedded in the tissues. In some species, a calcareous ring of ten plates surrounds the
esophagus and serves as a support for the tentacles.

The diet of most cucumbers consists of plankton and decaying organic matter found in the sea.
The digestive canal is held in definite position by mesenteries. The esophagus loads into a stomach,
which is then followed by a tubular intestine. From the walls of the cloaca, there is usually a pair
of minutely branched respiratory trees which, by the muscular action of the cloaca, are filled with
water and serve as respiratory organs.
1.4.2.4.4 Evisceration
Sea cucumbers expel their internal organs as a defence mechanism called evisceration, a reaction
that includes release of the respiratory tree, intestine, cuvierian tubules, and gonads through the anal
opening. It is hypothesized that the reason for this ingenious form of defence is because these organs
contain high levels of compounds that repel predators.
The sea cucumbers are dioecious with separate male and female individuals, which reproduce by
releasing sperm and eggs into the ocean water. The reproductive system consists of a single gonad,
consisting of a cluster of tubules emptying into a single duct that opens on the upper surface of the
animal, close to the tentacles. A larval form known as an auricularia is produced in the development
stages of the embryo.
1.4.2.5 Class Crinoidea

1.4.2.5.1 Habitats
The class Crinoidea includes feather stars or sea lilies. There are more than 600 species of crinoids
of which around 80 are sea lilies. Crinoids are the oldest of the living echinoderms, with a fossil
record stretching back 450 million years. These sessile organisms are usually found living in groups
of several thousand. Feather stars swim slowly by flapping their feathery arms through the water, or
they crawl along the ocean floor to find food. They often use rocks, corals, or sponges to raise them-
selves above the bottom when they feed and hide in caves and ledges during the day. Feather stars
are found in shallow and deep ocean waters but are most common in tropical reef environments. Sea
lilies live attached to the ocean floor in depths greater than 100 m.
1.4.2.5.2 Anatomy
Crinoids have an external skeleton made of calcium carbonate plates covered by a thin skin. The
plates are held together with ligaments or muscles. Shallow water forms are usually very colourful.
The skeleton is usually divided into four basic parts: the holdfast, a disc-like sucker, which anchors
the crinoid to the ocean bottom; the stem, filled with muscles, which raises the calyx above the
substrate; the calyx, a cup-shaped central structure, which contains the internal organs; and the
arms, which are from 5 to as many as 200 feeding arms (in multiples of five). The largest sea lily
(Metacrinus superbus) has a large calyx, which with its arms gives it a diameter of 1.5 m. The larg-
est feather star (Heliometra glacialis) has an arm span of 35 cm. The smallest crinoids are around
3 cm in diameter.

Most species are nocturnal filter/suspension feeders that consume plankton and decaying organic
matter. To feed, they spread their feeding arms to sieve the passing seawater for microscopic
organisms and detritus. Mucus on the tube feet traps their food, which is passed down the arms
into the mouth by beating cilia. Crinoids have a U-shaped digestive system with the anus next to
the mouth.
1.4.2.5.4 Predators
Fish and other echinoderms (especially sea urchins) are the known predators of crinoids. Sea lilies
have also been reported to crawl away from sea urchins.
Introduction 11
Crinoids have male and female individuals but have no true gonads, producing their gametes from
genital canals. The eggs and sperm are released into the surrounding seawater. The fertilized eggs
hatch, resulting in the formation of a free-swimming ciliated larva, which does not feed, and it lasts
only for a few days before settling in the bottom of the sea using an adhesive gland on its ventral
surface. The larva then metamorphoses into an adult in 8–12 months.
1.5 CONSERVATION MEASURES OF ECHINODERMS

In order to safeguard the diversity and biomass of echinoderms for sustainable fisheries, the follow-
ing measures need to be undertaken (Micael et al., 2009).
1. Controls of the management of echinoderm fisheries: These include closed seasons during
times of spawning; gear restrictions; designation of no-take, marine-protected areas; daily
catch limits; minimum legal size; prohibition of night fishing for nocturnal species; and
restrictions on the use of scuba for harvesting.
2. Rotational fishing is used as an appropriate harvest strategy for echinoderm fisheries.
3. The echinoderm biodiversity is ensured through the conservation of natural habitats and
wild fauna and flora.
4. Existing conservation mechanisms must be integrated and applied consistently on a global
scale.
5. Echinoderm fisheries management requires an ecosystem approach by which there is
improved information sharing among government agencies, nongovernmental organiza-
tions and academia, and a dialogue between all users, including the industry and commu-
nities dependent on echinoderm resources.
The development and integration of some of the suggested measures and the consideration of the
different spatial scales (local, regional, and global) will permit the sustainable use of echinoderm
species and conservation of this precious resource. Further, there is a clear need to improve our
biological knowledge about the target species to ensure that the diversity of this wonderful group of
marine animals is maintained. The following adage will also hold good for echinoderms.
In the end, we will conserve what we love; we will love only what we understand; and we will under-
stand only what we are taught
Baba Dioum
1.6 IMPORTANCE OF ECHINODERMS

Echinoderms offer important benefits to human beings. They are important nutritionally, ecologi-
cally, scientifically, medicinally, and aesthetically. Sea urchins are a food item for humans, which
are consumed either raw or briefly cooked. Sea urchin “roe” (the gonads) is a popular food in Korea,
Japan, Chile, and Spain. Apart from domestic consumption, Chile and a number of other countries
export the sea urchin to Japan in order to meet its demand. Similarly sea cucumber is considered a
delicacy in Far East countries such as Malaysia, China, Japan, and Indonesia. It is highly valued for
its supposed medicinal properties.
Ecologically, echinoderms play a key role in food webs. In the case of sea cucumbers, these are
typically scavengers, feeding on debris in the benthic layer, and consuming plankton and other
organic matter found in the sea. A variety of fish, most commonly pearl fishes, have evolved a
symbiotic relationship (commensalism) with sea cucumbers in which the pearl fish will live in the
sea cucumber’s cloaca, using it for protection from predation, a source of food (the nutrients passing
in and out of the anus from the water), and to develop into their adult stage of life. In the case of
starfish, their place in the food web also impacts the world economically, as they feed on oysters,
clams, and other organisms that humans consume.
Scientifically, the sea urchin occupies a special place in biology due to its long-time use as a
standard subject for studies in embryology.
Medicinally, some varieties of sea cucumber (known as gamat in Malaysia) are said to have excellent
healing properties. There are pharmaceutical companies being built based on this gamat product. Extracts
are prepared and made into oil, cream, or cosmetics. Some products are intended to be taken internally.
Claims have been made that sea cucumber helps a wound heal more quickly and reduces scarring.
Aesthetically, the diverse forms of the echinoderms and their sometimes brilliant colouring are
often a source of joy to humans observing them.
1.6.1 Echinoderms in Commercial Fisheries

Of the five extant classes of echinoderms, only sea urchins and sea cucumbers are heavily exploited (Kelly,
2005). There is a limited, localized commercial fishery for sea stars but reported global landings are small,
unreliable, and strongly dominated by the fishery for Asterias rubens in Denmark (Sloan, 1985). The latter
sea star is exported to West Germany as an additive to finfish meal for poultry feedstocks (Sloan, 1985). In
the United States, sea star fisheries are uneconomical due to high production costs and low product quality
(Loosanoff, 1961). Although Sloan (1985) considered that it was unlikely that sea stars would become an
object of appreciable new meal fisheries, there is an emerging demand for them in the Asian food markets
but, unfortunately, the extent of this demand is not documented (Micael et al., 2009).
Male and female sea urchins are collected for their gonads, referred to as “roe” (Kelly, 2005), a
food delicacy that fetches high prices in Asian and Mediterranean markets (Sloan, 1985). Individuals
are picked off the seabed by hand by divers operating in near-shore waters or are collected using
dredges (Bergman and van Santbrink, 1994). The quantity and quality of roe are vital to the market
and are considered critical to the profitability of the processing operation. Because these roe attri-
butes are not apparent externally, animals need to be cracked open, resulting in a high level of
mortality and many discarded individuals (Kalvass and Hendrix, 1997). There are established fisher-
ies in Chile; northern Asia (Japan and Korea); Maine, California, and Alaska (United States); Baja
California (Mexico); Russia; and British Columbia, New Brunswick, and Nova Scotia (Canada)
(Andrew et al., 2002). Japan consumes more than 80% of the world’s catch of sea urchin roe (Sonu,
1995; Hagen, 1996), followed by France (Hagen, 1996). Different sea urchin species are the tar-
get in different fishery regions (Table 1.1). World catches peaked in 1995 with global landings of
113,654 tons (t), representing a threefold increase in production over 25 years (Williams, 2002). As
a supplier market, Chile landed some 54,600 tons in 1995 making it the world’s largest producer
of sea urchins, but this level of production was regarded as unsustainable (Jimmy et al., 2003).
Smaller fisheries in Europe supply domestic markets in the main but Portugal, without a local
market, exported 15 tons of sea urchins in 2002 to the United States and Japan (Instituto Nacional
de Estatistica, 2003). The fishery statistics (Table 1.1) clearly demonstrate that most of the world’s
urchin fisheries are fully or overexploited, and it is generally accepted that further sea urchin fishing
grounds are unlikely to be discovered (Keesing and Hall, 1998).
World catches of sea urchins peaked in 1995 with global landings of 117,100 tonnes represent-
ing a threefold increase in production over twenty-five years (Williams, 2002). As a supplier mar-
ket, Chile landed 54,600 tons in 1995 making it the world’s largest producer of sea urchins, but this
level of production was regarded as unsustainable (Jimmy et al., 2003). Smaller fisheries in Europe
Introduction 13
supply domestic markets in the main but Portugal, without a local market, exported 15 tons of sea
urchins in 2002 to the United States and Japan (Instituto Nacional de Estatistica, 2003). The sea urchin
landings (along with their contributing species) of the different regions of the world are given below.
TABLE 1.1
World Sea Urchin Landings (as of 1995)
Region Contributing Species Landings (tons)
Northwest Atlantic Strongylocentrotus droebachiensis 18400
Northeast Pacific S. franciscanus 21500
S. purpuratus
S. droebachiensis
Japanese Islands S. intermedius 13700
S. nudus
Tripneustes gratilla
Pseudocentrotus depressus
S. pulcherrimus
Anthocidaris crassispina
South America Loxechinus albus 54700
Korean Peninsula 3800
Southeast Asia Anthocidaris crassispina 700
South Pacific Heliocidaris erythrogamma 1000
S. rodgersii
Evechinus chloroticus
Tripneustes gratilla
Northern Europe S. droebachiensis 3200
S. intermedius
S. nudus
S. polyanthus
Southern Europe Paracentrotus lividus 100
Total 117100
Source: Keesing and Hall, 1998.
The sea urchin fisheries statistics below clearly demonstrate that most of the world’s urchin fish-
eries are fully or overexploited, and it is generally accepted that further sea urchin fishing grounds
are unlikely to be discovered (Keesing and Hall, 1998).
1.6.1.1 Sea Urchin Species with a Major Contribution to the Modern Fishery
Species Fishery Distribution
Anthocidaris crassipina Japan, Korea, China
Centrostephanus rodgersii Australia, New South Wales
Echinometra spp. Circumpolar–Caribbean
Evechinus chloroticus New Zealand
Glyptocidaris crenulatus China
Heliocidaris erythrogramma New South Wales
Heliocidaris tuberculata New South Wales
Hemicentrotus pulcherrimus Japan, Korea, China
Loxechinus albus Chile, Peru
Lytechinus variegates West Atlantic, Caribbean
(Continued)

Paracentrotus lividusa Atlantic, Mediterranean
Pseudocentrotus depressus Japan, Korea
Strongylocentrotus droebachiensis Circumpolar–North
Strongylocentrotus franciscanus Chile, Mexico, Northeast Pacific
Strongylocentrotus intermedius Japan, Russia, Korea
Strongylocentrotus nudus Japan, China
Strongylocentrotus pallidus Russia
Strongylocentrotus polyacanthus Russia
Strongylocentrotus purpuratus Mexico, Northeast Pacific
Tripneustes esculentus Circumpolar–Caribbean
Tripneustes gratilla Indian Ocean
Source: Micael et al., 2009.
About 42 species of sea cucumbers (“bêche-de-mer” or “trepang”) are fished commercially

(Bruckner, 2005; Conand, 2005) for their proteinaceous body wall, which is boiled and dried.
The Chinese have imported sea cucumbers for more than 1,000 years from India, Indonesia, and
the Philippines, but traders began gathering them from a wider area in the eighteenth and nine-
teenth centuries (Conand and Byrne, 1993). During the 1990s, the number of producing countries
(and species exploited) increased worldwide and holothurian fisheries became established in many
nontraditional fishing areas such as Mexico, the Galapagos, and North America (Bruckner et al.,
2003). Growing evidence indicates that sea cucumber populations are declining worldwide, includ-
ing Australia, India, Madagascar, Thailand, Papua New Guinea, and the Galapagos.
1.6.1.2 Sea Cucumber Species that Contribute to the Modern Fishery

Actinopyga echinites South Pacific
Actinopyga lecanora South Pacific
Actinopyga mauritiana South Pacific
Actinopyga miliaris South Pacific
Apostichopus japonicas Pacific
Bohadschia argus South Pacific, Southeast Asia
Bohadschia graeffei South Pacific, Southeast Asia
Bohadschia marmorata marmorata Southeast Asia, South Pacific, Red Sea
Bohadschia marmorata vitiensis Southeast Asia, South Pacific, Red Sea
Bohadschia vitiensis South Pacific, Indian Ocean
Halodeima atra South Pacific
Halodeima edulis South Pacific
Halodeima fuscogilva South Pacific, Southeast Asia, India
Halodeima nobilis South Pacific, Southeast Asia
Halodeima scabra South Pacific, Southeast Asia, India
Halodeima scabra versicolor South Pacific, Southeast Asia
Holothuria fuscocinerea Pacific
Holothuria nobilis Pacific
Holothuria pervicax Pacific
Holothuria scabra Pacific
Isostichopus fuscus Indo and eastern Pacific
Parastichopus californicus Canada
(Continued)
Introduction 15

Stichopus chloronotus South Pacific, Indian
Stichopus hermanni Southeast Asia, South Pacific
Stichopus variegates Pacific
Thelenota ananas South Pacific
Thelenota anax South Pacific
1.6.2 Echinoderms in Aquaculture (Farming)

Among the echinoderms, sea urchins and sea cucumbers are widely cultivated as detailed below
Species Farming Countries
Sea Urchins
Strongylocentrotus droebachiensis East and west coasts of North America, Norway, China and Russia
Strongylocentrotus franciscanus West coast of North America
Strongylocentrotus purpuratus West coast of North America
Lytechinus variegates West coast of North America
Loxechinus albus Chile
Paracentrotus lividus Israel, Southern Ireland, France, Scotland
Evechinus chloroticus New Zealand
Echinus esculentus Scotland
Psammechinus miliaris Scotland
Centrostephanus rodgersii Australia
Heliocidaris erythrogramma Australia
Tripneustes gratilla Australia, Philippines
Sea Cucumbers
Apostichopus japonicas China, Korea, Russia
Holothuria scabra Australia, India, Indonesia, Maldives, Philippines, Solomon Islands, Malaysia,
Saudi Arabia, Vietnam
Holothuria fuscogilva Australia
Actinopyga Mauritania Indonesia
Source: Rahman, M.A. et al., Sea Urchins (Echinodermata: Echinoidea): Their biology, culture and bioactive compounds,
Proceedings of International Conference on Agricultural, Ecological and Medical Sciences, pp. 39–48, 2014a; Rahman,
M.A., Culture potentials of sea cucumbers (Echinodermata: Holothuroidea) and their biomedical applications,
Proceedings of International Conference on Chemical, Biological, and Environmental Sciences, pp. 46–50, 2014b.
1.6.3 Echinoderms as Food
In some countries, echinoderms are considered delicacies. Around 50,000 tons of sea urchins are
captured each year for food. They are consumed mostly in Japan, Peru, Spain, and France. Both
male and female gonads of sea urchins are also consumed. The taste is described as soft and melt-
ing, like a mixture of seafood and fruit. Sea cucumbers are considered a delicacy in some southeast-
ern Asian countries. In China, they are used as a basis for gelatinous soups and stews.
From a nutritional point of view, holothurians are considered a valuable food product, as they
contain nutrients that are important for human physiology.
The results show that the average ingredients value nutrition of fresh sea cucumber is as follows:
protein (21%–44.07%), fat (1.01%–1.19%), carbohydrate (0.5%–2.34%), ash (2.01%–3.07%), water
(76.03%–79.43%) (Oedjoe, 2017).
Sea cucumbers, informally named as bêche-de-mer or gamat, have long been used for food and
folk medicine in the communities of Asia and the Middle East. Nutritionally, sea cucumbers have
an impressive profile of valuable nutrients such as Vitamin A, Vitamin B1 (thiamine), Vitamin B2
(riboflavin), Vitamin B3 (niacin), and minerals—especially calcium, magnesium, iron, and zinc—
with potential uses for functional foods and nutraceuticals.
1.6.4 Echinoderms in the Aquarium and Souvenir Trades

Many marine invertebrates, including echinoderms, are popular in the aquarium trade. Mexico,
Indonesia, Singapore, Fiji, Sri Lanka, Philippines, and Vanuatu are the main exporters, accounting
for close to 17% of the global trade (Wabnitz et al., 2003). The most commonly collected sea star in
the aquarium trade is Linckia laevigata, which is taken mainly from the shallow waters of the tropical
Indo-Pacific (Shimek, 2005). According to the Global Marine Aquarium Database (GMAD), this spe-
cies accounts for 3% of the total global trade in marine invertebrates. Almost all L. laevigata are taken
from the wild, with few resulting from captive breeding (Wabnitz et al., 2003). Calado (2006) identified
11 echinoderm species in Portuguese waters as potential target species for the marine aquarium trade.
Little is known of the true extent of the global use of echinoderms as souvenirs, although dried
sea star candleholders, toy cars sporting sea urchins as wheels, and sand dollar necklace pendants
are seen frequently in souvenir shops.
1.6.4.1 Potential Echinoderms in the Ornament Industry
Class Species
Asteroidea Echinaster sepositus
Ophidiaster ophidianus
Ophiuroidea Ophioderma longicauda
Echinoidea Arbaciella elegans
Brissus unicolor
Centrostephanus longispinus
Cidaris cidaris
Diadema antillarum
Echinocardium cordatum
Spatangus purpureus
Crinoidea Antedon bifida
1.6.5 Echinoderms in Scientific Research (Experimental Model Systems)

Echinoderm embryos and larvae have been used as experimental model systems in several lines of
research for more than a century. Research on echinoderm embryos has led to significant advances
in the areas of developmental biology, cell biology, and immunology. Evolutionary biologists have
used echinoderms to test theories of life history evolution (Raffaelli, 2006). Congenic pairs of
sea urchin species, one exhibiting direct and the other indirect development, have been used to
investigate the developmental basis for changes in life history patterns (Amemiya et al., 2005).
Echinoderms are well-known for their striking regenerative capacity; asteroids and crinoids can rap-
idly and completely regenerate arms following self-induced or traumatic amputation (Clark, 1968).
Introduction 17
These echinoderm groups in particular, therefore, have provided a valuable experimental model to
investigate the regenerative process from the macroscopic to the molecular level (Carnevali and
Bonasoro, 2001) and for the identification of the genes involved in the process of neural regen-
eration (Thorndyke et al., 2001; Kelly, 2005). Echinoderm regeneration also provides a convenient
model for examining the effects of persistent micropollutants on the developmental physiology
(cell proliferation, morphogenesis, differentiation, tissue renewal) of marine animals (Kelly, 2005).
Regenerative medicine may benefit significantly from the extensive study of echinoderm models in
parallel with traditional mammal models, in the reasonable hope that what echinoderms can do so
easily may eventually become easy also for other animals, humans included.
1.6.6 Echinoderms in Bionics
The brittle star Ophiocoma wendtii possess single calcite crystals arranged to function as lenses.
These lenses focus light on to nerve bundles that run behind them and that presumably receive the
signal to be further processed. In total, thousands of lenses form a compound eye that covers
the upper surface of the animal, resulting in a function similar to a digital camera that builds up the
picture pixel by pixel (Aizenberg et al., 2001; Petzelt, 2005). At present, engineers in the photonic
industry are trying to imitate the perfect calcite lenses and their use in signal reception application
of biological methods and systems found in nature and apply them to the study and design of engi-
neering systems and modern technology.
1.6.7 Echinoderms in Bioindicators of Environmental Quality

Echinoderms are prime candidates for model toxicological test organisms for the marine ecosystem
for many reasons, including their ubiquitous distribution, their benthic and relatively sedentary life-
style, their susceptibility to the presence of micropollutants stored in marine sediments, and their
sensitivity to many types of contaminants. Echinoderms can be considered key organisms in both
basic and applied research in this field and can be usefully employed for developing new successful
experimental approaches and strategies. Sea urchin gametes, embryos, and larvae can be used for
fast, low-cost, and reliable screening and testing of toxic substances and for detailed studies of their
mechanism of action.
Sea urchins are also model organisms used in developmental biology research. They have been
used to study the mechanisms of fertilization and egg activation, physiological processes that occur
during early development, and the regulation of differentiation in the early embryo. In addition, the
molecular basis of early development was studied in sea urchins. Gametes can be obtained easily,
sterility is not required, and the eggs and early embryos of many commonly used species are beau-
tifully transparent. In addition, the early development of sea urchin embryos is a highly conserved
process. When a batch of eggs is fertilized, all of the resulting embryos typically develop at the
same time. This makes biochemical and molecular studies of early embryos possible in the sea
urchin and has led to a number of major discoveries.
1.6.8 Ecological Role of Echinoderms

Echinoderms play numerous ecological roles. Sand dollars and sea cucumbers burrow into the sand,
providing more oxygen at greater depths of the seafloor. This allows more organisms to live there.
In addition, starfish prevent the growth of algae on coral reefs. This allows the coral to filter-feed more
easily. And many sea cucumbers provide a habitat for parasites such as crabs, worms, and snails.
Echinoderms are also an important step in the ocean food chain. They are the staple diet of many
animals, including the sea otter. On the other hand, echinoderms eat seaweed and keep its growth in
check. Recall that the sea urchin is a grazer, mainly feeding on algae on the coral and rocks. Recently,
some marine ecosystems have been overrun by seaweed. Excess seaweed can destroy entire reefs.
Scientists believe that the extinction of large quantities of echinoderms has caused this destruction.
Most commercial sea cucumbers are detritivores, and the bioturbation they cause during feeding
plays a key role in nutrient cycling within the marine ecosystem (Bakus, 1973; Uthicke and Klumpp,
1998; Uthicke, 1999, 2001). In addition, sea cucumbers consume and grind sediment and organic matter
into finer compounds, turning over the top layers on the seabed, allowing oxygen to enter the sediment.
Basically, they are responsible for the extensive shifting and mixing of the substratum and recycling of
detrital matter (Bruckner et al., 2003). This constant cycling prevents the accumulation of organic matter
and may help control pathogens. Sea cucumbers such as Isostichopus badionotus rework the sediment
(Webb et al., 1977) and are important in determining habitat structure for other species; their removal
may result in the loss of these other benthic species (Lovatelli et al., 2004; Bruckner, 2005). In some
ecosystems, sea cucumbers represent a substantial portion of the ecosystem biomass (Bruckner et al.,
2003) and their eggs, larvae, and juveniles constitute an important food source for other marine species,
including crustaceans, fishes, and molluscs; rapid declines in sea cucumber populations may have seri-
ous consequences for the survival of other species that are part of the same complex food web. Finally,
several species of sea cucumbers have unique symbionts such as bacteria, protozoans, and metazoans,
including molluscs and fishes, which may disappear on removal of the host (Micael et al., 2009).
1.6.9 Echinoderms as Medicine
Echinoderms are also used in traditional medicine and in scientific research. For example, some
sea cucumber toxins slow down the growth rate of tumour cells, so there is an interest in using
these in cancer research. Traditional Chinese medicine commonly uses sea cucumbers in treating
weakness, impotence, debility of the aged, constipation due to intestinal dryness, frequent urina-
tion, and joint problems. Western medicine is using sea cucumbers to treat rheumatoid arthritis,
osteoarthritis, ankylosing spondylitis, and connective tissue disorders. Australia has approved
the use of sea cucumbers as an over-the-counter treatment for arthritis, and the Japanese have a
patent using sea cucumber chondroitin sulphate for HIV therapy. Sea cucumbers have been well
recognized as a tonic and traditional remedy in Chinese and Malaysian literature for their effec-
tiveness against hypertension, asthma, rheumatism, cuts and burns, impotency, and constipation.
1.6.10 Echinoderms as Sources of Pharmaceutical (Bioactive)

Compounds and in the Development of New Drugs
Echinoderms are exclusively marine. They represent an exceptional source of polar steroids of
an immense structural diversity, showing a range of biological activities. The steroids are organic
compounds that act as an integral part of the cell membrane. The steroidal components, namely
saponins, asterosaponins, and astropectenol are the major source of compounds abundantly found
in sea stars [1–3]. A basic study of these facts reveals the search for “drugs from the sea” pro-
gresses at the rate of a 10% increase in new compounds per year [4]. The isolation and charac-
terization of bioactive compounds from the sea stars in the marine ecosystem is serving a good
resource for the human population to fight against the deadly diseases such as cancer (Sumithaa
et al., 2017a).
The bioactive compounds derived from echinoderms are compounds of interest showing an
extensive application in the treatment of many diseases. Those compounds showed several biologi-
cal properties, such as antibacterial, anticoagulant, antifungal, antimalarial, antiprotozoal, antitu-
berculosis, anti-inflammatory, antitumour, anti-HIV, and antiviral activities.
The importance of echinoderms as a potential source of bioactive compounds for the develop-
ment of new therapeutic drugs/agrochemicals has been growing rapidly. Compounds isolated from
the different species of echinoderms showed numerous biological activities including antibacterial,
Introduction 19
anticoagulant, antifungal, antimalarial, antiprotozoal, anti-tuberculosis, anti-inflammatory, antitu-

mour, and antiviral activities.
Triterpene glycosides, Steroids, Saponins, Peptides, Sphingolipids and fatty acids, Carotenoids,
quinones, spinochromes, and pigment (Gomes et al., 2014).
Gymnochrinus richer—gymnochrome B, D, and isogymnochrome D—antiviral
Celerina heffernani—ptilomycalin A, celeromycalin—antiviral
Fromia monolis—fromiamycalin, crambescidin 800—antiviral
Rosaster sp.—(25S)-5α-cholestane-3β, 4β, 6β, 7α, 8, 15α, 16β, 26-octol—antifungal
1.6.10.1 Present Status of New Natural Products (NP) as

Pharmaceutical Compounds from Echinodermata
Taxon Total No. Species No. Species with New NP % of Species with NP
Phylum Echinodermata 7353 153 2
Subphylum Asterozoa 4018 72 2
Class Asteroidea 1849 74 4
Order Forcipulatida 273 22 8
Order Spinulosida 131 7 5
Order Valvatida 733 25 3
Subphylum Echinozoa 2890 58 2
Class Holothuroidea 1800 51 3
Order Dendrochirotida 794 25 3
Source: Leal, M.C. et al., PLoS One, 7, e30580, 2012.
1.6.10.2 Pharmaceutical (Bioactive) Compounds from Sea Stars

Taxon Compounds Activity
Leptasterias ocholensis Asterosaponins & Glycosides Cytotoxic activity
Ctenodiscus crispatus Steroidal compounds Cytotoxic and antitumour activities
Ophiocoma erinaceus Saponins Haemolytic and cytotoxic activities
Luidia maculata — Antioxidant and antimicrobial activities
Astropecten polyacanthus Asteropectinol Cytotoxic activity
Protoreaster linckii Crude compounds Antimicrobial, haemolytic, antinociceptive, and
cytotoxic activities
Stellaster equestris Steroidal compounds Antibacterial
Acanthaster planci Glycoprotein Cytotoxic and apoptotic effect on human breast cancer
Astropecten indicus Crude compound Antibacterial
Archaster typicus Archasterosides & Regularosides Cytotoxic activity against human and mouse myeloma
Asterina pectinifera Polysaccharides Antitumour against colon cancer
Asterina pectinifera Crude compounds Antifungal activity
Culcita novaeguineae Asterosaponins Cytotoxic and antitumour activities
Asterias rubens Coelomocytes fluid (peptides) Antibacterial activity
Asterias amurensis Glycosides Promotes osteoblastic proliferation
Anasterias minuta Steroidal glycosides Antifungal activity
Dermasterias imbricate Sulphated steroidal compounds Antifungal activity
Anthenea chinensis Anthenoside A Cytotoxicity against human tumour cell lines
Lysastrosoma anthosticta Lysaketotriol and lysaketodiol Immunomodulatory activity
Source: Sumithaa et al., 2017a.

1.6.10.3 Pharmaceutical (Bioactive) Compounds from Brittle Stars

Ophiocoma scolopendrina Ophiodilactones A and B Cytotoxic activity against P388 murine leukemia cells.
Ophiocoma erinaceus Dichloromethane extract Cytotoxic and apoptosis inducing potential against melanoma
cancers
Ophiocoma erinaceus Crude extract Antioxidant and anti-inflammatory properties
Ophiocoma erinaceus Methanol extract Antiangiogenic effects
Source: Gomes, A.R. et al., RSC Adv., 4, 29365–29382, 2014.
1.6.10.4 Pharmaceutical (Bioactive) Compounds from Sea Urchins

Strongylocentrotus droebachiensis Peptides, Centrocins 1 and 2 Antimicrobial activity
Strongylocentrotus droebachiensis Spinochromes Antioxidant and anti-allergic
activities
Strongylocentrotus nudus Naphthoquinone pigments Antioxidant activity
Glyptocidaris crenularis N-acyl taurine Cytotoxicity
Diadema setosum PUFA Functional food
Salmacis sphaeroides PUFA Functional food
Source: Gomes, A.R. et al., RSC Adv., 4, 29365–29382, 2014; Rahman, M.A. et al., Proc. Int. Conf. Agricultural, Ecological
and Medical Sciences, 2014a, doi:10.15242/ IICBE.C714075.
1.6.10.5 Pharmaceutical (Bioactive) Compounds from Sea Cucumbers

Sea Cucumber Species Bioactive Compounds Activity
Pentacta quadrangularius and Cucumaria frondosa Philinospides A, E Antiangiogenic
Holothuria scabra, Holothurialeucos pilota, Stichopus Frondoside A, Okhotosides B1-B3, Anticancer
chloronotus, Cucumaria frondosa, Cucumaria okhotensis, Intercedensides A-C, Frondanol A,
Mensamaria intercedens, Pearsonothuria graeffei, Stichopus Holothurin A,
japonicus, and Stichopus variegates 24-dehydroechinoside, and
Frondanol(R)-A5p
Ludwigothurea grisea and Thelenota ananas Fucosylated chondroitin sulphate Anticoagulant
Stichopus japonicas Polypeptides, acidic Anti-fatigue
mucopolysaccharides, collagen,
and bioactive amino acids
Actinopyga echinites, Actinopyga miliaris, Holothuria atra, Phosphate-buffered saline, Antimicrobial
Holothuria scabra, Bohadshia argus, Cucumaria frondosa, Hemoiedemosides A,B
Holothuria poli, Hemoiedema spectabilis, Psolus Patagonicoside A, Holothurin B,
patagonicus, Actinopyga lecanora, Holothuria atra, Psolus Marmoratoside A, 17α-hydroxy
patagonicus, Bohadschia marmorata, and Cucumaria impatienside A, Impatienside A,
frondosa and Bivittoside D
Cucumariafrondosa, Stichopus japonicus, Paracaudina chilensis, Gelatin hydrolysate, protein Antioxidation
, Cucumaria frondosa, Holothuria scabra, Holothuria hydrolysate, phenols, flavonoids,
leucospilota, Stichopus chloronotus, and Acaudina molpadioides and collagen polypeptides
Stichopus japonicas Glycosaminoglycan Antithrombotic
(Continued)
Introduction 21
Sea Cucumber Species Bioactive Compounds Activity

Mensamaria intercedens, Mensamaria intercedens, Intercedensides D–I, Hillasides A,B, Antitumour
Holothuria hilla, Pentacta quadrangularis, Holothuria Philinopside A Holothurinosides
forskali, Stichopus japonicus, Holothuria nobilis, Holothuria A-D, Desholothurin A, Nobilisides
fuscocinerea, Stichopus japonicus, Holothuria impatiens, A-C, Fuscocinerosides A-C,
Ludwigothurea grisea, and Cucumaria japonica Impatienside A, and Cumaside
Staurocucumis liouvillei Liouvillosides A, B Antiviral
Stichopus japonicas Fucan sulfate, Glucosamin, and Osteoarthritis
Chondroitin
Thyone briareus, Stichopus chloronotus, Stichopus herrmanni, Arachidonic acid, eicosapentaenoic Wound healing
Thelenota ananas, Thelenota anax, Holothuria fuscogilva, acid, and docosahexaenoic acid
Holothuria fuscopunctata, Actinopyga mauritiana,
Actinopyga caerulea, Bohadschia argus, Stichopus
chloronotus, Holothuria tubulosa, Holothuria polii, and
Holothuria mammata
Source: Bordbar, S. et al., Mar. Drugs, 9, 1761–1805, 2011.
1.6.10.6 Pharmaceutical (Bioactive) Compounds from Sea Lilies

Himerometra magnipinna 12-hydroxyhomoaerothionin, aerothionin and Inhibition assay in Streptomyces
crinemodin-rhodoptilometrin bianthrone
Proisocrinus ruberrimus Proisocrinins A–F No bioactivity
Holopus rangii Gymnochromes E and F Cytotoxic activity
Source: Gomes, A.R. et al., RSC Adv., 4, 29365–29382, 2014.
It has been reported that of the total natural compounds discovered so far from echinoderms, only
50% of the compounds were associated with some sort of bioactivity. For the remaining 50% of
compounds, their bioactivity has not yet been either studied or reported. Further, the most studied
bioactive compounds of this phylum were the triterpene glycosides and steroids, showing antifun-
gal activity and cytotoxicity against human tumour cell lines as the main biological properties.
Furthermore, among the different classes of Echindermata, a higher number of new natural com-
pounds has been isolated from only starfishes and sea cucumbers, which seem to be more popular
among researchers, probably due to the bioprospecting studies (Gomes et al., 2014). Further studies
should therefore be pursued on less studied species of Echinoidea, Crinoidea, and Ophiuroidea in
order to screen and search for other new potential bioactive compounds.
2 Biology and Ecology of
Pharmaceutical Sea Stars
(Class: Asteroidea)
2.1 FAMILY: ASTERIIDAE (CLASS: ASTEROIDEA; ORDER: FORCIPULATIDA)

Anasterias antarctica (Lütken, 1857) (= Anasterias minuta)
Common name(s): Cinderella starfish, subantarctic sea star
Global distribution: Polar—Southern Ocean; Temperate South America
Ecology: This benthic species is found with the brown algae, giant kelp (Macrocystis pyr
ifera), and on hard substrates such as rocks, cobbles, and boulders in intertidal and subtidal
areas to a depth of 190 m.
Biology
Description: Abactinal plates of this species are slender, delicate, and form an irregu-
lar reticulum, with very large meshes. Dorsal spinelets are few and are widely scattered.
Numerous, rather thickly lanceolate, subobtuse straight pedicellariae, decidedly longer
than broad, are scattered on the marginal and actinal plates in the intermarginal channel
and along edge of furrow. This medium-sized species attains an arm length of 96 mm.
Food and feeding: This species is a generalist or opportunistic predator and consumes a
wide range of prey, including molluscs and crustaceans. The purple mussel Perumytilus
purpuratus was the most abundant prey item (57.6%). Other important prey were the gas-
tropod Pareuthria plumbea, the isopod Exosphaeroma lanceolatum, and the mussels
Aulacomya atra atra and Mytilus edulis platensis (Gil and Zaixso, 2008). In this species,
feeding intensity is at a maximum before and after the reproductive period. Females might
23
occasionally resume feeding when they are still brooding a small number of juveniles.
Prey size increases with starfish size.
Reproduction: In this species, egg laying occurs between March and July, and juvenile
dispersal is mostly in October–November. Juvenile starfish are of ca. 2 mm arm length and
grow to 9–11 mm in one year.
Compounds and Activities:

Antifungal activity: Two new sulfated steroidal hexaglycosides, anasterosides A and B,
along with the known versicoside A isolated from this species displayed antifungal activity
against Cladosporium cucumerinum (Chludil et al., 2000, 2002a; Maier, 2008).
Chludil and Maier (2005) reported on the antifungal activity of sulfated polyhydroxylated
steroidal xylosides, minutosides A and B; and pycnopodioside B isolated from the etha-
nolic extract of this species. Pycnopodioside B and minutoside A were moderately active
(inhibition zones of 7–10 mm) against Cladosporium cucumerinum at the tested con-
centrations (10–60 µg/spot), while minutoside B was inactive at the lowest concentration
(10 µg/spot) and weakly active (inhibition zones of 3–4 mm) at the highest concentrations
(20–60 µg/spot). All these three glycosides were moderately active against A. flavus, show-
ing inhibition zones of 5–10 mm at the highest tested concentrations (20–60 µg/spot). While
minutoside B (2) was inactive at concentrations of 5–10 µg/spot, minutoside A and pycnopo-
dioside B were moderately active (inhibition zones of 5–7.5 mm) at these concentrations.
Biology and Ecology of Pharmaceutical Sea Stars (Class: Asteroidea) 25
Other compounds: Eight glucosylceramides have also been isolated from the water-
insoluble lipid fraction of a methylene chloride/methanol/water extract of this species.
The activities of these compounds are, however, yet to be known (Chludil et al., 2003).
Aphelasterias japonica (Bell, 1881)
Common name(s): Not designated

Global distribution: Temperate Northern Pacific; Tatar Strait to Yellow Sea, Korea
Ecology: It is an epibenthic, epizoic species occupying lower intertidal, shallow subtidal,
deep subtidal, and bathyal zones; oyster reef, mussel reef, kelp forest, macroalgal beds,
coralline algae; and sandy, rocky, and silty grounds up to a depth of 40 m.
Biology
Description: Not reported
Food and feeding: It is a predator and carnivorous feeding voraciously on worms, crusta-
ceans, snails, bivalves, small-sized starfishes, echinoderms, and fishes.
Reproduction: It is a broadcast spawner associated with external fertilization. The merg-
ing larvae are planktotrophic and planktonic. Asexual reproduction, which is often by
fission, is also seen in this species.
Associated species: It is an epibiont species living on the surface of the scale worm
Arctonoe vittata and copepod Scottomyzon gibberum.

Cytotoxicity: The asterosaponin, aphelasteroside F isolated from this species slightly
inhibited cell proliferation and colony formation of the cancer melanoma cell lines
SK-Mel-28, SK-Mel-5, and RPMI-7951 at nontoxic concentrations (Popov et al., 2016).
Haemolytic activity: The compounds isolated from this species include disulfated quinovo-
side aphelasteroside C and the monosulfated polyhydroxysteroid aphelaketotriol; chelifero-
side L1, 3-O-sulfoasterone, forbeside E3, and 3-O-sulfothornasterol A. All these compounds
except 3-O-sulfoasterone showed haemolytic activity to mouse erythrocytes (Ivanchina et al.,
2000).
Others: Four sulfated steroid compounds, viz. aphelasteroside D, pycnopodioside C,
3-O-sulfo-24, 25-dihydromarthasterone, and 3-O-sulfothornasterol A have been isolated
from this species. The activities of these compounds are, however, yet to be known (Kicha
et al., 2001).
Asterias amurensis (Lütken, 1871)
Common name(s): Northern Pacific sea star, Japanese common starfish
Global distribution: Northern China, Korea, Russia, and Japan; Tasmania, southern
Australia, Alaska, the Aleutian Islands, parts of Europe, and Maine
Ecology: It is typically found in shallow waters of protected coasts and is not found on

reefs or in areas with high wave action.
Biology
Description: It is yellow with red and purple pigmentation on its five arms, and a small
central disc. Its distinctive characteristic is its upturned tips. The undersides are completely
yellow and arms are unevenly covered with small, jagged-edged spines. These spines line
the groove in which the tube feet lie and join up at the mouth in a fan-like shape. This spe-
cies can grow up to 50 cm in diameter.
Food and feeding: It eats bivalves, gastropod molluscs, barnacles, crabs, crustaceans,
worms, echinoderms, ascidians, sea urchins, sea squirts, and other sea stars.
Reproduction: It reproduces sexually and asexually. Spawning occurs between July and
October in Australian waters. The female sea star is capable of carrying up to 20 million
eggs. Fertilization is external and larvae remain in a planktonic stage for up to 120 days
before settling and metamorphosing into juvenile starfish.

Antifungal activity: The crude saponin extracted from this species exhibited predominant
growth inhibitory activity against six human fungal pathogens. It showed the lowest mini-
mum inhibitory concentrations (MIC) effect among starfish species against Aspergillus fla
vus and Trichophyton mentagrophytes with the values of 31.2 and 41.6 µg/mL, respectively
(Farhana, 2016).
Antibacterial activity: The crude saponin from this species was found to be slightly active
against bacteria such as Escherichia coli and Edwardsiella tarda with the MIC values of
250 and 125 µg/mL, respectively (Farhana, 2016).
Antimicrobial and antioxidant activity: The organic extracts of the six different tissues
(muscle, gut, liver, tube feet, gonads, and body) of this species exhibited the antimicro-
bial and antioxidant activity. Extracts from all the tissues, except the body tissue, showed
potent antimicrobial activity against Escherichia coli D31. Further, increased antioxidant
activity was observed in the gut, liver, and body extracts (Go et al., 2014).
Haemolytic activity: The crude saponin extracted from this species exhibited haemolytic
activity against 2% mouse erythrocytes (Farhana, 2016).
Cholesterol-binding ability: The crude saponin extracted from this species exhibited
cholesterol-binding ability with the value of 34.3% (Farhana, 2016).
Neuritogenic and antitumour activity: Gangliosides, a complex family of sialylated gly-
cosphingolipids, are abundant in the vertebrate nervous system and play an important role
in the development of the central nervous system. There have been many reports indicating
that gangliosides can induce neuronal differentiation. In vitro evaluation indicated that the
synthesized neuritogenic ganglioside GAA-7 and its glycan moiety of this species showed
strong neuritogenic activity towards neuron-like rat adrenal pheochromocytoma (PC12)
cells in the presence of neurite growth factor (Tamai et al., 2015).
Antitumour activity: The cerebrosides of this species exhibited an inhibitory effect on
cell proliferation through induction of apoptosis in S180 cells. Moreover, administration
of these compounds (50 mg/kg BW) on S180 tumour-bearing mice reduced the tumour
weight by 35.71%. In the S180 ascites tumour model, the treatment (50 mg/kg BW) exhib-
ited a significant ascites fluid growth inhibition of 22.72% (Du et al., 2012a).
HIV-inhibitory activity: Two sulfated sterols isolated from this species showed
inhibitory activity against HIV-1 and HIV-2. While the first sterol showed IC50
values > 126 µM for both HIV − 1 and HIV − 2, the second sterol showed > 123 μM
against HIV −1 and 69 µM against HIV − 2 ( McKee et al., 1994).
Immune enhancement effect: The fatty acids isolated from three organs of this spe-
cies showed immune-enhancing effects on murine macrophages (RAW 264.7 cells). These
fatty acids boosted production of immune-associated factors such as nitric oxide (NO)
and prostaglandin E2 in RAW 264.7 cells. They also enhanced the expression of critical
immune-associated genes, including iNOS, TNF-α, IL-1β, and IL-6, as well as COX-2.
These results increase our understanding of how A. amurensis fatty acids boost immunity
in a physiological system, as a potential functional material (Monmai et al., 2018a).
Anti-inflammatory effect: The fatty acids extracted from the tissues of this species
showed anti-inflammatory effects on RAW264.7 macrophage cells. In lipopolysaccha-
ride-stimulated RAW264.7 cells, fatty acids from the skin, gonads, and digestive glands
exhibited anti-inflammatory activities by reducing nitric oxide production and inducing
nitric oxide synthase gene expression. Further, these fatty acids effectively suppressed the
expression of inflammatory cytokines such as tumour necrosis factor-α, interleukin-1β,
and interleukin-6 in lipopolysaccharide-stimulated cells. Cyclooxygenase-2 and prosta-
glandin E2, which are critical inflammation biomarkers, were also significantly suppressed.
Furthermore, these fatty acids reduced the phosphorylation of nuclear factor-κB p-65, p38,
extracellular signal-related kinase 1/2, and c-Jun N-terminal kinase, indicating that these
fatty acids ameliorated inflammation through the nuclear factor-κB and mitogen-activated
protein kinase pathways. These results provide insight into the anti-inflammatory mecha-
nism of the fatty acids of this species on immune cells and suggest that the species is a
potential source of anti-inflammatory molecules (Monmai et al., 2018b).
Anticancer and apoptotic activity: The sphingoid bases of the glycosylceramides (com-
plex lipids) isolated from this species exerted apoptotic activity on human colon carcinoma
Caco-2 cells (Shaw et al., 2008).
Osteoblastic proliferation: Among the sulfated steroidal compounds, 3β-O-sulfated-
cholest-5-ene-7α-ol and (E) 25-O-β-d-xylopyranosyl-26, 27-dinor-24(S)-methyl-22-
ene-15α-O-sulfated-5α-cholest-3β, 6α-ol isolated from this species, the latter has been
reported to significantly promote the osteoblastic proliferation at 0.01–100 μM (Liu et al.,
2008).
Asterias forbesi (Desor, 1848)
Common name(s): Forbes sea star, common sea star
Global distribution: Northwest Atlantic Ocean and the Caribbean Sea
Ecology: It inhabits shallow waters of the intertidal zone of rocky shores.
Biology
Description: This species usually has five arms but occasionally has four or six. The upper
surface is covered in blunt conical projections, giving it a rough feel. Some of these are
pedicellariae, minute pincers that can grip objects. The arms are plump, broad at the base,
and tapering to a blunt tip. This starfish grows to 15 cm in diameter with an arm length of
about 6 cm. The madreporite is usually pink and is visible near the edge of the disc. There
are several rows of tube feet on the underside on either side of the ambulacral groove that
runs down the centre of each arm. Colour of the upper side is variable, ranging from brown
or tan to reddish purple and the underside is usually pale brown. Near the tip on the under-
side of each arm there are small eyespots.
Food and feeding: It feeds on bivalve molluscs and other marine invertebrates. It can open
shells by gripping the two halves with its tube feet and forcing them apart. It then inserts
its stomach, turning it inside out in the process, and secretes enzymes, digesting the body
of the mollusc in situ.
Disease: A densovirus, named sea star-associated densovirus (SSaDV), has been associ-
ated with the sea star wasting disease (SSWD), which has caused widespread mass mortal-
ity of this species on the Atlantic Coast (Bucci et al., 2017).
Parasite: The ciliate parasite Orchitophrya stellarum has been found in the gonads of up
to 20% of male Asterias forbesi in Long Island Sound. The parasite feeds on the tissue of
the gonad and effectively castrates its host. A small number of females were also found to
contain the parasite.
Associated species: Caprella grahami, an amphipod—an obligate commensal—was
found abundant on individuals of this species. These commensals were found on all parts
of the starfish and seemed to feed mainly on detritus that settled on the starfish or was
caught up in host secretions (Patton, 1968).

Haemolytic and bactericidal activities: The cell-free coelomic fluid of this species
showed haemolytic activity against unsensitized rabbit erythrocytes. Further, the lytic-
free coelomic fluid demonstrated bactericidal activity against Vibrio tubiashii (Leonard
et al., 1990).
Others: Saponins, viz. forbesides A–E, E1-E3, and F–H have been isolated from this
species. Their activities are, however, yet to be known (Findlay et al., 1987a, 1989,
1990a; Findlay and He, 1991; Findlay et al., 1991; Buckingham, 1994; Ahmad and
Baha 2006).
Atta-ur-Rahman (1995) and Findlay and He (1991) reported on the isolation of forbesides
I, J, K, and L from this species.
A sulphated glycolipid, forbesin, and a disodium salt of eicosane-1,16-disulfate have been

isolated from this species (Findlay et al., 1990).
Asterias rathbuni (Verrill, 1909)

Global distribution: Arctic: Northern Bering Sea and Chukchi Sea
Ecology: It is a benthic species.
Biology
Food and feeding: Not reported
The steroidal 24-O-xylosides, designated as rathbuniosides R1 and R2, and the already
known amurensoside A and 3-O-sulfomarthasterone have been isolated from this species.
The activities of these compounds are, however, yet to be known (Ivanchina et al., 2001).
Rathbunioside R1 Rathbunioside R2
Amurensoside A
Asterias rollestoni (Bell, 1881)
Global distribution: Yantai (China)
Ecology: Coastal waters
Biology

Antioxidant activities and neuroprotective effects: The crude polysaccharide SF-2 of this
species displayed the highest antioxidant activity among the polysaccharides. Moreover,
SF-1 and SF-2 exhibited neuroprotective activities in a neurotoxicity model of Parkinson’s
disease (PD) (Zhang et al., 2013; Walag, 2017).
Phosphorylase activity: The compound, purine-2′-deoxyriboside, xanthosine isolated from
this species is involved in purine-nucleoside phosphorylase activity (Huang et al., 2014).
Antitumour effects: The liposomes containing cerebrosides and phosphatidylcholine sep-

arated from this species have significant antitumour effect in vitro and in vivo. In vitro, the
liposomes showed prominent inhibitory effect on the growth of S 180 cells with the 50%
inhibitory concentration (IC50) of 199.89, 148.95, and 126.65 μg/mL, respectively, after
24, 48, and 72 hours in vivo; the liposomes obviously inhibited the growth of tumours in
S 180-bearing mice and the tumour inhibition rate was 72.6% (Du et al., 2012b).
Antidiabetic activity: The compound, enolic glycoside, asterolloside isolated from this
species showed moderate α-glucosidase inhibitory activity with an inhibitory rate of 37.9%
at a concentration of 0.12 mg/mL (Yang et al., 2015).
Cytotoxic activity: Of the two alkaloids, viz. fellutanine A and N-(2-(1H-indol-3-yl)

ethyl)-2-phenylethanamine isolated from this species, the former exhibited potent cyto-
toxic activity against MGC803 with the inhibition rate of 53.99% (Song et al., 2016).
As raw materials for marine drugs and functional foods: This species is a rich source of
valuable bioactive components, such as lipids, proteins, asterosaponin, and polysaccha-
rides. Therefore, it has high nutritional values and development potentials as raw materials
for marine drugs and functional foods. The predominant components of the gonad lip-
ids of this species were triglycerides and phospholipids, while the contents of cholesterol
and free fatty acids were relatively low. Meanwhile, C14:0, C16:0, C18:1n-7, C20:1n-11,
C20:5n-3 (eicosapentaenoic acid, EPA), and C22:6n-3 (docosahexaenoic acid, DHA) were
the main fatty acids, and the content of C20:5n-3 and C22:6n-3 accounted for 21.56% of
total fatty acids. Moreover, the contents of n-3 and n-6 polyunsaturated fatty acid (PUFA)
were respectively 23.62% and 2.71%, and the ratio of n-3 to n-6 polyunsaturated fatty
acids was up to 8.72, which was significantly higher than the daily dietary ratio of n-3 to
n-6 polyunsaturated fatty acids recommended by the Food and Agriculture Organization
(FAO) of the United Nations and the World Health Organization (WHO). Polyunsaturated
fatty acids, especially C20:5n-3 and C22:6n-3, had positive effects on promoting the devel-
opment of the nervous system and curative effects on preventing cardiovascular disease
and anti-inflammatory. Additionally, the atherogenic index (AI) and thrombogenic index
(TI) of the gonad lipids of this species were 0.81 and 0.29, respectively. In conclusion,
the gonad lipids of this species have high nutritional values and great exploitation pros-
pects, and they can effectively prevent atherosclerosis and thrombosis; they can also be the
important dietary sources of C20:5n-3 and C22:6n-3 (Lou et al., 2018).
Asterias rubens (Linnaeus, 1758)
Common name(s): Common starfish, common sea star
Global distribution: It is native to the northeastern Atlantic Ocean and its range extends
from Norway and Sweden, through the North Sea, round the coasts of Britain, France, Spain,
and Portugal and southwards along the coasts of Africa to Senegal; it is also known from the
western Atlantic where it occurs between Labrador and Florida and the Gulf of Mexico.
Ecology: It inhabits rocky and gravelly substrates, and it is capable of surviving in brackish
water too.
Biology
Description: The common starfish normally has five arms, broad at their base, and gradu-
ally tapering to a point at their tips, which are often turned up slightly. There is a line of
short white spines running along the centre of the aboral (upper) surface of the arms. The
oral (lower) surfaces of the arms have rows of small tube feet, used in locomotion and
feeding. This starfish is usually orange or brick red on the aboral surface and paler on the
oral surface but can also be purple or pale brown. Individuals from deep water are usually
paler. It grows to a maximum diameter of 52 cm.
Food and feeding: It largely feeds on molluscs and other benthic invertebrates.

Antibacterial activity: The extracts of the gastrointestinal organs and eggs of this species
have shown high antibacterial activity. Similarly, the antimicrobial peptides of its coelo-
mocytes fluid have also shown similar activity (Haug et al., 2002).
Li et al. (2010a) reported that the peptides/proteins of the body wall—fragments of actin,
histone H2A, filamin A, peptides, and lysozyme of its celomocytes—have shown antimi-
crobial activities.
Lysozyme-like activity: Several tissues of this species displayed lysozyme-like activity
(Haug et al., 2002).
Haemolytic activity: The body wall extracts of this species showed haemolytic activity
(Haug et al., 2002). Maier (2008b) reported that its steroid, viz. thornasterol A sulphate
has shown haemolytic activity to mouse erythrocytes with an ED50 value of 1.1 × 10 –4 M.
Others: Four compounds, viz. ruberosides A−D have been isolated from this species.
Their activities are, however, yet to be known (Sandvoss et al., 2000).
Ruberosides B, A Ruberosides C
Coscinasterias tenuispina (Lamarck, 1816)
Common name(s): Blue spiny starfish, white starfish
Global distribution: The range of this subtropical species includes the Mediterranean
Sea, France, Spain and Portugal, the Azores and other Atlantic Islands; Bermuda, Cuba,
and the American coast between North Carolina and Santos, Brazil.
Ecology: This benthic species inhabits shallow waters on the lower shore and down to a
depth of 165 m as well as hard bottoms and under stones and seaweed.
Biology
Description: It has six to twelve arms (usually seven), often of varying lengths, and it
grows to 20 cm in diameter. It is a creamy, slightly bluish colour, variously blotched with
brown, and it is rough textured with short spines.
Food and feeding: It is a predator and an omnivore. It mainly feeds on other echinoderms
and on bivalve molluscs.
Reproduction: In most of its range, this species undergoes sexual reproduction in the win-
ter, while in the summer it proliferates by asexual reproduction. In the process of asexual
reproduction or “fission,” the disc tears itself into two sections, with each part eventually
growing extra arms and developing into a new individual. In Brazil, all individuals of this
species seem to be male and fission occurs throughout the year.

Antitumour activity: The acetylenic lipid compound asterinic acid, isolated from this
species, showed antitumour activity (Kilimnik et al., 2016).
Other compounds: Steroidal glycosides, viz. tenuispinosides A–C and coscinasterosides

A–F have been isolated form this species. The activities of these compounds are, however,
yet to be known (Riccio et al., 1986).
Coscinateroside A Coscinateroside B
Diplasterias brucei (Koehler, 1907)
Global distribution: Antarctica: Antarctic Peninsula and in East Antarctica; Pacific

Ocean and Southern Ocean
Ecology: This benthic, polar species is found on the seabed at a depth range of 105–407 m.
Biology
Description: It has five arms, but the six-armed form is common. This starfish varies in
colour from a pale bluish green to a yellowish or orange hue. The maximum size is about
25 cm across.
Food and feeding: It is a predator and scavenger primarily feeding on bivalve mollusc.
Limatula hodgsoni.
Predator: It is sometimes preyed on by the sea anemone Urticinopsis antarctica.
Reproduction: Developing embryos of this species are brooded by the female until they
have developed into juvenile starfish.

Cytotoxic activities: Two new asterosaponins, diplasteriosides A and B, and the previ-
ously known asteriidoside A of this species have shown cytotoxic activities against human
colon cancer cell line HCT-116, human breast cancer cell line T-47D, and human mela-
noma cancer cell line RPMI-7951 (Ivanchina et al., 2011a; Atta-ur-Rahman, 2016).
Distolasterias nipon (Döderlein, 1902) (= Asterias nipon)

Global distribution: Northwest Pacific: East and South China Seas, Hong Kong, Taiwan,
and Japan
Ecology: This subtropical, benthic species lives on sandy substrate at depths of 100–200 m.
Biology
Description: It has five rays that are usually of different length. Rays are subpetagonal in
cross section. Carinal plates are imbricated in regular line. Marginals are also imbricated.
There are 1–2 straight pedicellariae between the subambulacral spine and inferomarginal
spines. Sparse straight pedicellariae are present on furrow margin.
Neuritogenic and neuroprotective effects: The steroidal glycoside compounds distolas-
terosides D1–D3 isolated from this species showed neuritogenic effects at concentrations
of 1–50 nM. These steroids also act as neuroprotectors against oxygen-glucose deprivation
(OGD) by increasing the number of surviving cells (Palyanova et al., 2013).
Other compounds: Lorizzi et al. (1993) reported on the occurrence of steroidal glycoside
compounds such as pycnopodioside C and pisasteroside A; asterosaponins, viz. versico-
side A, thornasteroside A, and nipoglycosides A–D; and glycosides of polyhydroxysteroids,
distolasterosides D1, D2, D4, and D5. The activities of these compounds are, however,
not known.
Kicha et al. (2008a) reported on the occurrence of distolasterosides D6, D7, D1, D2, and
D3, and echinasteroside C. Among these compounds, distolasterosides D1, D2, and D3
were found to induce neuroblast differentiation in a mouse neuroblastoma C 1300 cell
culture.
pycnopodioside C pisasteroside A
Evasterias echinosoma (Fisher, 1926)

Common name(s): Spiny net starfish
Global distribution: East Coast of Korea (Uljin, Sacheon, Jinjae, Jangho); Hokkaido,
Abakaman, Kamchatka—Okhotsk Sea, and Nombo Bering Sea.
Ecology: It is known to inhabit reef environments and sandy areas ranging in depth from
20 to 50 m (or more).
Biology
Description: The body of this species is very large, about 20 cm long from the small central
disc to the end of the ray. There are five, long, and very strong rays. The length of each
ray becomes narrower toward the end. The epidermis is very hard and rough, with many
protrusions. Body colour is dark red, and the dorsal pole is almost white. The underside is
dark yellow.
Food and feeding: It feeds on infaunal and small epifaunal residents of the benthos.
The steroid glycosides, viz. evasteriosides D and E; pycnopodiosides A and C; lurido-
side A; 5α-cholestane-3β,6α,8,15β,16β,26-hexaol; 5α-cholestane-3β,6α,8,15β,24-pentaol
24-sulfate sodium salt and marthasterone sulfate sodium salt have been identified from
this species. The bioactivities of these compounds are to be known (Levina et al., 2009a).
Evasterias retifera (Djakonov, 1938)

Common name(s): Western Star
Global distribution: Pacific Ocean: Sea of Japan
Ecology: It is a fairly deepwater species present at depths of 33–68 m.
Biology
Description: It is a large red star, with bluish needles. Rays are thick, sharpening towards
the ends, 25.5 cm long and more. The needles of the dorsal side form groups connected
to each other by jumpers. In this species, there are several morphs (E. retifera retifera,
E. retifera retata, E. retifera tabulata), differing in features of colouring and location of
needles on the back. Along the dorsal side are many marginal needles forming double rows
on each ray. This species differs in high fecundity.
Food and feeding: It feeds on free-living and sedentary benthic animals or their remains.
Associated species: Young red king crab (Paralithodes camtschaticus)

The sulphated xylosides, viz. evasteriosides A, B, and C have been isolated from this species.
Their activities are yet to be known (Ivanchina et al., 2011).
Evasterioside A Evasterioside B
Leptasterias fisheri (Djakonov, 1929)
Global distribution: Pacific Ocean: Sea of Japan
Ecology: It inhabits silt and stones at a depth of 84 m.
Biology
Food and feeding: Not known

The glycoside, fisherioside A, has been isolated from this species (Kicha et al., 2012).
Leptasterias hylodes (Fisher, 1930)
Common name(s): Aleutian star
Global distribution: Northeast Pacific: Alaska
Ecology: It is a benthic species found on rocky and sandy regions at depths of 10–180 m.
Biology
Description: Rays of this species are slender and gently tapering. Dorsal spines are uni-
form, slender, and tapering, about 1 mm long. Straight pedicellariae are few on the dorsal
surface, and crossed pedicellariae are larger. Ventrolateral plates are in a series.

Cytotoxic activity: The pentasaccharide, hylodoside A, isolated from this species
displayed cytotoxic activity against mouse erythrocytes carcinosoma cells (Atta-ur-
Rahman, 2016).
Antibacterial and haemolytic activities: The steroid glycosides, hylodoside A and

novaeguinoside Y, along with previously known five polyhydroxylated steroids have been
isolated from the ethanolic extracts of this species. Among these compounds, the polyhy-
droxylated steroid was shown to inhibit the growth of Staphylococcus aureus up to 10%
from the control at a concentration of 1 mg/mL. The compounds hylodoside A and novae-
guinoside Y and another polyhydroxylated steroid showed moderate haemolytic activity in
the mouse erythrocytes assay. Further, the polyhydroxylated steroids also displayed pH-
depended haemolytic properties (Levina et al., 2010).
Leptasterias ochotensis (Brandt, 1851)
Global distribution: Okhotsk Sea and northern Japan Sea
Ecology: It inhabits both shallow and deep waters with a depth range of 3–2601 m.
Biology
Description: Dorsal spines are small, spaced, cylindrical, or clavate. Straight pedicellar-
iae are small and subtriangular, and they are distributed on the dorsal and ventral sides.
Madreporite is circular, situated about midway between the centre of the disc and the
margin. The size of this species ranges from 15 to 28 mm.
Food and feeding: It feeds on a variety of diets such as barnacles, mussels, chitons, and
limpets.
Reproduction: These animals deposit their eggs on the substratum, and the adult covers
them until the metamorphosed juveniles grow to a considerable size.
Cytotoxicity: Three new sulphated steroid monoglycosides—leptaochotensosides A–C
and a new sulphated polyhydroxylated steroid—have been isolated from the alcoholic
extract of this species. Although the isolated compounds did not show any apparent cyto-
toxicity against melanoma RPMI-7951 and breast cancer T-47D cell lines, leptaochoten-
soside A demonstrated inhibition of T-47D cell colony formation in a soft agar clonogenic
assay at nontoxic doses. In addition, this compound decreased the epidermal growth factor
(EGF)-induced colony formation of mouse epidermal JB6 Cl41 cells. The cancer preven-
tive action of leptaochotensoside A is realized through regulation of a mitogen-activated
protein kinase (MAPK) signalling pathway (Malyarenko et al., 2015).
Anticancer and cytotoxic activities: Six new asterosaponins, leptasteriosides A–F, and
two asterogenins have been isolated from the alcoholic extract of this species. While
leptasteriosides A–F showed slight or moderate cytotoxic activities against cancer cell
lines RPMI-7951 and T-47D, the asterosaponins demonstrated a significant inhibition of
RPMI-7951 and T-47D cell colony formation in soft agar clonogenic assay in nontoxic
doses (Malyarenko et al., 2014). (Top to bottom: Leptasterioside B, A, C, D, E, F.)
Lethasterias fusca (Djakonov, 1931)
Common name(s): Far Eastern starfish, warm-water sea stars
Global distribution: Korean coastal seas
Ecology: It lives on rocky reefs, rocky soils at shallow depths (2–50 m); it is less commonly
found on silted sands with an admixture of pebbles and stones; it is also found on the thalli
of algae-macrophytes, often in oysters or mussel banks.
Biology
Description: This seaside five-ray star can be easily distinguished by its black or almost
black colour of the central disc and the rays from the dorsal side. There are also dark grey
flies, and on the rays on a dark background there may be yellowish and whitish spots,
sometimes located in the form of bandages.
Food and feeding: They lead a predatory life, attacking small-sized molluscs.

Antitumour activity: Its asterosaponins and other steroid glycosides exhibited a signifi-
cant suppression of the human tumour HT-29, HCT-116, RPMI-7951, and T-47D cell col-
ony formation in a soft agar clonogenic assay (Malyarenko et al., 2015).
Blunt et al. (2014) reported that its lethasterioside A, while being weakly cytotoxic, had
pronounced ability to inhibit colony formation of tumour cells.
Cytotoxicity: Two new asterosaponins, lethasteriosides A (1) and B (2), were isolated along
with previously known thornasteroside A (3), anasteroside A (4), and luidiaquinoside (5) from
the ethanolic extract of this species. Compounds 1 and 3–5 did not show any apparent cyto-
toxicity against cancer cell lines T-47D, RPMI-7951, and HCT-116, but glycoside 1, at con-
centration of 20 μM, demonstrated considerable inhibition of the T-47D (97%), RPMI-795I
(90%), and HCT-116 (90%) cell colony formations in a soft agar clonogenic assay (Ivanchina
et al., 2012).
Lethasterias nanimensis chelifera (Verrill, 1914)

Common name(s): Russian starfish, black-spined star
Global distribution: From Saghalien to Bering Strait thence to Gulf of Alaska and south-
ward to Hokkaido
Ecology: It occurs on fine grey sand and green mud areas at depths of 76–284 m.
Biology
Description: In this species, dorsal spines are tipped black. The spines are thickly wreathed
with crossed pedicellariae. Marginal plates are 4-lobed, arranged in regular longiseries.
Superomarginal plates are situated low on the side of the ray. The madreporite is circular.
The colour in a dry specimen is blackish grey in the dorsal side, and the ventral side is a
little paler. Radius of rays is about 170 mm.
Food and feeding: It mainly feeds on small gastropods.

A total of 13 steroidal compounds have been isolated from this species. Activities of these
compounds are, however, to be known (Kicha et al., 2004).
Kicha et al. (2003) also isolated 4 alkaloidosteroids, viz. 1-methyl-6,7-dihydroxy-

1,2,3,4-tetrahydroisoquinolinium salts of 3-O-sulfoasterone (1), 3-O-sulfoisoasterone
(2),3-O-sulfothornasterol A (3) and an alkaloid, 1-methyl-1,2,3,4-tetrahydro-β-carboline-
3-carboxylic acid (4) from this species.
Anticancer activity: The endogenous neurotoxin 1-methyl-6,7-dihydroxy-1,2,3, 4-tetrahy-

droisoquinoline (salsolinol) of this species has been reported to show anticancer activity in
human dopaminergic neuroblastoma SH-SY5Y cells by inhibiting the mitochondrial com-
plex I (NADH-Q reductase) activity. The cell death induced by salsolinol in the present
investigation was largely due to impairment of cellular energy supply, caused in particular
by inhibition of mitochondrial complex II (succinate-Q reductase) (Storch et al., 2000).
Lysastrosoma anthosticta (Fisher, 1922)
Common name(s): Pacific starfish
Global distribution: Sea of Japan
Ecology: It occurs in areas representing major intertidal habitats.
Biology
Description: Disc of this species is small, and rays are marked off from the disc by a slight
constriction at the base. The whole body is very weak and flabby. The ambulacral and
adambulacral plates are rather loosely articulated, and the plates are not hard and firm but
rather spongy. Radius of the rays is 63 mm. The madreporic body, sometimes invisible, is
situated near the edge of the dis and surrounded by several spinelets. One of the character-
istic features of this species is the fact that the rays are slightly spaced on the circumference
of the disc.
Food and feeding: It is a predator feeding on live food.

Anti-HIV and immunomodulatory activity: Lysaketotriol (1) and lysaketodiol (2),
along with two previously known polyhydroxylated steroidal sulfates (3,4), were iso-
lated from the ethanolic extract of ambulakrums separated from the arms of this species.
Compound 1 produced moderate stimulation of lysosomal activity in mouse splenocytes.
The lysosome acts as a safeguard in preventing pathogens from being able to reach the
cytoplasm. Lysosomal activity also results in a decrease in viral infectivity, including
HIV. Compounds 1-2, on the other hand, showed immunomodulatory activity (Levina
et al., 2009b).
Others: A steroidal glycoside, lysastroside A, has been isolated from this species (Jha and
Zi-rong, 2004).
Levina et al. (2001) reported on the isolation of four steroid compounds, viz 3β, 6α-dihydroxy-
5α-cholesta-9(11),24-dien-23-one 3-sulfate(1) 3β,6α-dihydroxy-5α-cholest-9(11)-en-23-
one 3-sulfate (2), sodium 24-O-β-d-glucopyranoside 6″-sulfate (pycnopodioside C)(3) and
sodium 24-O-β-d-xylopyranoside 4″-sulfate (luridoside A)(4) from this species.
Marthasterias glacialis (Linnaeus, 1758)
Common name(s): Spiny starfish

Global distribution: Widely distributed throughout northwestern Europe, the Atlantic,
and the Mediterranean Sea
Ecology: It is essentially a sublittoral species extending to depths of about 180 m, but some
specimens are found at low water on rocky shores and sheltered muddy sites.
Biology
Description: This starfish is large and has five very spiny arms. Each arm bears three lon-
gitudinal rows of spike-like spines surrounded by large cushions of pedicellariae. Smaller
spines may be scattered between these rows. The spines are white and usually purple-
tipped, and the animal varies in colour from dirty brown to greenish grey with purple tips
to the arms. It grows up to 35 cm or more in diameter.
Food and feeding: It is a mobile carnivorous species, feeding on a wide range of organ-
isms (dead or alive) such as molluscs, shellfish, fish, or other echinoderms.

Antibacterial and lysozyme-like activity: The eggs from this species exert antibacterial
action on marine bacterial strains and show a lysozyme-like activity (Stabili and Pagliara,
1995).
Antitumour activity: The acetylenic lipid compound, asterinic acid, isolated from this
species showed antitumour activity (Kilimnik et al., 2016).
Anti-inflammatory activity: Its predominant compounds cis 11-eicosenoic and cis 11,
14 eicosadienoic acids and the unsaturated sterol, ergosta-7, 22-dien-3-ol displayed anti-
inflammatory activity (Pereira et al., 2014a, 2014b).
Others: Pereira et al. (2014b) reported on the presence of compounds such as cis 11-eicose-
noic acid, cis 11,14 eicosadienoic acid, palmitic acid, ergosta-7,22-dien-3-ol, cholesterol,
astaxanthin, lutein, and zeaxanthin in this species.
Dini et al. (1983) reported on the occurrence of saponins, marthasteroside A1, A2, B, and
C from this species.
Pisaster giganteus (Stimpson, 1857)

Common name(s): Giant sea star, giant spined star
Global distribution: Western coast of North America from Southern California to British
Columbia
Ecology: It is usually found around the protected coastlines with low tide; it is often found
attached to rocks, pier supports, or in the sand.
Biology
Description: This species has a dense body with wide arms. Its surface is either tan or
brown, but on occasion it can have a yellowish or greyish surface. There are thick, blunt
spines that are bluish in colour with white, pink, or purple tips that are swollen and sur-
rounded by brown fuzz and pedicellariae. These pedicellariae are used as a protective
mechanism against predators. It can grow as large as 61 cm in diameter.
Food and feeding: The species preys on several kinds of sea organisms including bar-
nacles, gastropods, bivalves and limpets. It eats its prey by extending its stomach so it can
fit into tiny gaps, such as mussel shells.
Predators: The giant sea star only has a few predators. Sea otters and sea birds feed on
giant sea stars, and their larvae are eaten by certain types of sea snails.
Reproduction: Giant sea stars have small eggs, and their sperm contain spherical heads.
Once their larvae are born, they are bilaterally symmetrical. By the time they mature and
reach adulthood, they are centred on a set point with radial symmetry to their bodies. The
gonads of the giant sea star grow in wintertime, just in time for spawning season between
the months of March and April.

Four steroidal monoglycoside sulphates—viz. pycnopodioside B, pisasterosides D, E, and
F—and two known asterosaponins, thornasteroside A and versicoside A, have been iso-
lated from this species (Zollo et al., 1990).
2.2 FAMILY: HELIASTERIDAE (ORDER: FORCIPULATIDA)

Labidiaster annulatus (Sladen, 1889)
Common name(s): Antarctic sun starfish
Global distribution: Cold waters around Antarctica; Antarctic Peninsula, South Georgia
and the South Sandwich Islands
Ecology: This polar, benthic species occurs on the seabed and is found on sand, mud, and
gravel and among rocks from the intertidal zone down to 554 m.
Biology
Description: This species has a wide central disc and 40–45 long narrow rays and can
reach a diameter of 60 cm. The disc is slightly inflated and is raised above the base of the
rays. The madreporite is large and near the edge of the disc. The aboral or upper surface
is covered in a meshed network of small, slightly overlapping plates. These are covered by
a membrane with numerous raised projections called papulae, some small spines, and a
few large triangular pedicellariae. The oral or lower surface of the disc has a central mouth
surrounded by additional scales. The ambulacral grooves are wide and run down the centre
of the oral side of each ray. There are widely separated narrow scales on either side of the
grooves with two spines on each, one overlapping the groove and the other projecting from
the side of the ray. There are rows of tube feet on either side of the groove, each foot having
a button-like suction pad at the tip.
Food and feeding: It is an opportunistic predator and scavenger. Krill and amphipods are
the most frequent diet items. Other dietary items are varied, including smaller starfish of
this species and brittle stars such as Ophionotus victoriae.
Reproduction: Little is known of the reproduction of Labidiaster annulatus, but the
larvae pass through at least one bipinnaria and one brachiolaria stage, as has been dem-
onstrated by DNA analysis. The larvae are planktonic and spend many months drift-
ing with the currents before settling on the seabed and undergoing metamorphosis into
juveniles.

De Vivar et al. (2000) reported on the occurrence of two sulfated pentaglycosides, viz.
Labidiasteroside A and ovarian asterosaponin 1 from this species.
2.3 FAMILY: STICHASTERIDAE (ORDER: FORCIPULATIDA)

Cosmasterias lurida (Philippi, 1858)
Common name(s): Common fjord starfish
Global distribution: Subtropical to polar; Southeast Pacific, Southwest Atlantic, and

Atlantic Antarctic
Ecology: This benthic, cold-water species is found on solid substrates such as rock, cobble,
and boulders in intertidal areas to a depth of 650 m.
Biology
Description: It is a large, purplish sea star, with rows of small, cylindrical bumps along the
arms. It reaches up to 40 cm diameter.
Food and feeding: It is known to feed largely on the kelp, Macrocystis pyrifera.

Steroidal oligoglycosides, viz. cosmasterosides A–D, ophidianoside F, and forbeside H
have been isolated from this species (Roccatagliata et al., 1994).
Maier et al. (1998) reported on the isolation of three glucosylceramides, viz. (2S,3R,4E,8E,
10E)-1-(β-d-glucopyranosyloxy)-3-hydroxy-2-[(R)-2-hydroxyheptadecanoyl)amino]-
9-methyl-4,8,10-octadecatriene(1), (2S,3R,4E,8E,10E)-1-(β-d-glucopyranosyloxy)-3-
hydroxy-2-[(R)-2-hydroxyoctadecanoyl)amino]-9-methyl-4,8,10-octadecatriene(2), and
ophidiacerebroside E(3) from the water-insoluble lipid fraction of the methylene chloride/
methanol extract of this species.
Neosmilaster georgianus (Studer, 1885)
Common name(s): Antarctic starfish
Nothing is known about its distribution, ecology, biology, and food and feeding.
Reproduction: This species has been reported to reproduce throughout the year at low
rates, but the pseudocopulatory behaviour was observed in the austral spring/summer in
September/October. Before the action starts, there is “exploratory contact behaviour” as
the animals touch each other up, with males initiating contact with females. Because the
attraction of males to females is thought to be rooted in pheromones or some other chemo-
tactic stimulus, it is possible that one male pairing might actually attract other males to a
reproductively active female. This whole process takes place over the course of minutes to
hours. Spawning happens a few hours later.

Vázquez et al. (1992) reported on the occurrence of an asterosaponin, named santiagoside
from this species.
2.4 FAMILY: ZOROASTERIDAE (ORDER: FORCIPULATIDA)

Myxoderma platyacanthum (Clark, 1913)
Global distribution: Eastern Central Pacific: California, US

Ecology: This benthic, subtropical species occurs on soft bottoms at depths of 820–1030 m.
Biology
Food and feeding: It feeds on ophiuroid ossicles and bivalves.

Seven polyhydroxy steroids (1–7) and asteroid glycoside, myxodermoside A have been
isolated from this species (Finamore et al., 1991).
2.5 FAMILY: ASTROPECTINIDAE (ORDER: PAXILLOSIDA)

Astropecten indicus (Döderlein, 1888)
Common name(s): Plain sand star
Global distribution: Indo-Pacific: United Arab Emirates, Qatar, and Thailand
Ecology: This benthic, tropical species occurs in intertidal to subtidal mud, sand and shell
habitats. It comes out in large numbers at sunset. During the day, it usually remains buried
in the sand or silt. Depth range is 181–196 m.
Biology
Description: The body of this species is rather flat. Arms are long and tapered to a sharp
tip. Along the sides of the arms are stout, flat, long spines. The spines are usually tinged a
bright orange at the base with white tips. The marginal plates on the sides of the arms are
not so large. The white tube feet are pointed. Colours are generally a plain bluish brown,
with a darker brown centre and stripes down the length of the arms. The tips of the arms
are black. Diameter of arms is 4–6 cm.
Food and feeding: It feeds on a variety of bivalves and gastropods, as well as crustaceans.
Sometimes, tiny white snails are found on the upper side. These are parasitic snails (Family
Eulimidae).

Antibacterial activity: The crude methanol extracts of the tissue sample of this spe-
cies exerted antimicrobial activity on human microbial pathogens, viz. Escherichia
coli, Klebsiella pneumoniae, K. oxytoca, Staphylococcus aureus, Streptococcus sp.,
Pseudomonas aeruginosa, Salmonella paratyphi, and S. typhi. The maximum inhibiting
zone of 13.4 mm was observed in the methanol extracts for bacteria Pseudomonas aeru
ginosa, followed by ethyl acetate 11.3 mm against Klebsiella pneumoniae at the concentra-
tion of 1000 µL. Moderate activity was found for all the tested bacterial strains at 500 µL
concentration of both the solvents (Chamundeeswari et al., 2012).
Others: Atta-ur-Rahman (1995) reported on the occurrence of galactofuranosides, indico-
sides A–C from this species.
Astropecten irregularis (Pennant, 1777)
Common name(s): Sand sea star
Global distribution: Northeast Atlantic and the Mediterranean
Ecology: This subtropical, epibenthic, and free-living species is found in all kinds of
mobile seabeds from 1 to about 1000 m deep. This species is active and easy to find during
the night; sometimes it is possible to find it in the late afternoon.
Biology
Description: The body of this species has a fairly large disc and five rather short, stiff, and
tapering arms that are up to 100 mm in diameter (occasionally 200 mm). The upper surface is
covered with paxillae. The upper and lower rows of marginal plates are well marked. The upper
marginal plates generally have a larger, conical spine. The lower marginals are flattened, rect-
angular, and at their upper end bear 4–5 large marginal spines in an oblique series. In the oral
side, the adambulacral plates each bear three equal-sized furrow spines; and there are no pedi-
cellariae. The tube feet are pointed and suckerless, and they are provided with double ampullae.
These animals are coloured reddish violet or yellowish, with or without purple marks.
Food and feeding: This sea star is a carnivore and feeds on molluscs, which it catches with
its arms and then takes to the mouth. The prey is then trapped by the long, moving prickles
around the mouth cavity.

Antioxidant activity: The organic extracts of this species showed DPPH antioxidant activ-
ity with a value of 38.5 mg TE/g edw (the concentration of standard Trolox with the same
antioxidant capacity of the extract under investigation) (Marmouzi et al., 2018).
Antidiabetic activity: The organic extracts of this species showed antidiabetic activity.
The IC50 inhibition values of α-Amylase and α-Glucosidase were found to be 147.0 and
540.0 µg/mL, respectively (Marmouzi et al., 2018).
Antibacterial activity: The organic extracts of this species showed antibacterial activity
against Staphylococcus aureus CIP 483, Bacillus subtilis CIP 5262, Escherichia coli CIP
53126, Pseudomonas aeruginosa CIP 82118, and Salmonella enterica CIP 8039 with MIC
values of 12.5, 3.1, 6.3, 3.1, and 6.3 mg/mL, respectively (Marmouzi et al., 2018).
Astropecten latespinosus (Meissner, 1892[1])
Common name(s): Sea star
Global distribution: Seas around Japan
Ecology: It lives on a mobile seabed (sandy, muddy, or gravel seabed) and remains largely
buried under sediment during the day.
Biology
Description: It is a five-armed starfish with a brownish flattened body and a fringe of pale-
coloured spines round the margins of the short, tapered arms.
Reproduction: Unlike many members of this genus, the larva of Astropecten latespinosus
is not a brachiolaria larva but is barrel-shaped and undergoes metamorphosis at a very
early stage of development.

Antitumour activity: Four cerebrosides, astrocerebroside A–C and acanthacerebroside
A, have been isolated from this species. Further, two gangliosides, LG-1, and LG-2, which
occur as mixtures of various alkyl chains, were also isolated from this species. Among
these compounds, LG-2 showed antitumour activity against murine lymphoma L1210 cells
in vitro (Higuchi et al., 1995).
Atta-ur-Rahman (1995) reported on the occurrence of versicoside A in this species.
Astropecten monacanthus (Sladen, 1883)
Common name(s): Vietnamese starfish
Global distribution: Indo-Pacific: Iran, Thailand, China, India, Eastern Mrica,

Madagascar, Mozambique, and the Red Sea
Ecology: This tropical, benthic species is found lying on the sand; the depth range is
0–112 m.
Biology
Description: Peripheral margins of the body of this species are fringed with conspicuous
large spines. Supero-marginal plates are relatively narrow. The paxillar area at the base of
the arm is more than half the total arm breadth. The actinal (ventral) surface of the infero-
marginal plates is very smooth and covered with short, rounded (more or less) apressed squa-
mules. The colour of the body is purple, reddish yellow, or light grey. It is an edible species.
Food and feeding: It is carnivorous, feeding on invertebrates and seashells.

Anti-inflammatory activity: Four new asterosaponins, astrosteriosides A–D (1–3 and 5),
and two known compounds, psilasteroside (4) and marthasteroside B (6), have been iso-
lated from the MeOH extract of this species. Compounds 1, 5, and 6 exhibited potent anti-
inflammatory activity. Such potent anti-inflammatory activities render compounds 1, 5,
and 6 important materials for further applications including complementary inflammation
remedies and/or functional foods and nutraceuticals (Thao et al., 2013a).
Anticancer activity: Six asterosaponins (1–6) isolated from the MeOH extract of this species
displayed cytotoxic activities against human cancer cell lines, HL-60 (promyelocytic leuke-
mia), PC-3 (prostate cancer), and SNU-C5 (colorectal cancer) with IC50 values ranging from
0.84 to 3.96 µg/mL. Further, its compound astrosterioside D exhibited potent cytotoxic effects
against all these cell lines with IC50 values ranging from 4.31 to 5.21 µM. In addition, the
MeOH extract and astrosterioside D had an effect on leading to apoptosis (Thao et al., 2014).
Astropecten polyacanthus (Müller and Troschel, 1842)
Common name(s): Sand sifting starfish, comb sea star, brown spotted combstar
Global distribution: Throughout the Indo-Pacific region; from the Red Sea and Zanzibar
to Hawaii, and from Japan to Australia and New Zealand
Ecology: It is often found on silty sand bottoms in harbours and estuaries at depths down
to about 185 m. This tropical, reef-associated species is buried in the sand during daytime
and crawls on the sand to search for food at night.
Biology
Description: The upper surface of this starfish is a dark purplish colour, while the under-
side is orange. On the upper surface paxillae, little pillars with flattened summits are
cream, grey, or brown; the colours sometimes making a chevron pattern. Along the edges
of the five arms there is a fringe of long, sharp marginal spines, usually with brown bases
and pale tips. The arms are fairly broad and have a maximum length of 9 cm. The tube feet
are pointed rather than having suckers, an arrangement that is more suitable for digging.
The arm spread is up to 20 cm.
Food and feeding: It feeds on detritus and bivalve and gastropod molluscs, which it swal-
lows whole. It also sometimes engulfs pebbles and digests the biofilm and small inverte-
brates adhering to the surface.
Aquarium values: The comb star is sometimes kept in reef aquaria where it is efficient at
clearing detritus and uneaten food from the sand or gravel. It is mostly nocturnal and needs
to be acclimatized gradually to the conditions in the tank.

Anti-inflammatory activity: The crude extracts and steroids isolated from this spe-
cies showed inhibitory effects on pro-inflammatory cytokine (Interleukin-12 (IL-12) p40,
interleukin-6 (IL-6), and tumour necrosis factor α (TNF-α)) production in lipopolysaccha-
ride (LPS)-stimulated bone marrow-derived dendritic cells (BMDCs). Among compounds
tested, compounds 5 and 7 showed potent inhibitory effects on the production of all three
pro-inflammatory cytokines with IC50 values ranging from 1.82 to 7.00 μM. Potent inhib-
itory activities were also observed for compound 1 on the production of IL-12 p40, and
IL-6 with values of 3.96 and 4.07 μM, respectively, and for compounds 3 and 4 on the pro-
duction of IL-12 p40 with values of 6.55 and 5.06 μM, respectively. Moreover, compounds
2 (IC50 = 34.86 μM) and 6 (IC50 = 79.05 μM) exhibited moderate inhibitory effects on the
production of IL-12 p40, whereas compounds 3 (IC50 = 22.80 μM) and 4 (IC50 = 16.73 μM)
moderately inhibited the production of TNF-α and IL-6, respectively (Thao et al., 2013b).
Cytotoxic activity: Four new steroids, astropectenols A-D (1–4), along with three known
compounds, viz. 5α-cholest-7-ene-3β,6α-diol (5), 5α-cholest8(14)-ene-3β,7α-diol (6), and
5α-cholest-7,9(11)-diene-3β-ol (7) have been isolated from this species. The CH2Cl2 frac-
tion and compound 7 exhibited potent cytotoxic effects against HL-60 human leukemia
cells with the IC50 of 8.29 µg/mL and 2.70 µM, respectively. On the other hand, the
CH2Cl2 fraction and compound 7 induced the apoptosis of HL-60 cells via the inactivation
of ERK 1/2 and the decrease of C-myc. These findings suggested the potential use of the
CH2Cl2 fraction and compound 7 of this species for leukemia treatment (Thao et al., 2013c;
Sumithaa et al., 2017a).
Astropecten scoparius (Müller and Troschel, 1842)
Global distribution: Western Central Pacific: South China Sea
Ecology: This benthic, tropical species lives in shallow water at depths of 14–64 m; it
burrows in the muddy sediments on the seabed.
Biology
Description: It is a grey starfish, and each of its five arms has a narrow pale margin.
Food and feeding: It largely feeds on molluscs.

Reproduction: It mostly breeds between June and August. Both females and males liber-
ate gametes into the sea where fertilization takes place. The bipinnaria larvae that hatch
from the eggs are planktonic.
Toxicity: In some circumstances, this species contains the neurotoxin tetrodotoxin, also
known as TTX, which is associated with cases of paralytic shellfish poisoning of humans
in Japan caused by consumption of the trumpet shell Charonia lampas. It is believed that
the toxin is passed through the food chain, the trumpet shell having acquired it through
feeding on the starfish. The starfish may themselves have incorporated TTX into their tis-
sues through feeding on certain tiny gastropod molluscs of the species Umborium suturale.

growth inhibitory activity against human fungal pathogens (Farhana, 2016).
Antibacterial activity: The crude saponin extracted from this species exhibited antibacte-
rial activity (Farhana, 2016).
cholesterol-binding ability (Farhana, 2016).
Others: Atta-ur-Rahman (1995) reported on the isolation of scopariosides A–D, polyhy-
droxysteorids (4 compounds), and an asterosaponin, hexaglycoside, viz. marthasteroside
A1 from this species.
Craspidaster hesperus (Müller and Troschel, 1840)

Common name(s): Bordered sea star
Global distribution: Indo-West Pacific: Bay of Bengal, Japan to the Philippines
Ecology: This subtropical demersal species has a depth range of 2–195 m; it spends its
time buried in silty seabeds.
Biology
Description: It is a flat sea star with elegant tapered arms. The upper surface of the body
is covered with special flat-topped, pillar-like structures called paxillae. The body edges
are bordered with large, wide marginal plates. Its tube feet are pointed (not tipped with
suckers). Diameter with arms is about 10 cm.
Food and feeding: It feeds largely on detritus and small invertebrates.
Parasite: The sea snail, a marine gastropod Asterolamia cingulata, parasitizes this species.

Cytotoxic activities: The polyhydroxysteroidal glycosides, viz. hesperusides A–C and
novaeguinoside A isolated from this species showed cytotoxic activities against human
leukemia MOLT-4, human hepatoma BEL-7402, and human lung cancer A-549 cell lines.
While the compound hesperuside B exhibited cytotoxicity against all the tested human
tumour cells, compounds hesperuside A and novaeguinoside A were partially active
against A-549 and BEL-7402 cells (Kang et al., 2016).
In vitro Cytotoxicity (IC50: µM) of Glycosides 1–3

Against Three Tumour Cell Lines
Cell Line Hesperuside A Hesperuside B Hesperuside C
A-549 3.62 1.84 2.40
MOLT-4 2.59 0.68 2.12
BEL-7402 5.26 2.67 5.72
Source: Kang, J. et al., Mar. Drugs, 14, 189, 2016.

Psilaster cassiope (Sladen, 1889)
Global distribution: Gulf of Mexico
Ecology: It lives in deep-sea corals and deep-reef habitats at depths of 800–1,000 m.
Biology
Description: This robust species has five arms that taper smoothly to an acute point. The
disc is broad, and the paxillar area on the arms, relatively broad at the base, becomes
extremely narrow by the end of the arm. The oval paxillae are in regular transverse rows
across the arm and are small. They are short, bearing a flat, even covering of 15–25 granules.
Cytotoxic activities: The asterosaponin, psilasteroside, isolated from this species showed
cytotoxic activities against rat basophilic leukemia RBL-2H3 cell lines (Ivanchina et al.,
2011b).
2.6 FAMILY: CTENODISCIDAE (ORDER: PAXILLOSIDA)

Ctenodiscus crispatus (Bruzelius, 1805)
Common name(s): Mud star, cookie-cutter sea star

Global distribution: Polar to tropical regions; throughout North Atlantic and North Pacific
Ocean, including Canada, Greenland, Japan, Mexico, Panama, Russia, and US
Ecology: This benthic species is found on soft mud and rock or sand; the depth range is
10–1890 m.
Biology
Description: The shape of the body of this species is star-shaped, with a wide flattened
disc on the dorsal side. Specimens with five rays are relatively rare, and there are instances
with four and six rays. Ambulacral legs are conical, devoid of suckers, and are arranged
in two rows. The dorsal side is covered with tightly spaced low paxillae. There is a clear
disc in the middle of the disc. Marginal plates are high, forming a rigid side frame of the
rays. The colour is pale and flesh coloured, yellowish, dirty/sandy, sometimes light brown.
Normal size of the specimen is 2–4 cm and rarely up to 5 cm in diameter.
Food and feeding: It is a nonselective deposit feeder.

Cytotoxic and apoptotic effects: The methanol extract of this species has yielded five steroids,
(22E,24ξ)-26,27-bisnor-24-methyl-5α-cholest-22-en-3β,5,6β,15α,25-pentol 25-O-sulfate (1),
(22E,24R,25R)-24-methyl-5α-cholest-22-en-3β,5,6β,15α,25,26-hexol 26-O-sulfate (2), (28R)-24-
ethyl-5αcholesta-3β,5,6β,8,15α,28,29-heptaol-24-sulfate (3), (25S)-5α-cholestane-3β,5,6β,
15α,16β,26-hexaol (4), and Δ7-sitosterol (5), Among these compounds, compound 4 showed
cytotoxicity against human hepatoma HepG2 and glioblastoma U87MG cells via inhibition
of cell growth and induction of apoptosis (Quang et al., 2014).
2.7 FAMILY: LUIDIIDAE (ORDER: PAXILLOSIDA)

Luidia clathrata (Say, 1825)
Common name(s): Grey sea star, lined sea star

Global distribution: Temperate and subtropical; Western Atlantic: From US to Brazil and
Belize
Ecology: This benthic species is found at depths from 0 to 175 m but usually encountered
in shallow waters less than 40 m, on soft bottom habitats.
Biology
Description: This sea star has a small central disc surrounded by five long, flat arms. The
arms of this species are two to three times the size of the disc diameter. The surface of the
body is covered in plates. The upper surface of the body ranges from grey to light brown,
rose, or salmon colour. It has a dark grey or black stripe on the dorsal midline of each arm.
It is large, growing to 20–30 cm in length.
Food and feeding: It is a forager that can feed on a variety of different taxa including fora-
miniferans, nematodes, ostracods, gastropods, bivalves, crustaceans, as well as sediment
and detritus. It prefers to feed on infaunal bivalve Mulinia lateralis. It may shift from intra-
oral macrofaunal feeding to intraoral and extraoral detrital feeding. This sea star obtains
its food by ingesting sand and mud and then straining this material through oral spines.
When it is buried, it will invert its stomach to feed on detritus.
Reproduction: It spawns annually and has one larval stage before metamorphosis, a large
bipinnaria larva (2 mm long) that is competent (ready for metamorphosis) within one
month.
Associated species: Adults have a commensal polychaete worm, Podarke obscura Verrill,
living in the ambulacral groove.
Regeneration: The arms of this species can be lost as a result of predation. Regeneration
of the exposed end of the damaged arm begins by immediately sealing the damaged area.
A new tip appears in approximately one week.
Antibacterial activity: The sulphated sterol compound, isolated from this species, inhib-
ited the growth of both bacterial species (B. subtilus and S. aureus) at 50 µg/disc (Carvalhal
et al., 2018).
Iorizzi et al. (1995) reported on the isolation of polyhydroxysteroids (1–10); hydroxylated

steroids (11–13); and asterosaponins (14–17) from the EtOH body wall extract of this spe-
cies. Among these compounds, the polyhydroxysteroid (compound 10) and the hydroxyl-
ated steroid (compound 12) inhibited the growth of Bacillus subtilis and Staphylococcus
aureus. At a concentration of 50 µg per disc, compound 10 inhibited the growth of both
bacterial species, with 1- and 3-mm zones of growth inhibition for S. aureus and B. sub
tilis, respectively. At a concentration of 50 µg per disc, compound 12 caused 1- and 2-mm
zones of inhibition for S. aureus and B. subtilis, respectively.
Luidia maculata (Müller and Troschel, 1842)
Common name(s): Eight-armed Luidia sea star
Global distribution: Tropical; Indo-West Pacific
Ecology: This reef-associated species is found on soft, silty shores, near seagrass meadows
and coral rubble. It is usually seen alone and not in large groups. It moves rapidly and is
usually more active at night. It is also exposed or buried in the sand at depths of 0–90 m.
Biology
Description: It has five to nine (usually eight) arms. The arms are long, somewhat rounded
in cross section, and tapered to a sharp tip, edged with small sharp spines along the sides.
The upper surface of the body is covered with special flat-topped, pillar-like structures
called paxillae. The underside is pale, and from grooves along the arms emerge large tube
feet with club-like, pointed tips. Colours and patterns on the upper side are highly variable
in shades of greyish blue, to brown and beige. Size of the specimen is up to 60 cm.
Food and feeding: It is carnivorous, feeding on small buried animals such as molluscs and
other echinoderms.

Antioxidant activity: Among the five fractions of this species studied, fractions 3, 4, and
5 exhibited antioxidant potential. In DPPH assay, fraction 3 showed the highest IC50 value
0.27 mg/mL and hydrogen peroxide scavenging assay showed the highest IC50 value of
0.07 mg/mL in fraction 5. Whereas, fraction 3 showed the highest absorbance (0.3) in
reducing power assay when compared to fractions 4 and 5 (Suguna et al., 2014).
Antimicrobial properties: The ethanolic extract of this species displayed antibacterial
and antifungal activities against five bacterial and five fungal pathogens. In antibacterial
assay, Escherichia coli was the most sensitive pathogen against n-butanol extract as well as
fractions, and the highest zone of inhibition was shown in fraction 3 on E. coli (7.67 mm).
In antifungal assay, the highest zone of inhibition (23.33 mm) was observed in n-butanol
extract against Penicillium sp. and the lowest zone of inhibition (0.33 mm) was observed in
fraction 4 against Penicillium sp. (Anbukkarasu et al., 2014).
Others: Anon. (http://shodhganga.inflibnet.ac.in/bitstream/10603/174118/13/13_chapter_03.
pdf) reported on the isolation of polyhydroxy and epoxy-polyhydroxy steroids from this species.
Luidia quinaria (von Martens, 1865)

Common name(s): Spiny sand sea star
Global distribution: Western Pacific: Japan, China, Taiwan, and the Philippines
Ecology: This benthic, subtropical species lives in shallow water on soft sediments; the
depth range is 0–218 m.
Biology
Description: It has a small central disc and five long, slender arms fringed with short
spines. The aboral (upper) surface is covered in small paxillae, pillar-like spines with flat
tops giving a smooth, table-like surface. The usual colour of the aboral surface is grey with
contrasting pink or orange margins to the arms. The oral (under) surface is paler.
Food and feeding: It largely feeds on other echinoderms, in particular the brittle star
Ophiura kinbergi.

Cytotoxic activities: The glycoside, luidiaquinoside (an asterosaponin), isolated from this
species showed cytotoxic activities against rat basophilic leukemia RBL-2H3 cell lines
(Ivanchina, et al., 2011b).
cholesterol-binding ability with the value of 31.7% (Farhana, 2016).
Others: Andriyashchenko et al. (1996) reported on the isolation of five steroid compounds
from this species viz. 5α-cholestane-3β,5,6β,15α,16β,26-hexaol 3-sulfate (1), 5α-cholestane-
3β,5,6β,15α,26-pentaol 15-sulfate (2), sodium (24S)-O-(β-d-giucopyranosyll-5α-cholestane-
3β,6α,8,15β,24-pentaol 6′-sulfate (3), sodium (24S)-5α-cholestane-3β,6α,8,15β,24-pentaol
24-sulfate (4), and sodium tornasterol A sulfate (5).
Luidia sarsii (Düben and Koren in Düben, 1844)
Global distribution: Temperate species; Northeast Atlantic and the Mediterranean: North
and Celtic Seas
Ecology: It is a deeper water, epibenthic species; the depth range is 10–1292 m. The spe-
cies is usually found on muddy sediment and is most active at night, burying itself under
the sand during the day.
Biology
Description: This sand-coloured species has a velvety texture. Adults express pentamerism
or pentaradial symmetry. The five gently tapering arms have conspicuous bands of long,
white marginal spines in groups of three. This species grows to approximately 20 cm across.
Food and feeding: It feeds exclusively on the brittle stars Ophiura albida and Amphiura
filiform and the sea urchin Echinocardium cordatum. It exhibits diurnal rhythmic activity,
which is controlled by light, and it seeks its food actively.
Reproduction: The planktonic bipinnaria larva of this species attains a rather large size.
In the late bipinnaria, the larva develops a five-armed rudiment—concordant with the
number of arms of the adult—which develops from the rudiment. The larva has a “stalk”
with extensive ciliary bands; the larva also has larval arms and ciliary girdles. The juvenile
disc has tube feet that are already active before the release of the starfish from the larva.
The larval development is long, more than one year.

53126, Pseudomonas aeruginosa CIP 82118, and Salmonella enterica CIP 8039 with MIC
values of 12.5, 3.1, 6.3, 6.3, and 6.3 mg/mL, respectively (Marmouzi et al., 2018).
2.8 FAMILY: PORCELLANASTERIDAE (ORDER: PAXILLOSIDA)

Styracaster caroli (Ludwig, 1907)
Global distribution: Indian Ocean, north of Madagascar and in the Bay of Bengal
Ecology: It lives on a bottom of globigerina ooze and brownish ooze; the depth range is
2600–4820 m.
Biology
Description: This species is characterized by its naked ventrolateral plates, which are
arranged in about 5–6 tangential rows and farther in radiating rows, not extending outside
the disc area. Radius of the arm is 65 mm.

HIV-Inhibitory activity: Two sulphated sterols isolated from this species displayed
anti-HIV activity. While the first sterol inhibited HIV-1 and HIV-2 with IC50 values
of l showed IC50 values of >174 μM, the second sterol showed IC50 values of >164 μM
(McKee et al., 1994).
Others: De Riccardis et al. (1993) reported on the isolation of polyhydroxysteroids, caro-

listerols A-C from this species.
2.9 FAMILY: ECHINASTERIDAE (ORDER: SPINULOSIDA)

Echinaster (Othilia) brasiliensis (Müller and Troschel, 1842)
Common name(s): No common names found
Global distribution: Western Atlantic: Caribbean; Florida to Brazil
Ecology: Benthic. This tropical, benthic species lives in moderately shallow waters with a
depth range of 20–100 m.
Biology
Description: The disc of this species is small, and the five arms are stout, cylindrical, and
taper to a subacute tip. The regular rows of roughly triangular plates are connected by oval
or rectangular secondary plates. The spines of the marginals are larger and more thornlike
than those of the other plates. The madreporite is small, oval, raised, covered with radiat-
ing gyri, and without spinules or with a few small ones. The tips of the arms turn up, and
the oculars are small and broader than long.
Associated species: Three species of janirid isopods; Janaira gracilis, Carpias nereus and
Carpias asterophilus

Anti-HIV activity: The sulphated sterols isolated from this species showed inactivation
against HIV at the concentration of 100 µg/mL (McKee et al., 1994; Kim, 2013).
Others: This species serves as an ethnomedicine for the treatment of asthma (Anon.,
https://getd.libs.uga.edu/pdfs/narchi_nemer_e_201105_phd.pdf).
Two asterosaponins, viz. marthasteroside A1 brasiliensoside and ten glycosides of polyhy-
droxysteroids including echinasterosides and laeviusculosides C and I have been isolated
from this species. Further, a series of anthraquinones (animal pigments) have also been
isolated from this species (Iorizzi et al., 1993).
Echinaster luzonicus (Gray, 1840)

Common name(s): Luzon sea star
Global distribution: It is found in the tropical and subtropical Western Indo-Pacific region.
Its range extends from Madagascar and the east coast of Africa to Northern Australia,
Indonesia, and the Philippines.
Ecology: It is found on both reef crests and in the intertidal zone. It sometimes lives symbi-
otically with a copepod or a comb jelly, and it is prone to shed its arms, which then regener-
ate into new individuals.
Biology
Description: It is normally a six-armed starfish but is often rather asymmetrical in appear-
ance because of its habit of shedding arms. It is somewhat variable in colouring, ranging
from red to dark brown. Individuals seem to change their colour from red to brown and
back again, possibly as a response to the amount of ambient light they receive.
Food and feeding: It feeds on bacterial and algal films that it extracts from the sediment.
Reproduction: This species is unique in its genus in that it reproduces asexually by autoto-
mizing its arms; the shed arm then regenerates, growing a new disc and additional arms
[5]. This species has not been recorded breeding in any other way.
Associated species: A species of copepod, Paramolgus sp., lives symbiotically on the oral
(under) surface of Echinaster luzonicus; it is so cryptically coloured as to be almost indis-
tinguishable from its host. Another associate of this starfish is the comb jelly, Coeloplana
astericola, which grows in abundance on its aboral (upper) surface.

Anticancer activity: The cyclic steroid glycosides, viz. luzonicosides A, D isolated
from this species have shown anticancer activity in human melanoma RPMI-7951 and
SK-Mel-28 cell lines by inhibiting the proliferation and the formation of colonies, and the
migration of cells. The molecular mechanism of action appears to be related to the regula-
tion of the activity of cleaved caspase-3 and poly(ADP-ribose) polymerase (PARP), along
with Survivin, Bcl-2, p21, and cyclin D1 level (Malyarenko et al., 2017).
Lysosomal activity: Kicha et al. (2015) reported on the isolation of six steroid glycosides,
viz. luzonicosides A-F from this speices.Among these compounds, luzonicoside A at concen-
trations of 0.01-0.1 μM was shown to be potent in lysosomal activity stimulation, intracellu-
lar ROS level elevation, and NO synthesis up-regulation in RAW 264.7 murine macrophages.
Echinaster (Echinaster) sepositus (Retzius, 1783)
Common name(s): Mediterranean red sea star, purple starfish, or blood star
Global distribution: East Atlantic, including the Mediterranean Sea
Ecology: It is found at depths of 1–250 m in a wide range of habitats, including rocky,

sandy, and muddy bottoms, and seagrass meadows (Posidonia oceanica and Zostera sp.).
Biology
Description: It has five relatively slender arms around a small central disc. It usually has a
diameter of up to 20 cm but can exceptionally reach up to 30 cm. It is a bright orange-red
in colour and has a soapy surface texture. The surface is dotted with evenly spaced pits
from which the animal can extend its deep red gills (papullae). It reaches a diameter of up
to 25 cm.
Food and feeding: It feeds on algae.
Reproduction: The eggs of this species are also red. They do not live through a larval
stage but are immediately small starfish.
Predator: One of the predators of this species is the Atlantic Triton Trumpet.

Antibacterial and antioxidant activities: This species has shown lysozyme-like activity
(mean diameter of lysis of 13.4 mm), an antimicrobial activity against the human emerging
pathogens Staphylococcus aureus, Pseudomonas aeruginosa, and Candida famata, and a
strong lytic activity (100%) towards the human red blood cells. Further, this species has
shown the highest antioxidant activity (1765.65 nmolTE/mL) (Stabili et al., 2018).
Others: Free sterol mixture of this species has shown several Δ5 and Δ7-sterols includ-
ing (22E)-24-nor-5α-cholesta-7,22-dien-3β-ol (asterosterol), (22E)-27-nor-24(S)-methyl-
5α-cholesta-7,22-dien-3β-ol (amuresterol), and 5α-cholesta-7,22-dien-3β-ol (Desimone
et al., 1980).
Echinaster (Othilia) echinophorus (Lamarck, 1816)
Common name(s): Orange knobby star
Global distribution: Caribbean Sea and along the Atlantic coast of South America
Ecology: It occurs on shallow water areas such as reefs, rocks, and areas of coral rubble
and sometimes among mangroves at depths ranging from 24 to 73 m.
Biology
Description: It has a small central disc and five spatulate arms, tapering very little and
with rounded tips. The arms on the aboral (upper) side have one or two rows of bluntly
conical spines and more rows of spines on the sides of the arms and on the oral (under)
surface, on either side of the ambulacral grooves. The colour is usually some shade of red
or orange. It is a small species with a diameter of up to 7 cm.
Food and feeding: It feeds on many species of sponges.
Reproduction: It spawns in late spring and early summer. Two types of eggs are produced,
some being dark coloured and planktonic while others are bright orange and immediately
sink to the seabed. These soon start to develop into modified brachiolaria larvae, which have
larval arms and attach with a sucker. By the seventh day, they have two pairs of tube feet and
begin to move around. The development of the pelagic eggs takes place much more slowly.

Antileishmanial activity (antiprotozoal activity): The methanolic extract of this spe-
cies has shown antileishmanial activity by inhibiting the proliferation of promastigote and
amastigote forms with IC50 values of 62.9 and 37.5 μg/mL−1, respectively. This extract
also showed a moderate toxicity on macrophages from BALB/c mice. A dose of 100 mg/
kg/day was effective when administered during 15 days by intraperitoneal route to BALB/c
mice infected experimentally (ParraI et al., 2010; Oliveira et al., 2016).
Antimalarial activity: The crude extracts of this species showed antimalarial activity
against Plasmodium falciparum FcB1 strain and the recorded IC50 value was µg/mL
(Alonso et al., 2017).
Anticancer activity: The clarified extracts of this species showed anticancer activity
against 3LL and PC3 cancer cell lines with IC50 values of 265.7 and 405.6 µg/mL, respec-
tively (Alonso et al., 2017).
Henricia downeyae (A. M. Clark, 1987)
Common name(s): Slender-armed sea star
Global distribution: North Atlantic Ocean
Ecology: It can be seen on the beach, under rocks, in tidal pools when available, and on
gravel. Its aquatic biomes are the coastal and benthic zones.
Biology

Antimicrobial activity: The steroid glycosides, viz. downeyosides A-L; 26-nor echin-
asteroside A desulfated, 22(23)-dihydroechinasteroside A desulfated, echinasteroside A
desulfated, echinasteroside B desulfated, and echinasteroside C desulfated; laeviuscolo-
sides G, H, and 22,23-dihydro, laeviuscoloside I; and 5α-cholest-7-en-3β-yl-sulfate iso-
lated from this species have shown antimicrobial activity (Palagiano et al., 1996).
Liu (2011) isolated the compound, desulfonylated26-norechi-nasteroside A, from this

species.
Antifungal activity: The ethanolic body-wall extracts of this species have shown antifun-
gal activity against Sordaira fimicola (Bryan et al., 1994).
Cytotoxic activity: The sulphated steroid glucuronides, downeyoside A and B of this spe-
cies, have shown cytotoxic activity against two non-small cell lung human carcinoma cell
lines with IC50 of 60 and 36 μg/mL, respectively (Palagiano et al., 1995a).
Henricia leviuscila (Stimpson, 1857)
Common name(s): Pacific blood star, blood star, blood starfish

Global distribution: From Alaska to Baja California
Ecology: Its habitat is the intertidal zone under rocks and protected places from the low-
tide line to about 400 m deep.
Biology
Description: This species is identified by its bright orange-red colour, but there can also
be many variations from tan to almost purple. The disc can be a mottled grey colour.
There can also be a saddle-like marking of lilac blotches between the rays, but the rays are
not mottled. Usually, it has five rays (occasionally four to six). The rays are smooth and
appear smooth due to the lack of pedicellariae and spines. The species is relatively small
and its diameter is usually more than 8 cm and rarely gets larger than 12 cm.
Food and feeding: It mainly feeds on sponges and small bacteria. This sea star moves
these tiny particles, which are captured in mucus and swept to the mouth by ciliated tracts.
It may also feed by applying the stomach to the surfaces of sponges and bryozoa.
Reproduction: In this species, the sexes are dioecious and females are not known to brood
young. Embryonic stages do not adhere to one another but float freely. Post-hatching larvae
are ciliated and swim.
Associated species: It often has a commensal scale worm, Arctonoe vittata.

Anticarcinogenic action: The steroid biglycoside, leviusculoside G, from this species dem-
onstrated anticarcinogenic action by the induction of p53-dependent apoptosis and inhibition
of activator protein 1 (AP-1), nuclear factor kappa-light-chain-enhancer of activated B cells
(NF-κB), and extracellular signal-regulated protein kinases (ERKs) activities in human leu-
kemia HL-60, THP-1, and mouse epidermal JB6 Cl41 cells (Malyarenko et al., 2015).
Haemolytic activity: Five polar steroids, polyhydroxysterols (2–5), a glycoside leviuscu-
loside J (7), and another 4 steroids (1, 6, 8, and 9) have been isolated from the alcoholic
extract of this species. Among them, the compounds, 1, 3, 6, 7, and 9 showed moderate
haemolytic activity in the mouse erythrocytes assay (Ivanchina et al., 2006).
Ivanchina et al. (2011b) also reported on the occurrence of sanguinosides A–C and levi-
usculoside J from this species. Of these compounds, leviusculoside J showed a moderate
haemolytic activity in the mouse erythrocytes assay.
2.10 FAMILY: PTERASTERIDAE (ORDER: SPINULOSIDA)

Diplopteraster multipes (M. Sars, 1866)
Common name(s): Pincushion star
Global distribution: Circumpolar: from Artic to Chesapeake Bay and Ireland in the
Atlantic and from Japan to US in the Pacific
Ecology: This temperate, benthic species is found on sand, mud, and gravel; the depth
range is 57–1225 m.
Biology
Description: This species is large and looks like a pentagonal cushion with a rather thick
“cushion cover” (supradorsal membrane). It is also reinforced with 8–10 spikes that pro-
trude under the membrane (paxilles). In addition to the size and pillow shape, it has four
rows of suction feet. On the underside, along the edge of the suction feet there are two dif-
ferent rows across the fence; and 4–5 spikes turn down to the fence while 3–4 spikes turn
the opposite way. All rows of spikes are woven together with a thin membrane. Along the
side of the mouthplates, there are 5 spikes (oral spikes), all of which are woven together
with a thin membrane. The species is light brown and is about 20 cm in diameter.
Food and feeding: In the stomach of this species, snake stars (ophiuroid, Astrotoma agas
sizii) and sediments have been found as a diet.

Blood clotting and the immunity: The cholesterol sulphate of this species has been reported
to regulate the activity of serine protease, the main function of which is digestion in humans.
However, this enzyme also functions in processes such as inflammation, blood clotting, and
the immune system in both prokaryotes and eukaryotes (Jha and Zi-rong, 2004).
Others: Sodium salt of (20R)-3α,4β-dihydroxycholest-5-ene-21-yl sulfate and disodium
salts of (20R)-4β-hydroxycholest-5-ene-3α,21-diyl disulfate, (20R)-24-methylcholest-
5,24(28)-diene-3α,21-diyl disulfate, (20R)-24-methyl-5α-cholest-24(28)-ene-3α,21-diyl
disulfate, (20R)-cholest-5-ene-3α,21-diyl disulfate, (20R)-5α-cholestane-3α,21-diyl disul-
fate, and (20R)-3α-hydroxycholest-5-ene-2β,21-diyl disulfate have been isolated from this
species (Levina et al., 2002).
Pteraster pulvillus (M. Sars, 1861)
Common name(s): Orange cushion star
Global distribution: It is a circumboreal species and is found in the Arctic and Northwest
Atlantic to Cape Cod; it is also available in the Pacific and Bering Sea.
Ecology: This benthic, temperate species is found in rocky areas at depths of 36–3700 m.
Biology
Description: It has clusters with spikes (paxilles) on the upper side, with 6–15 spikes per
cluster. It also has 6–7 oral spikes (oral spikes) per mouthplate, all of which are woven
together. The species looks like a small five-legged pillow, with “cushion cover,” or supra-
dorsal membrane. There are 6–15 spikes in each of the spikes. The species is yellowish
brown on the upper and lower sides. On the underside, along the fist with the suction feet
(adambula beads), there are 4–6 spikes per plate across and they are all woven together
with a thin membrane. Alongside each mouthplate, there are 6–7 spikes that are also woven
together with a thin membrane. The species is 4–5 cm in diameter.
Food and feeding: It eats sponges, hydroids, and tunicates. Some deep-water species are
also thought to eat sediment.

Haemolytic activity: Six steroidal disulfates, namely, disodium salts of (20R)-cholest-5-
ene-3α,4β,21-triol 3,21-disulfate (1), (20R)-5α-cholestane-3α,4β,21-triol 3,21-disulfate (2),
(20R)-5α-cholestane-2β,3α,21-triol 3,21-disulfate (3), and (20R)-5α-cholestane-3α,21-diol
3, 21-disulfate (4), the dityrammonium salt of (20R)-5α-cholestane-3α,21-diol 3,21-disul-
fate (5), and a mixture of sodium and tyrammonium salts (1:1) of (20R)-cholest-5-ene-
3α,21-diol 3,21-disulfate (6) have been isolated from the ethanolic extracts of this species.
Among these steroids, compounds 1, 2, and 4–6 showed haemolytic activity to mouse
erythrocytes with HC50 values of 8.0 × 10 −5, 4.5 × 10 −5, 1.0 × 10 −5, 1.8 × 10 −5, and
3.3 × 10 −5 M, respectively (Ivanchina et al., 2003).
2.11 FAMILY: ACANTHASTERIDAE (ORDER: VALVATIDA)

Acanthester planci (Linnaeus, 1758)
Common name(s): Crown-of-thorns starfish
Global distribution: It has a very wide Indo-Pacific distribution. It is perhaps most com-
mon in Australia but can occur at tropical and subtropical latitudes from the Red Sea and
the east African coast across the Indian Ocean, and across the Pacific Ocean to the west
coast of Central America.
Ecology: It occurs where coral reefs or hard coral communities exist.
Biology
Description: It is one of the largest starfish in the world. The body form of this species is
fundamentally the same as that of a typical starfish, with a central disc and radiating arms. It
has disc-shaped, multiple-armed, flexible, prehensile, and heavily spined characteristics, and
it has a large ratio of stomach surface to body mass. Though it is multiple armed, it has lost
the fivefold symmetry (pentamerism) typical of starfish. These organisms have up to 21 arms.
Although the body of the crown of thorns has a stiff appearance, it is able to bend and twist to
fit around the contours of the corals on which it feeds. The underside of each arm has a series
of closely fitting plates, which form a groove and extend in rows to the mouth. The long, sharp,
and venomous spines on the sides of the starfish’s arms and upper (aboral) surface resemble
thorns. The spines are stiff and very sharp, and they readily pierce through soft surfaces. The
organisms are usually of subdued colours, pale brown to grey-green, but they may be garish
with bright warning colours. Adult starfish normally range in size from 25 to 35 cm.
Food and feeding: It preys upon hard or stony coral polyps.
Predators: The organisms are preyed upon by species of pufferfish and triggerfish; Triton’s
trumpet, a very large gastropod mollusc; small painted shrimp Hymenocera picta; and cni-
darian Pseudocorynactis sp.

Anticancer actiivty: Three new steroid biglycosides, plancisides A–C (1–3), have been
isolated from the ethanolic extract of this species. Among them, compound 2 showed
anticancer activity against HCT-116, T-47D, and RPMI-7951 cancer cell lines (Kicha
et al., 2014a).
Cytotoxic activity: Two isobenzofuranone derivatives, pseudaboydins A (1) and B

(2), along with five known compounds—including (R)-2-(2-hydroxypropan-2-yl)-2,
3-dihydro-5-hydroxybenzofuran (3), (R)-2-(2-hydroxypropan-2-yl)-2, 3-dihydro-
5-methoxybenzofuran (4), 3,3′-dihydroxy-5,5′-dimethyldiphenyl ether (5), 3-(3-methoxy-
5-methylphenoxy)-5-methylphenol (6), and (-)-regiolone (7)—have been isolated from the
marine fungus, Pseudallescheria boydii, associated with this species. Among these com-
pounds, pseudaboydin A (1) showed moderate cytotoxic activity against human nasopha-
ryngeal carcinoma cell line HONE1, human nasopharyngeal carcinoma cell line SUNE1
and human glandular lung cancer cell line GLC82 with IC50 values of 37.1, 46.5, and
87.2 μM, respectively (Lan et al., 2014).
Vien et al. (2018a) reported on the isolation of attenuatoside B-1, planciside A, and cul-
citoside C2 from this species. Among these compounds, culcitoside C2 (3) showed weak
cytotoxicity against five human cancer cell lines including HepG2, KB, LNCaP, MCF7,
and SK-MEL-2.
Regulating nitric oxide production: Nitric oxide (NO) is a widespread signalling mole-
cule that participates in virtually every cellular and organ function in the body. Pyrrole oli-
goglycosides such as plancipyrrosides A, B produced by this species have been reported to
suppress lipopolysaccharide-induced nitric oxide production in RAW264.7 macrophages
(Blunt et al., 2014).
Others: Itakura and Komori (1986) reported on the isolation of four new genuine oligo-
glycoside sulphates, named acanthaglycoside B, C, D, and F from this species (Itakura and
Komori, 1986).
Kicha et al. (2014b) reported on the presence of a new steroidal glycoside called planciside
D from this species.
Maier (2008) reported on the isolation of an asterosaponin, thornasteroside A, from this

species.
D’Auria et al. (1993) reported on the presence of 5-deoxyisonodososide, isonodososide,

and acanthaglycoside F from this species.
2.12 FAMILY: ARCHASTERIDAE (ORDER: VALVATIDA)

Archaster typicus (Müller and Troschel, 1840)
Common name(s): Common sea star, sand-sifting star, and sand star
Global distribution: Indo-Pacific region: Maldive Islands, the Bay of Bengal, Singapore,
northern Australia, New Caledonia, the Philippines, China, southern Japan, and Hawaii
Ecology: It inhabits areas of the seabed with soft sediments including sand, silt, and sea-
grass meadows and mangroves at depths down to 60 m.
Biology
Description: The body of this species is somewhat rounded (not flat). Arms are long and
tapered to a sharp tip and edged with short, flat, blunt spines. Most organisms have five
arms, but those with three, four, and six arms are also seen. The body is slightly inflated
and there is a whitish madreporite near the centre of the disc. The small armour plates
that cover the upper surface of the arms are lined up in neat parallel rows. The spines,
arranged in a marginal fringe, are short, flat, and blunter and the tube feet have suckers and
not points. Colours and patterns on the upper side are highly variable in shades of greyish
blue to brown and beige. The size is up to 15 cm across. The underside is pale, with large
tube feet tipped with suckers. The diameter of adults with arms measures 12–15 cm.
Food and feeding: It feeds on detritus, decaying plants, and tiny animals. To feed, it everts
its stomach through its mouth, which is situated centrally on its underside. The food item
is engulfed and brought inside the starfish when its stomach is kept in its normal position.
Reproduction: In this species, sexes are separate. It is a broadcast spawner and the male
and female starfish each liberate their gametes into the sea where fertilization takes place.
However, in contrast to most other starfish, this species performs pseudocopulation in
which males are often found stacked on top of the females in an unusual aggregation.
Individuals reach sexual maturity at a radius of 29 mm. About two months ahead of
spawning, the starfish begin to congregate. The male or female is identified probably by
chemotactic recognition. On recognizing a female, the male will climb on top of her and
may remain there for about two months. During this time, they synchronize their gonadal
activity so that when the female is ready to spawn, so is the male. When she releases her
eggs, he releases his sperm almost simultaneously, thereby ensuring successful fertiliza-
tion will take place. Larval settlement occurs among mangroves, while individuals gradu-
ally move to seagrass and sandy habitats as they grow.

Cytotoxic activities: The asterosaponins archasterosides A and B and regularoside A iso-
lated from this species showed moderate cytotoxic activities against HeLa and mouse JB6
P+ Cl41 cell lines (Kicha et al., 2010).
Others: Eight highly hydroxylated steroids (1–8), including three new compounds as
sodium salts of (24S)-5α-cholestane-3β,4β,5,6α,7β,8,14,15α,24-nonaol 6-sulfate (1),
(24E)-5α-cholest-24-ene-26-yde-3β,6α,8,14,15α-pentaol 15-sulfate (2), and 5α-cholest-
3β,6α,8,14,15α,24,25,26-octaol 15-sulfate (3), have been isolated from the methanol
extract of this species (Hanh et al., 2016).
2.13 FAMILY: ASTERINIDAE (ORDER: VALVATIDA)

Patiria pectinifera (Muller and Troschel, 1842) (= Asterina pectinifera)
Common name(s): Blue bat star
Global distribution: Northern Pacific Ocean along the coasts of Japan, China, and Russia
Ecology: It inhabits a shallow subtidal zone on stony seabeds and other substrates down to
depths of 40 m. It prefers living on coarse sediment to fine sediment.
Biology
Food and feeding: It feeds on algae and seagrasses, detritus, and small invertebrates.
Predators: It may be preyed on by the carnivorous starfish Luidia quinaria.
Reproduction: This starfish breeds twice a year, in the autumn and the spring. Females
spawn about 500,000 eggs each year.
Research uses: Individuals of this species are used as model organisms in developmental
biology.

The methanol and water extracts were found to be the most active, and Aspergillus spp. and
Cryptococcus neoformans proved to be sensitive species (Choi et al., 1999).
cholesterol-binding ability (Farhana, 2016).
Neuritogenic and neuroprotective effects: The polar steroids, viz. asterosaponin
Р1, (25S)-5α-cholestane-3β,4β,6α,7α,8,15α,16β,26-octaol and (25S)-5α-cholestane-
3β,6α,7α,8,15α,16β,26-heptaol (1–3) of this species have been reported to enhance neu-
rite outgrowth in NB cells. Dose-dependent responses to compounds 1–3 were observed
within the concentration range of 10–100 nM. Further, these compounds also served as
neuroprotectors against oxygen-glucose deprivation (OGD) (Palyanova et al., 2013).
Anti-inflammatory activity and cytotoxicity: Purified cerebrocides isolated from this

species displayed cytotoxic activity against two human cancer cells in a dose- and time-
dependent manner up to 400 µg/mL. Further, a polysaccharide of this species inhibited the
growth of human breast and colorectal cancer cells. A fraction of this species exhibited
strong anti-inflammatory activity and cytotoxicity against cancer cells in a dose-dependent
manner. At 25 µg/mL, this fraction inhibited NO release by 92.4% (IC50, 14.6 µg/mL)
without causing cytotoxicity (cell viability >90%), which suggested that this fraction con-
tained NO-inhibitory compounds. However, at 50 µg/mL, the said fraction was cytotoxic
(cell viability approximately 10%). Two polyhydroxysteroids of this species were also cyto-
toxic to the HL-60 cells, with IC50 values of 80.3 and 40.5 μM (Wikarta and Kim, 2016).
Anti-melanogenic effect: The control of melanogenesis is an important strategy in the
treatment of abnormal skin pigmentation for cosmetic purposes. The extracts of this spe-
cies inhibited melanogenesis by reducing tyrosinase activity and melanin production via
subsequent downregulation of tyrosinase-related proteins. These extracts of this species
may therefore be promising candidates for the treatment of hyperpigmentation disorder
and useful for self-tanning cosmetic products (Jeong et al., 2013).
Antibacterial activity and human skin cell line protection: The compound phytosphingo-
sine isolated from this species has been reported to protect the HaCaT cell line (cell line from
adult human skin) against injuries caused by stimulation to 10 µg/mL mite antigen for 1 hour.
Furthermore, the above compound could significantly inhibit the growth of S. aureus. This
result implied that the application of phytosphingosine isolated from this species of starfish
might be a promising therapeutic option of atopic dermatitis (Choi et al., 2010).
Antiviral activity: A monosulfated sterol and the polyhydroxylated sterol 333 asterosapo-
nin P2 isolated from this species exhibited activity against HSV-1, with MIC values of
0.2 and 0.07 µM, respectively. Similarly, another compound—halistanol trisulfate C—
showed similar activity against HSV-1 (KOS 337 strain), with an IC50 value of 6.09 µg/mL
(Carvalhal et al., 2018).
Chemopreventive agent: From the butanol fraction of the starfish Asterina pectinifera
(Müller and Troschel, 1842) (Asteriidae), we have isolated a new component. The com-
pound 5α-cholest-7-en-3β-ol isolated from the butanol fraction of this species exhibited
antigenotoxic and antimutagenic activities with Escherichia coli PQ37 and Salmonella
typhimurium TA1538, respectively. These results suggest that 5α-cholest-7-en-3β-ol might
be useful as a chemopreventive agent (Han et al., 2000).
Others: A novel pyrrole oligoglycoside 3-{O-β-d-fucopyranosyl-(1→3)-β-d-fucopyranosyl-
(1→4)-[β-d-quinovopyranosyl-(1→2)]-β-d-quinovopyranosyl}-2-acetyl-pyrrole was isolated
from the whole body of this species (Zhang et al., 2006a).
Sugita (1977) reported on the isolation of glycosphingolipids, ceramide mono- and di-
hexosides, from this species.
2.14 FAMILY: ASTEROPSEIDAE (ORDER: VALVATIDA)

Asteropsis carinifera (Lamarck, 1816)
Common name(s): Sheriff-badge sea star
Global distribution: Indo-West Pacific Ocean: Red Sea, Mozambique, Seychelles, and
Kenya to the Solomon Islands
Ecology: This tropical, benthic species is found in intertidal areas, particularly on rocky
bottoms and protected coral reefs. It emerges at night to hunt prey.
Biology
Description: Size: No other information is available for this aspect except its size, which is
15 cm.
Food and feeding: It is a deposit feeder and feeds occasionally on cone shells on experi-
mental observations.
Cytotoxicity: A total of 12 steroidal biglycosides, viz. 6 new compounds, cariniferosides
A-F (1–6) and 6 already known compounds (7–12) [halitylosides A, B, D, E, (7, 8, 9,
10); and halityloside A 6-O-sulfate (11) and 4″-O-methylhalityloside A 6-O-sulfate(12)]
have been isolated from the alcoholic extract of this species. The in vitro cytotoxicity of
glycosides 1 and 6–12 against human cancer HCT-116, T-47D, and RPMI-7951 cells was
evaluated. Compounds 6 and 9–12 did not show any apparent cytotoxicity against all of the
tested cells within a 10–160 µM concentration range. Glycoside 8 was nontoxic towards
the HCT-116 cell line in the same concentrations and exhibited slight cytotoxicity against
T-47D and RPMI-7951 cells with IC50 = 154 and 128 µM, respectively. Compound 7
demonstrated a similar cytotoxic effect against only HCT-116 cells with IC50 = 150 µM.
Glycoside 1 displayed some stronger cytotoxic action in all of the investigated cell lines,
with IC50 values that ranged from 32 to 66 µM in comparison with the other studied com-
pounds (Malyarenko et al., 2011).
In vitro Cytotoxic Activity and Inhibition of Cancer Cell Formation

by Compounds 1 and 6–12
Compound HCT-116 IC50 (µM) T-47D IC50 (µM) RPMI-7951 IC50 (µM)
1 32 37 66
6 >160 >160 >160
7 150 >160 >160
8 >160 154 >160
9 >160 >160 128
10 >160 >160 >160
11 >160 >160 >160
12 >160 >160 >160
Source: Malyarenko, T.V. et al., Steroids, 76, 1280–1287, 2011.
Dermasterias imbricata (Grube, 1857)

Common name(s): Leather star
Global distribution: The range of the leather star includes the western seaboard of North
America from central Alaska to northern Mexico.
Ecology: This temperate, benthic species lives in intertidal areas at depths of about 100 m.
It is found mostly on rocks, sand, or mud. It seems to prefer at least partially sheltered
areas.
Biology
Description: It has a broad central disc and five plump, short arms, which taper broadly
from the central disc. The arms have two rows of tube feet and there are no bordering mar-
ginal plates. The upper surface is smooth and velvety, covered with a reticulated pattern
in reddish brown, often with patches of greyish blue. No pedicellariae are seen, but the
madreporite can be seen. This starfish can grow to 25 cm in diameter. It has a distinctive
smell that resembles garlic and sulphur.
Food and feeding: It feeds on algae and a range of invertebrates, including other asteroids,
bryozoans, sea urchins, sponges, sea cucumbers, hydroids, sea pens, and colonial tuni-
cates. It usually swallows its prey whole and digests them internally.
Predator: It is preyed on by the morning sun star (Solaster dawsoni).
Reproduction: Off the Washington coast, spawning of this species is from April to August.
The females release yellow eggs, which are fertilized in the water column. The larvae then
become part of the zooplankton.
Associated species: The leather star sometimes lives symbiotically with the scale worm
Arctonoe vittata.
Parasite: The parasitic barnacle genus Dendrogaster is sometimes an endoparasite of the
leather star.

Antifungal activity: The saponins isolated from this species have shown antifungal activ-
ity (Sumithaa et al., 2017a).
Others: Jha and Zi-rong (2004) reported on the isolation of a benzyltetrahydroisoquinoline
alkaloid imbricatine, the activity of which is yet to be known.
2.15 FAMILY: GONIASTERIDAE (ORDER: VALVATIDA)

Fromia heffernani (Livingstone, 1931) (= Celerina heffernani)
Common name(s): Heffernan’s starfish, Heffernan’s sea star, dark sea star
Global distribution: Western Central Pacific: Philippines to Solomon Islands and the
Great Barrier Reef
Ecology: This tropical, reef-associated species has a depth range of 5–40 m.

Biology
Description: In the centre of this species, there is a small disc from which five long,
thin, cylindrical but angular arms radiate. All these five arms are well separated and are
arranged around the disc at regular intervals. There may be four or six arms because of
growth or regeneration problems. The central disc is dark with predominantly violet or
purple. The radius of the disc is 5 cm maximum. On the oral side (substrate side), the arms
are dug out of a furrow from which the podias emerge. Each furrow is bordered on either
side of a single row of bifurcated, cream-coloured spines.
Food and feeding: It feeds on detritus and small invertebrates. It devails its stomach
to feed itself. The gastric juices directly attack the fixed organisms. Digestion is partly
external.
Reproduction: It is a gonochoric species in which the sexes are separated. Fertilization is
external. There is also a possibility of asexual reproduction (schizogony) in which the star
splits to give two individuals.

Antiviral (anti-HIV) activity: The compounds ptilomycalin A and celeromycalin isolated
from this species have shown antiviral activity with IC50 values of 10.11 and 10.32 µg/mL,
respectively (Motuhi et al., 2016).
Cytotoxic activity: Ptilomycalin A, crambescidin 800, and celeromycalin isolated from

this species were found to be highly cytotoxic to the target cells with CC-50 (mean 50%
cytotoxic concentration) of 0.11 µg/mL (Palagiano et al., 1995b).
Fromia monilis (Perrier, 1869)
Common name(s): Necklace starfish, tiled starfish, red tile starfish
Global distribution: Indian Ocean and Western Pacific, from the Andaman Islands up to
Australia and Japan
Ecology: This tropical, reef-associated species has a depth range of 0–51 m and is found
largely on reef slopes with rocky bottoms.
Biology
Description: In this species, the tips of the arms and the disc centre are bright red, while
the remaining parts are paler, forming large plates. The appearance of this sea star can be
highly variable (colours, plates, presence of plates on the central disc, armtips, etc.). It can
reach a diameter of about 30 cm.
Food and feeding: It feeds on encrusting sponges, detritus, or small invertebrates.
Aquarium values: This species is also considered in reef aquariums.

Antiviral (anti-HIV) activity: The compounds fromiamycalin and crambescidin 800 iso-
lated from this species have shown antiviral activity with IC50 values of 10.11 µg/mL
(Motuhi et al., 2016).
Cytotoxic activity: The compounds ptilomycalin A, crambescidin 800, and celeromycalin

of this species have shown cytotoxic activity to the target cells with CC-50 of 0.11 µg/mL.
Further, another compound (made up from the hydroxyspermidine residue linked to long-
chain ω-hydroxy acid) also exhibited a weaker cytotoxicity with CC-50 of 2.7 µg/mL
(Palagiano et al., 1995b).
Hippasteria phrygiana (Parelius, 1768) (= Hippasteria kurilensis)
Common name(s): Kurile spiny star, Pacific starfish, rigid cushion star
Global distribution: Northeast Pacific: Alaska
Ecology: This benthic species lives mostly in cold and deep waters at depths of 20–800 m.
In the North Sea it is found from the Shetland Islands down to Northumberland on the
British east coast. Elsewhere, it is northerly distributed from Scotland to Greenland,
Iceland, and Finmark.
Biology
Description: The body of this species has a large disc and five small, narrow, and tapering
arms. In the dorsal side, the plates are of various sizes. The larger plates bear a thick spine
or a pedicellaria in the middle and a series of graines around the edge. The smaller plates
are disposed irregularly around the larger ones. The marginal plates each bear 1–3 thick
spines in the middle and a series of graines along the edges. In the oral side, the interradial
area is very large, with numerous large pedicellariae. Along the ambulacral furrow, the
pedicellariae are arranged in a fairly regular longitudinal series. This species grows up to
20 cm in diameter.
Food and feeding: This species feeds mostly on cnidarians, especially deep-sea corals.

Cytotoxic activity: Cyclopropane-containing steroid phrygiasterol (1) and steroid glyco-
side phrygioside B (2), along with previously known borealoside C and (20R,24S)-5α-
cholesta-3β,6α,8,15α,24-pentaol (3), have been isolated from this species. Compound 1
inhibited the growth of Ehrlich carcinoma cells with an IC50 of 50 μg/mL, whereas 2
induced apoptosis of the same cells (EC50 = 70 μg/mL) and inhibited Ca2+ influx into
mouse spleenocytes (EC50 = 20 μg/mL) (Levina et al., 2005).
Anticancerogenic properties: Kicha et al. (2011) reported on the isolation of sulfated ste-
roidal glycosides (asterosaponins), hippasteriosides A–D, from the alcoholic extract of this
species. Among these compounds, hippasterioside D demonstrated a remarkable inhibition
of the HT‐29 colony formation in soft‐agar clonogenic assay, suggesting its anticancero-
genic properties.
Others: Three novel steroidal triglycosides, designated as kurilensosides A, B, and C

(1–3), were isolated along with a new steroidal diglycoside, kurilensoside D (4), and two
new (6,7) and one known (5) polyhydroxysteroid from the alcoholic extract of this species
(Kicha et al., 2008b).
Five new steroidal monoglycosides, kurilensosides E, F, G, and H and 15-O-sulfate of echin-

asteroside C have been isolated from the alcoholic extract of this species (Kicha et al., 2009).
Kichaa et al. (2009) reported on the isolation of compounds such as kurilensoside I and J,
lincoside F, echinasteroside A desulfated, forbeside L, lLinckoside LI, granuloside A, and
leviusculoside G (= forbeside J) (6) from this species.
Rosaster sp.
Nothing is known about its ecology and biology.

Antifungal activity: The compound (25S)-5α-cholestane-3β,4β,6β,7α,8,15α,16β,26-octol
isolated from this species has shown antifungal activity against Clodosporium cucumeri
num (Motuhi et al., 2016).
Stellaster childreni (Gray, 1840) (= Stellaster equestris)
Common name(s): Galloping sea star
Global distribution: Indo-Pacific from East Africa to Australia and southern China and
Japan
Ecology: Individuals of this species are found at a mixture of sandy and muddy substra-
tum, followed by a mixture of sandy and coral, muddy substratum, coral substratum, and
sandy areas at depths of 45–185 m.
Biology
Description: It has five long, skinny arms with large, smooth marginal plates on the edges
and sparse, short, stumpy spines. The oral side is white and smooth with colourful mark-
ings. It has short tube feet tipped with suckers. The abactinal (dorsal) plate of each arm
contains granules and blunted small spines and pedicellariae. Tube feet of the oral side are
relatively small and are embedded inside the adambulacral groove. The diameter of this
species with arms is 8 cm.
Food and feeding: It feeds on a variety of flora and fauna including Mollusca (gastropod,
bivalves, and scaphopods), sponge seagrass, and seaweed as well as benthic Foraminifera.

Antibacterial activity: The crude methanol and ethanol extracts of this species have shown
antibacterial activity. The crude methanol extract showed the maximum zone inhibition
(9.7 mm) against Escherichia coli and Vibrio parahaemolyticus at 100% concentration and
minimum was Staphylococcus aureus (4.06 mm). The crude ethanol extract showed the
maximum zone of inhibition in E. coli (9.70 mm) and the lowest concentration exhibited
the minimum inhibition activity against all bacterial pathogens. The fractions showed the
maximum inhibition zone in Klebsiella oxytoca (5.00 mm), Salmonella typhi (5.00 mm),
and Staphylococcus aureus (2.3 mm) (Prabhu and Bragadeeswaran, 2013a). Sumithaa et al.
(2017b) reported on the antibacterial activity of its crude and fractioned steroidal compound.
Cytotoxicity: The chloroform and the methanolic extracts of this species showed cytotox-
icity against human ovarian teratocarcinoma cell line PA 1 with an IC50 value of 35 μg/
mL (Sumitha et al., 2017b).
2.16 FAMILY: MITHRODIIDAE (ORDER: VALVATIDA)

Mithrodia bradleyi (Verrill, 1867) (= Mithrodia enriquetacasoi)
Common name(s): Bradley’s sea star
Global distribution: Eastern Pacific Ocean; Gulf of California and Lower California
Panama, Coast of Columbia and Galapagos Islands
Ecology: It inhabits rocky shallow coastal areas and tidepools.
Biology
Description: Dried specimens of this species show flat arms that are slightly constricted at
the base. Arm angles are rather sharp, except when two arms stand wide apart. Spines are
clearly spiniform, sometimes more cylindrical. The skeleton is more compact. Pedicellariae
vary in place and in number, but they are more numerous. The size of the specimen is
35 cm.
Antimicrobial activity: The ethanol extracts of this species showed antimicrobial activity
against Mycobacterium tuberculosis with the values of 10 (under BACTEC 460 system) at
concentration of 300 µg/mL (Encarnación et al., 2000).
Mithrodia clavigera (Lamarck, 1816)
Common name(s): Nail starfish, nail sea star
Global distribution: Indo-Pacific from the sea Rouge, to the Indian Ocean, to French
Polynesia and South of Japan, to the New Caledonia and Isle of Man Howe (Australia)
Ecology: This tropical starfish is found on all types of substrates rich in encrusting organ-
isms in lagoons or coral reefs from 10 to 80 m deep. It is active mainly at night.
Biology
Description: The disc (central part of the body) of this species is quite small. The five arms
are more or less cylindrical and are slightly narrowed at their base. They wear long spines
on their sides and shorter, more rarely, on their upper face. These tips are club-shaped, with
small tubercles at their ends. The pedicellaries have a form of forceps with 3–8 jaws. On
the underside of the arms, the ambulacral sulcus is lined with small prickles and there is
only one row of podia. It is a large starfish up to 50 cm wide. The base colour is beige to
dark brown, often with alternating lighter areas and darker areas. The body is covered with
vaguely triangular dark spots.
Food and feeding: This starfish feeds on encrusting organisms (sponges, corals, bryozo-
ans) and bivalves at night. It can also consume algae and organic waste. To feed, it devours
its stomach on its prey.
Reproduction: Sexual reproduction of this species has not been studied. However, as with
other starfish, the sexes must be separated, the gametes released into the water, and exter-
nal fertilization occurs. The regenerative power of starfish, and Mithrodia clavigera in par-
ticular, is very important. Each injured or amputated arm is able to grow back. Likewise,
a snatched arm is able to regenerate an entire starfish.
Associated species: It is common to observe small Zenopontonia soror shrimps on the
surface of this starfish, preferably on the ventral side. This starfish is host to ectoparasites,
Stellicola longiseta and Synstellicola longiseta.

Cytotoxic activity: The compounds such as sulfated polyoxide steroid (20S)-3β,6α,20-
trihydroxi-5α-cholest-9(11)-ene 3-sulfate (sodium salt) named mithrotriol, glycosides
echinasteroside B, granulatoside A, linckoside K, and forbeside L, as well as sulfates of
thornasterol A and cholesterol isolated from this species have shown cytotoxic activity in
human melanoma cells SK-MEL-28, SK-MEL-5, and RPMI-7951 (Levina, et al., 2012).
Thromidia catalai (Pope and Rowe, 1977)
Common name(s): Catala’s sea star, brown-tipped sea star, heavy starfish, fat sea star
Global distribution: Western Central Pacific: New Caledonia; South China Sea; from
Indonesia to Hawaii
Ecology: This tropical, benthic species inhabits lagoons and outer reefs and is seen on all
kinds of substrates; the depth range is 10–105 m.
Biology
Description: It is a large species of starfish with a maximum size of 100 cm in diameter. It
ranges in colour from white to yellow and orange with brown tips at the ends of its legs.
Food and feeding: It feeds mainly on detritus.

Three known sulphated “asterosaponins,” thornasteroside A, ophidianoside F, and regu-
laroside B; one known steroidal diglycoside, granulatmide A; and one new minor steroidal
monoglycoside, thromidioside, have been isolated from this species (Riccio et al., 1988).
2.17 FAMILY: OPHIDIASTERIDAE (ORDER: VALVATIDA)

Certonardoa semiregularis (Müller and Troschel, 1842)
Global distribution: East China Sea; in Japan, in particular, Honshu Island, Kyushu
Ecology: It inhabits exposed intertidal and sheltered subtidal habitats and coral reefs.
Biology
Description: This species has five, slender, tapering arms. There are regular, longitudinal
and transverse rows on the dorsal side. The ventral side is paler than the dorsal side.

Cytotoxicity and antibacterial activity: Thirteen new polyhydroxysterols and two known
polyhydroxysterols isolated from this species displayed considerable cytotoxicity against
a small panel of human solid tumour cell lines. Further, these compounds also showed
weak antibacterial activity against Streptococcus pyogenes 308A, Pseudomonas aerugi
nosa 1771, and Pseudomonas aeruginosa 1771M (Wang et al., 2003a).
Cytotoxicity: Eleven new polyhydroxysterols and eight new saponins isolated from the
brine shrimp active fraction of this species displayed considerable cytotoxicity against a
small panel of human solid tumour cell lines (Wang et al., 2004a).
Antitumour and cytototoxic activities: A total of 14 saponins designated as certonardo-
sides A–N along with culcitoside C6 have been isolated from this species and tested for
cytotoxicity against a small panel of human solid tumour cell lines. Among these saponins,
certonardoside C was most active against the SK-MEL-2 skin cancer cell line (ED50,
3.8 µg/mL). Certonardosides L and N showed considerable cytotoxicity against all five cell
lines, while certonardosides A, H, K, and M were weakly active only against the skin can-
cer cell line. Other congeners did not show any activity against the five human solid tumour
cell lines. Further, certonardosides K–M and certonardoside N were also evaluated for anti-
bacterial activity against 20 clinically isolated strains. Certonardosides K–M showed weak
antibacterial activity against Streptococcus pyogenes 308A, Pseudomonas aeruginosa 1771,
and Pseudomonas aeruginosa 1771M (MIC, 25.0 µg/mL). Certonardoside N displayed
weak antibacterial activity against Pseudomonas aeruginosa 1771 and Pseudomonas aeru
ginosa 1771M (MIC, 25.0 µg/mL). Certonardosides K–M and certonardoside N were inac-
tive (MIC, 25.0 µg/mL) against the rest of the strains (Wang et al., 2003b).
Antitumour activity: Among the two new sulphated saponins, viz. certonardosides P2
and I3 isolated from this species, Compound P2 displayed significant cytotoxicity against
the SK-MEL-2 skin cancer cell (Wang et al., 2005).
Others: Anon. (https://www.genome.jp/db/pcidb/kna_species/23055#metabolite) reported

on the isolation of 24 other bioactive compounds (as given below) in this species. Their
pharmaceutical activities are to be known.
Maier (2008) reported on the presence of certonardosides O1, P1, J2, J3, I2, H2, B2, B3,
H3, and H4 from this species.
Leiaster teres (Verrill, 1871)
Common name(s): Sunstar starfish, purple linckia starfish, purple starfish, smooth sea star
Global distribution: Eastern Pacific
Ecology: It is a coral reef-associated species.
Biology
Description: Individuals of this species are with thick dermis. Abactinal (aboral/dor-
sal) plates and marginal are lacking crystalline bodies. Papular areas are poorly defined.
Cylindrical, non-furrowed adambulacral spines are found arranged in pairs or triplets.

Antimicrobial activity: The ethanol extracts of this species showed antimicrobial activ-
ity against Staphylococcus aureus with the values of zone of inhibition of 10–15 mm in
diameter. Further, these extracts were found active against Mycobacterium tuberculosis
with the values of 82 and 8 (under BACTEC 460 system) at concentrations of 300 and
100 µg/mL, respectively, and Mycobacterium avium with 21 at concentration of 300 µg/mL
(Encarnación et al., 2000).
Linckia laevigata (Linnaeus, 1758)
Common name(s): Blue Linckia, blue star
Global distribution: Tropical Indo-Pacific
Ecology: It is an inhabitant of coral reefs and seagrass beds. These blue stars live subtid-
ally or sometimes intertidally on fine (sand) or hard substrata and move relatively slowly.
Biology
Description: This species is known for its variation in colour. These specimens may be
pure, dark, light blue, aqua, purple, or orange. These sea stars have rounded tips at each
of the arms. Some individuals may bear lighter or darker spots along each of their arms.
Individual specimens are typically firm in texture, possessing the slightly tubular, elon-
gated arms and short, yellowish tube feet. The species may grow up to 30 cm in diameter.
Food and feeding: It is a detritivore; it also feeds on sponges, algae, and Asterina starfish.
Reproduction: It may be able to reproduce asexually.
Parasite: This species is also prone to parasitization by a species of the parasitic gastropod
Thyca crystallina.
Aquarium values: This sea star is fairly popular with marine aquarium hobbyists, where
it requires a proper, slow acclimatization before entering the tank system, and an adequate
food source similar to that found in its natural habitat.

Neuritogenic activity: The steroidal glycosides, viz. linckosides A and B isolated from
this species showed neuritogenic activity (Jha and Zi-rong, 2004).
Neuroprotective activity: The compounds linckosides F–K isolated from this species
have shown neuroprotective activity (Grosso et al., 2014).
Antibacterial activities: The chloroform, methanol, and hexane extracts of this species
showed antibacterial activities after 8 hours of exposure (Layson et al., 2014).
Narcissia canariensis (d’Orbigny, 1839)
Common name(s): It has no common names
Global distribution: Its distribution area extends across Cape Verde, the Gulf of Mexico,
Madeira, the Azores, and the Congo.
Ecology: It lives on rocky and sandy soils, from 5 m down to a depth of 100 m.
Biology
Description: This species has an even orange colour with a yellow tip at the end of each of
its arms. Its subambulacral spines are granuliform and are in two rows of four. In diameter
it reaches a maximum size of 40 cm.
Food and feeding: It feeds on algae.

Cytotoxicity and antitumour activity: Its fraction F13-3 containing glucosylceramide
compounds, viz. ophidiacerebrosides B–D displayed cytotoxic activity over 24 hours
on cancerous cell lines, namely human myeloma cell line, KMS-11, glioblastoma (brain
tumour) and HCT-116 (colorectal adenocarcinoma) with an IC50 of around 20 μM (Farokhi
et al., 2010).
IC50 vaues (µM) of F13-3 fraction for different cancer cell lines after 24 hours of treatment.
IC50 KMS-11 HCT-116 GBM

F13-3 15.2 18 34.6
Source: Farokhi, F. et al., Mar. Drugs, 8, 2988–2998, 2010.

Narcissia gracilis malpeloensis (Downey, 1975)
Global distribution: North Pacific Ocean
Ecology: It occurs near the base of a vertical rock wall at a depth of 49 m.
Biology
Description: In this species, the whole surface of the body is covered by a thick granular
layer, formed by semispherical or polygonal granules, which completely hide the contours
of the lower plates. It is star-shaped with long, thin radii, sharpened regularly and with
the distal ends raised upwards. The body is covered with polyhedral granules of different
sizes. In the centre of the abactinal (upper) surface of the disc, there is a set of primary
plates. The diameter of the specimen is from 6.4 to 8.5 cm.

Antimicrobial activity: The ethanol extracts of this species showed antimicrobial activity
against Staphylococcus aureus and Bacillus subtilis with the values of zone of inhibition
of 10–15 mm in diameter. Further, these extracts were found active against Mycobacterium
avium with the value of 13 (under BACTEC 460 system) at concentration of 300 µg/mL
(Encarnación et al., 2000).
Pharia pyramidata (Gray, 1840)
Common name(s): Yellow-spotted starfish, Bradley’s sea star
Global distribution: Eastern Pacific: Costa Rica, El Salvador, and Panama
Ecology: This is an epibenthic, tropical species found on rocky bottoms in subtidal coral
reef areas at depths of 1–130 m.
Biology
Description: It is a large five-armed sea star, with long tubular arms. It has lavender/brown
base colour covered in yellow-green spots, which leave a clear line down the centre of each
arm. Its maximum size is about 30 cm.
Food and feeding: It feeds mainly on pocilloporid corals.

Cytostatic and cytotoxic effects: The whole body extracts of this species showed strong
cytostatic (growth inhibition) and cytotoxic effects against two human cell lines, lung car-
cinoma A-549 and colon carcinoma HT-29 (Petzelt, 2005).
Phataria unifascialis (Gray, 1840)
Common name(s): Blue sea star, tan starfish, blue phataria starfish
Global distribution: Eastern Pacific Ocean: from the Gulf of California and Magdalena
Bay (Mexico) to northwest Peru, including various eastern Pacific island groups such as the
Galápagos
Ecology: This is an epibenthic, tropical species found on rocky bottoms, particularly in

subtidal areas at depths of 1–50 m. It remains fully active at temperatures down to 17°C
but becomes inactive when it drops to 14°C.
Biology
Description: All the five rays of this species are tapering. There is only one row of ambu-
lacral spines that is wider in the base and tapering. It reaches a diameter of about 30 cm.
Food and feeding: It feeds mainly on pocilloporid corals.

Antimicrobial activity: The ethanol extracts of this species showed antimicrobial activ-
ity against Mycobacterium avium with the values of 17 (under BACTEC 460 system) at
concentration of 300 µg/mL (Encarnación et al., 2000).
Anti-inflammatory activity: This species has been reported to be an ethnomedi-
cine with anti-inflammatory activity (Narchi, https://getd.libs.uga.edu/pdfs/narchi_
nemer_e_201105_phd.pdf).
2.18 FAMILY: OREASTERIDAE (ORDER: VALVATIDA)

Anthenea aspera (Döderlein, 1915)
Common name(s): Cake sea star
Global distribution: Western Pacific; North Australia, Southern Japan, China, Indonesia,
and Singapore
Ecology: This benthic, tropical species is seen in seagrass meadows while larger ones
are usually seen on coral rubble, sometimes wedged under large rocks; the depth range is
18–18 m. It is usually seen alone and usually more active at night.
Biology
Description: It has a stiff body and the upper side is usually slightly convex. Arms are
short with rounded tips. Large, neat marginal plates are seen all around the edges. The
upper side is covered with tiny pedicellariae (pincer-like structures). The underside is flat,
usually with a pattern of bars that form chevrons around the arms, with large bivalved
pedicellariae. The tube feet are short tipped with suckers. These specimens have a wide
variety of patterns and colours, from black, brown, red, orange, yellow to even green.
Diameter of this species with arms is 10–20 cm.
Food and feeding: It feeds on invertebrates that grow on the grass-blades.

Anticancer activity: Polyhydroxysteroidal glycosides, anthenosides A1 and A2, and
anthenoside A isolated from this species showed anticancer activity by slightly inhibiting
the cell viability of human cancer T-47D cells. Further, anthenoside A1 slightly inhibited
colony formation of human cancer RPMI-7951 cells by 16% while compound, anthenoside
A2 decreased the number of colonies of T-47D cells by 40% (Malyarenko et al., 2018a,
2018b).
Haemolytic and immunomodulatory activities: Ten polyhydroxysteroidal glycosides,

anthenosides L-U have been isolated from this species. Of these compounds, the mix-
ture of anthenosides T and U showed haemolytic activity with an EC50 = 8 μM. Further,
the compound anthenoside O at a dose of 10 μM exhibited a potential immunomodu-
latory action, decreasing by 24% the intracellular ROS content in RAW 264.7 murine
macrophages, induced by pro-inflammatory endotoxic lipopolysaccharide from E. coli
(Malyarenko et al., 2016).
Blunt et al. (2017) reported that the steroidal glycosides anthenosides L–U of this species
have shown bioactivities as detailed below.
Anthenoside L: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside M: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside N: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside O: Inhibition of ROS in stimulated macrophages
Anthenoside P: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside Q: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside R: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside S: Cytotoxicity, haemolytic activity and ROS inhibition
Anthenoside T: Haemolytic activity
Anthenoside U: Haemolytic activity
Others: Malyarenko et al. (2018a) reported on the occurrence of anthenosides V–X from
this species.
Anthenea pentagonula (Lamarck, 1816) (= Anthenea chinensis)
Global distribution: Western Pacific: South China Sea, Thailand, and Hong Kong
Ecology: This subtropical, benthic species inhabits shallow waters; the depth range is
0–60 m.
Biology
Description: The back of this species is less convex and obscurely netted. Both the mar-
ginal and distal ossicules of the oral surface are furnished with large, elongated two-lipped
pores. There are five arms that are broad and are in half the length of body width. Arms
are broad and rounded at the tips. A very large, two-lipped pore is present on the ventral
surface of each plate. The ambulacral spines are disposed in three rows. In the innermost
row, there are five spines on each plate. Marginal plates are not tuberculated.

Antitumour and cytotoxic activities: Ma et al. (2010) and Malyarenko et al. (2018a)
reported on the isolation of ten new polyhydroxysteroidal glycosides, anthenosides B−K
(2–11), from this species. Among these compounds, compounds 5, 7, a mixture of 8 and 9,
and a mixture of 10 and 11 showed inhibitory activity against human tumour K-562 and
BEL-7402 cells. Furthermore, the mixture of 10 and 11 also exhibited cytotoxicity against
human tumour U87MG cells and promoted tubulin polymerization.
Ma et al. (2009) also reported on the occurrence of a new polyhydroxysteroidal glycoside,

(20R,24R)-16-O-(4-O-methyl-2-acetamido-2-deoxy-β-d-galactopyranosyl)-24-ethyl-5α-
cholest-8(14)-en-3β,6β,7β,16α-tetrol (anthenoside A), from this species. This compound
exhibited significant cytotoxicity against human tumour K-562, BEL-7402, and U87MG cells.
Anthenea sibogae (Döderlein, 1915)
Common name(s): Not reported
Global distribution: Not available
Ecology: It inhabits at depths of 12.8–162.5 m.
Biology
Food and feeding: Not available

Anticancer activity: Polyhydroxysteroidal glycosides, anthenosides S1–S6 (1–6), along
with a mixture of two previously known related glycosides, 7 and 8, have been isolated
from the methanolic extract of this species. The mixture of 7 and 8 slightly inhibited the
proliferation of human breast cancer T‐47D cells and decreased the colony size in the
colony formation assay (Kicha et al., 2018).
Others: Hung et al. (2018) reported on the occurrence of two steroidal diglycosides, namely
anthenoside R and anthenoside S from the methanol extract of this species.
Culcita novaeguineae (Müller and Troschel, 1842)
Common name(s): Cushion star
Global distribution: This species is found in tropical warm waters in the Indo-Pacific.
Its range extends from Madagascar and the Seychelles to the Philippines, New Guinea,
Australia, and Hawaii.
Ecology: This species occurs on living coral reefs at the reef edge and slope at depths of
0–90 m.
Biology
Description: It resembles a pentagonal pincushion. Arms of this species are very short
and the body is almost globular. The upper side has a texture of circular shapes and little
bumps. The underside is flat with five grooves and short tube feet with sucker-shaped tips.
The individuals have a wide range of colours and patterns. Juveniles are flatter, more star-
shaped with short arms edged with large marginal plates. The diameter of the specimen
with arms is 30–40 cm.
Food and feeding: It feeds on detritus and small invertebrates, including stony corals.
Associated species: A number of organisms live in or on this species. The small shrimp
Periclimenes soror lives as a commensal hiding under the cushion star. Astroxynus culci
tae, Stellicola oreastriphilus, and Stellicola parvulipes are copepods that live parasitically
on the outside of the cushion star. The star pearlfish (Carapus mourlani) sometimes lives
as a commensal inside the cushion star.
Aquarium value: It can be kept in a reef aquarium of sufficient size, stocked with suitable
corals on which it can feed.

Cytotoxicity: Two new sulphated steroidal pentaglycosides (asterosaponins), novaeguino-
sides I and II, along with the known regularoside B, have been isolated from this species.
The new asterosaponins showed marginal in vitro cytotoxicity against two human tumour
cell lines (Tang et al., 2005).
Polyhydroxysteroidal glycosides, culcinosides A–D (1, 2, 4, and 7) along with three known
compounds—echinasteroside C (3), linckoside F (5), and linckoside L3 (6)—isolated from
the ethanol extract of this species exerted cytotoxicity against human glioblastoma cell
lines U87, U251, and SHG44 (Lu et al., 2018).
Cytotoxic Activity of the New Compounds in vitro

Cytotoxic Activity (IC50, µM)
Compound U87 U251 SHG44
1 9.35 11.28 8.04
2 33.52 40.76 36.54
4 26.33 22.66 35.26
7 43.25 28.93 26.22
Source: Lu, Y. et al., Mar. Drugs, 16, 92, 2018.
Sima and Vetvicka (2011) reported that the sulphated glycosides, viz regularosides and
novaeguinosides of this species exhibited marginal cytotoxicty.
The n-BuOH extract of this species resulted in the isolation of one sulphated steroidal gly-
coside (asterosaponin) (1), along with three asterosaponins, viz. thornasteroside A (2), mar-
thasteroside A(1) (3), and regularoside A (4), as active compounds. All these saponins showed
moderate cytotoxicity against cancer cell lines K-562 and BEL-7402 (Tang et al., 2006).
Halityle regularis (Fisher, 1913)
Common name(s): Mosaic cushion star, regular cushion star
Global distribution: Tropical Western Central Pacific: Philippines, New Caledonia, and
Palau; South China Sea
Ecology: This benthic species is fond of sandy or sandy-sandy bottoms between 5 and
30 m deep, but it has also been observed up to 275 m.
Biology
Description: It is a very rounded (very domed) starfish with pentagonal shape. The arms
are very short and are slightly bent upwards at their end. Respiratory papules are grouped
in numerous papular triangular areas, arranged in six hexagonal patterns that can be remi-
niscent of six-petalled flowers. This pattern is repeated very evenly over the entire upper
(aboral) surface of the star and has a lighter colour than the rest of the body the colour of
which varies from orange to brown. The madreporic plaque is usually visible on the dorsal
surface. Pedicellaria are of two types: clip-shaped or coffee bean shaped. On the underside
(oral), the five ambulacral furrows meet at the mouth in the central position. This face
is made up of plates resembling small, rectangular, cream-coloured, well-ordered pads.
Around it, five squares of plates are blue in colour and edged with yellow. The species is
up to 40 cm in diameter and up to 15 cm thick.
Food and feeding: It feeds on sponges.
Reproduction: No information is available on the reproduction of the regular cushion star,
which is rarely encountered and is still poorly known.
Associated species: It is common to observe small Zenopontonia soror shrimps on the sur-
face of this starfish, preferably on the ventral side. Similarly, copepods such as Astronomes
indica and Stellicomes tumidulus are external parasites of this starfish.

Nonsulphated steroidal diglycosides 1–6 (1: Halityloside E; 2: Halityloside D; 3: Halityloside
F; 4: Halityloside B; 5: Halityloside A; 6: Halityloside H); the sulfated steroidal diglyco-
sides 7 and 8 (7: Halityloside H,6-O-sulfate; 8: Halityloside I); and the polyhydroxysterols
10 and 11 (10: (25S)-5α-Cholestane-3β,6α,8,15β,16β,26-hexol; 11: (25S)-5α-Cholestane-
3β,4β,6α,8,15β,16β,26-heptol) have been isolated from this species. Activities of these
compounds are, however, to be known (Iorizzi et al., 1986).
Pentaceraster regulus (Müller and Troschel, 1842)
Common name(s): Spotted sea star
Global distribution: Western Central Pacific: Thailand and New Caledonia; India: Gulf
of Mannar; Lakshadweep, Andaman, and Nicobar Islands
Ecology: This tropical, benthic species is usually found in sandy slopes; the depth ranges
from 2–115 m.
Biology
Description: The hull of this species consists of a disc with arms. The primary plates of
the upper side are elevated that tend to form regular longitudinal series. Pore-areas are
usually well defined. Spines are absent on the first two or four supero-marginal plates in
each interradial angle. Dorsolateral elevations or spines are developed along with the arms.
Body colour is yellow with a reddish spine on the upper side.
Food and feeding: This corallivore feeds on coral using a variety of unique adaptations
and strategies.
Reproduction: In this species, sexes are separate. Fertilization is external and its spawn-
ing frequency is one clear seasonal peak per year.

Immunomodulatory antioxidant activity: Three new polyhydroxysteroid glycosides,
viz. regulusosides A, B, and C isolated from this species have shown immunomodulatory
antioxidant activity by increasing reactive oxygen species (ROS) production in macro-
phages (Blunt et al., 2017).
Antimicrobial activity: The methanolic and n-butanol extracts of this species have shown
antimicrobial activity against bacterial pathogen Klebsiella oxytoca (inhibition zone dia.,
10 mm) and fungal pathogen Trichophyton rubrum (inhibition zone dia., 8 mm) (Kumaran
et al., 2011).
Cytotoxic and immunomodulatory activities: Seven new asterosaponins, pentaregulo-
sides A–G (1–7), including two furostane-type steroid oligoglycosides (2, 3), along with
four previously known compounds (8–11) have been isolated from the ethanolic extract
of this species. Compound 1 exhibited cytotoxic activity with an IC50 value of 6.4 μM
against RAW 264.7 murine macrophages. In contrast, nontoxic asterosaponins 3, 4, and
5 showed a potential immunomodulatory action at a concentration of 5 μM, reducing by
40%, 28%, and 55%, respectively, reactive oxygen species formation in the RAW 264.7
cells, co-stimulated with the pro-inflammatory endotoxic lipopolysaccharide from E. coli
(Kicha et al., 2017).
Others: Two new sulphated steroidal glycosides, viz. regulusoside S1 and regulusoside S2
have been isolated from the EtOH extract of this species (Kicha et al., 2017).
Protoreaster lincki (Blainville, 1834)
Common name(s): Red knob sea star, red spine star, African sea star, African red knob sea
star
Global distribution: Indo-Pacific; Western Indo-Pacific
Ecology: Its habitat ranges from shallow tidal pools to reefs up to 100 m deep.
Biology
Description: It has numerous tubercles located along its five arms. These tubercles are
bright red and extend upward from the arms. It has a grey body with red stripes that con-
nect the tubercles. It grows to a maximum diameter of 30 cm.
Food and feeding: It eats soft corals, sponges, tube worms, clams, and other starfish. It is
active in the daytime.
Aquarium values: It is a popular aquarium species but is considered incompatible with

many other invertebrates.

Cytotoxic and immunomodulatory activities: Four new steroidal glycosides, protolinck-
iosides A–D (1–4) and four previously known glycosides, 5–8, have been isolated from the
MeOH/EtOH extracts of this species. Among these compounds, compounds 1 and 5 at a
concentration of 100 lM showed cytotoxic activity and reduced cell viability by 20% and
50%, respectively. Compounds 2 and 4 induced ROS formation about 30% in the RAW
264.7 cells at a dose of 1 lM compared with untreated cells. Further, compounds 1, 2, 3,
and 4 significantly decreased the ROS content in the RAW 264.7 cells at a concentration
of 10 lM by around 50%, 48%, 44%, and 35%, respectively, suggesting their potent anti-
inflammatory properties (Malyarenkoa et al., 2016; Blunt et al., 2017).
Antimicrobial, haemolytic, antinociceptive, and cytotoxic activities: The crude compound

methanolic extract of this species has shown antimicrobial, haemolytic, antinociceptive,
and cytotoxic activities (Sumithaa et al., 2017a).
Antibacterial activity: Extract of this species displayed antibacterial activity with maxi-
mum inhibition zone of 9.5 mm against Bacillus subtilis (MIC of 75 µg/mL) and minimum
inhibition zone of 6.5 mm against Klebsiella pneumoniae, Salmonella typhimurium, and
Shigella sonnei (MIC value of 100 µg/mL) (Mariya and Ravindran, 2015).
Antimicrobial activity: The n-butanol extract of this species showed antimicrobial activ-
ity against the human bacterial and fungal pathogens S. paratyphi and K. pneumoniae
(inhibition zone dia., 6 mm); and fungal pathogen A. flavus (12 mm) (Kumaran et al., 2011).
Protoreaster nodosus (Linnaeus, 1758) (= Oreaster nodusus)
Common name(s): Horned sea star, chocolate chip starfish, Knobbly sea star
Global distribution: Indo-Pacific region: Cargados Carajos, Eastern Africa, Madagascar,

Kenya, New Caledonia, Palau Islands, Seychelles, Tanzania; West Indian Ocean; Andaman
and Nicobar Islands
Ecology: It primarily inhabits seagrass meadows in warm, shallow, intertidal waters.

Adults are usually seen in coral rubble areas.
Biology
Description: The hard, heavy body of this species is calcified. The disc is markedly ele-
vated. Arms are long, tapering to a rounded tip, thick and triangular in cross-section.
Although their arms appear stiff, these can bend quite extensively. This species is easily
identified by its single row of knobs along the upper side of the arms. The shape, colour,
and number of knobs may vary. Underneath, from grooves under the arms, emerge tube
feet with sucker-shaped tips. These tube feet can be bright red or purple. These sea stars
are mostly red, orange, or brown, but sometimes white, pink, blue or green ones are also
encountered. Diameter of this species with arms is about 30 cm.
Food and feeding: These organisms are opportunistic carnivores. Adults are known to
prey on most sessile life forms including hard corals and sponges in aquariums. In this
same setting, they will hunt down snails and eat them. An individual of horned sea star
also has been observed eating a sea urchin in its natural habitat.
Associated species: As with other tropical echinoderms, commensal animals like shrimps
(of genus Periclimenes), tiny brittle stars and even juvenile filefish can be found on the
surfaces of a horned sea star. This may be attributed to its protective nature, as there are
few predators that would dare to eat this echinoderm.
Commercial/aquarium uses: They are sometimes seen in the marine aquarium trade. In
many tropical Asian and Pacific countries, horned sea stars are collected for the seashell
trade for their ornate skeletons.

Anti-inflammatory activity: The methanolic extract and dichloromethane fraction of
this species showed anti-inflammatory activity by exerting potent inhibitory effects
on the production of all three pro-inflammatory cytokines, with IC50 values ranging
from 0.60 to 26.19 μg/mL. Four steroid derivatives (1–4) isolated from the dichloro-
methane fraction and water layer of this species also showed potent inhibitory activi-
ties. That is, (25S) 5α-cholestane-3β,4β,6α,7α,8β,15α,16β,26-octol (3) inhibited the
production of IL-12 p40 and IL-6 (IC50s = 3.11 and 1.35 μM); (25S) 5α-cholestane-
3β,6α,8β,15α,16β,26-hexol (1) and (25S) 5α-cholestane-3β,6α,7α,8β,15α,16β,26-
heptol (2) inhibited the production of IL-12 p40 (IC50s = 0.01 and 1.02 μM); and the
compound nodososide (4) exhibited moderate inhibitory effects on IL-12 p40 and IL-6
production (Thao et al., 2015).
Others: Ten new compounds, viz PNC-1-3a, PNC-1-3b, PNC-1- 4a/PNC-1-4b, PNC-1-4c,
PNC-1-5b, PNC-1-5c, PNC-1-6a, PNC-1- 6b/PNC-1-6c, PNC-1-6d, PNC-1-7a, PNC-1-7b,
PNC-1-8a, PNC-1- 8c, and PNC-1-10; and three new ganglioside molecular species, viz.
PNG-1, PNG2A, and PNG-2B have been isolated from the pyloric caeca of this species.
Their bioactivities are, however, yet to be known (Gomes et al., 2014).
2.19 FAMILY: ASTERINIDAE (ORDER: VALVATIDA)

Tremaster mirabilis (Verrill, 1880) (= Tremaster novaecaledonia)
Global distribution: Geographical distribution: western Atlantic and adjacent waters from
Nova Scotia, south of Cuba, east and south of the Falkland Islands, and in the northeast off
central Norway, also in Arctic from Labrador, Southeast Greenland, Southeast Iceland and
the Barents Sea, and in the Southern Ocean from Kerguelen
Ecology: This benthic, polar species has a depth distribution of 150–1060 m.
Biology
Description: Like other starfish, it has eyes at the end of each arm, which it bends to direct
its gaze. The furrow spines are in vertical series on adambulacral plates. There are four
series of tube feet.
Food and feeding: It feeds on food fragments and algal or bacterial mat covering the
substratum.

HIV-inhibitory activity: The two sulphated sterols isolated from this species exhibited
anti-HIV-1 and HIV-2 activity with IC50 values of >161 µM; and >125 μM, respectively
2.20 FAMILY: SOLASTERIDAE (ORDER: VALVATIDA)

Solaster endeca (Linnaeus, 1771)
Common name(s): Purple sun star, northern sun star, smooth sun star
Global distribution: Circumboreal: Arctic Ocean, Northern Pacific, and Atlantic

Ecology: The typical habitats of this temperate, benthic species are muddy sand, gravel, or
rocky areas with deposited sediment. It is found in both sheltered and fairly exposed loca-
tions; the depth range is 0–475 m.
Biology
Description: The body of this species has a large disc and 9–10 (rarely 7–13) long and
tapering arms. The upper surface is crowded with small paxillae with short spines. The
marginal paxillae, which are not very conspicuous, are arranged in a double row in which
the upper ones are smaller than the lower ones. In the oral side, the adambulacral plates
have 2–3 small furrow spines (rarely 4), which are almost concealed in the furrow. There
are transverse series of 6–8 short spines on the outer surface of the adambulacrals. The
species is up to 400 mm in diameter.
Food and feeding: A predator, it feeds on other starfish and bivalve molluscs, sea cucum-
bers, and other invertebrates.
Reproduction: In Britain, breeding of this species takes place in the spring. Over a period
of about a week, the female lays several thousand eggs in batches. These rise to the surface
of the water where they are fertilized by sperm liberated by the male. The developing lar-
vae become less buoyant after three days, feed on the yolks of their eggs, swim with cilia,
and develop a pair of larval arms. After about 18 days, they sink to the seabed, where each
one attaches itself to the substrate with a sucker. Here, it undergoes metamorphosis during
which it develops a disc and first five and then more arms, a pair of tube feet, relatively
long spines, red eyespots on the tips of the arms, a mouth, and an anus. After six weeks,
the sucker is resorbed and the juvenile starfish begins to move about with its tube feet.

Cytotoxic activity: Four polyhydroxylated steroids, viz. (20R)-5α-cholestan-3β,6α,8,
15α,24,26-hexaol, (20R,25S)-5α-cholestan-3β6α,8,15β,16β,26-hexaol, (20R,25S)-5α-cholestan-
3β,6α,15β,16β,26-pentaol, and marthasterone sulfate have been isolated from this species.
These steroids were found to be weakly cytotoxic in a human HeLa cell culture, and some
of them were inhibitors of nonspecific esterase from mouse Ehrlich carcinoma (Levina
et al., 2010).
Pharmaceutical Brittle
Stars and Basket Stars
(Class: Ophiuroidea)
3.1 FAMILY: OPHIOLEPIDIDAE (CLASS: OPHIUROIDEA;

ORDER: AMPHILEPIDIDA)
Ophiolepis superba (H. L. Clark, 1915)
Common name(s): Banded brittle star, superb brittle star, painted serpent starfish
Global distribution: Indo-West Pacific: Western Indian Ocean to New Caledonia and the
Philippines
Ecology: This tropical, benthic species inhabits protected reef areas; it occurs under coral
rubble in shallow reef areas; depth range is 0–30 m.
Biology
Description: It is a small ophiure with a pentagonal disc 3 cm in diameter. It is slightly
convex and with five relatively short arms of 10–12 cm length. The plates on the basal disc
are large and conspicuous. At the anchorage of the arms, two parallel plates larger than the
others are present. The arms consist of articulated segments that are also clearly visible.
Each segment carries on each side 1–5 small discrete spines. The background colour of
the specimen is beige with some purple to brown rings on the arms. The ventral side of the
disc and arms is beige. A purple pattern is also present on the ventral side of the disc. The
maximum length of the specimen is 20 cm.
171
Food and feeding: It is a detritory species that comes out at night in search of food; it also
feeds on microalgae (e.g., spirulina) and small crustaceans (krill, mysis, artemia).
Reproduction: It is a gonochoric species (the sexes are separated). Fertilization is external.
The eggs and then the larvae are pelagic. Like other ophiuroids, it has great regenerative
power. Its arms are able to push back. Similarly, in case of danger, stress, or sudden change
in water quality, this species may voluntarily lose some or all of one or more arms. This is
called autotomy.

HIV-inhibitory activity: The sulphated sterols isolated from this species exhibited HIV-1 and
HIV-2 inhibitory activity with IC50 values ranging from >157 to 161 µM (McKee et al., 1994).
3.2 FAMILY: OPHIOPHOLIDAE (ORDER: AMPHILEPIDIDA)

Ophiopholis aculeata (Linnaeus, 1767)
Common name(s): Crevice brittle star, daisy brittle star

Global distribution: It has a circumpolar distribution and is found in the Arctic Ocean,
the northern Atlantic Ocean, and the northern Pacific.
Biology and Ecology of Pharmaceutical Brittle Stars and Basket Stars (Class: Ophiuroidea) 173
Ecology: It is generally found on rocky substrates where it has a tendency to hide inside
shells, in hollows, and crevices. It generally occurs at depths less than 300 m but has been
found as deep as 1880 m.
Biology
Description: It has a central pentagonal disc of about 2 cm in diameter, and this is clearly
set off from the five robust, twisted arms, which are four times as long as the disc is wide.
The mouth is on the oral or underside of the disc and is surrounded by five toothed jaws.
The aboral or upper side of the disc is granular, and a pair of large scales, the radial shield,
covers the base of each arm. The radial shield is covered with small spines. The many-
jointed arms are covered with more scales both on the oral and aboral surfaces; and on the
aboral surface, there is a regular series of oval scales, each surrounded by smaller scales.
There are six or seven stout spines (with one short, hooked spine) per segment. The colour
of the individuals varies; it is often reddish or variegated and sometimes purplish. The
central scales on the disc often form a ten-pointed star, and there are often darker bands
on the arms.
Food and feeding: It feeds on detritus and small organisms that it traps with its tube feet
and with mucus secreted by glands on its arms.
Predator: It is preyed on by fish and birds.
Reproduction: The sexes are separate in this species, and fertilization is external. Mass
spawning events have been seen, with all the individuals of this species in a locality releas-
ing their spawn at the same time in response to some environmental cue. Several spawning
events have occurred when warmer surface waters have down-welled into deep, colder
water layers. The larvae of brittle stars are known as ophiopluteus larvae and form part of
the plankton. When they are fully developed, they settle on the seabed and undergo meta-
morphosis into juvenile brittle stars.

Cardiovascular activity: Its disulfated polyhydroxysteroids, which are potent Ca2+ ago-
nists in mammalian cell systems, have vasoconstricting and positive inotropic properties.
(Positive inotropes strengthen the force of the heartbeat.) (Petzelt, 2005)
Ophiopholis mirabilis (Duncan, 1879)


Global distribution: Cold temperate areas
Nothing is known about its biology and ecology

Antioxidant and antimicrobial activity: The compound ophiurasaponin extracted from this
species showed antioxidant and antimicrobial activity. The values of the IC50 of hydroxyl-
radicals, superoxide anions, and peroxide were 25.54, 9.98, and 1.37 mg/mL, respectively.
The refined ophiurasaponin also showed good inhibitory effect on Escherichia coli, Bacillus
subtilis, Staphylococcus aureus, Aerobacter aerogenes, and Proteusbacillus vulgaris, and
the minimum inhibitory concentration (MIC) was 0.0443 mg/mL (Wang et al., 2014).
Capacities of Antioxidation of Ophiurasaponin

Superoxide Anion Hydroxyl Hydrogen Peroxide
Concentration Concentration Concentration
(mg/mL) Inhibition (%) (mg/mL) Inhibition (%) (mg/mL) Inhibition (%)
22 98.3 30 77.3 5 94.0
4.4 32.1 27 59.6 3 72.1
2.2 19.8 25 47.5 1 49.2
1.5 16.3 23 34.4 0.5 38.6
1.1 10.2 20 10.1 0.05 31.9
IC50 (mg/mL) 9.98 25.54 1.37
Source: Wang, R. et al., J. Chem., 5, 2014.
Inhibition of Ophiurasaponin from Ophiopholis Mirabilis to Bacteria

Ophiurasaponin Ophiurasaponin
Species Crude (mm) MIC (mg/mL) Refined (mm) MIC (mg/mL)
Bacillus subtilis 12.2 1.1 26.9 0.0443
Escherichia coli 11.5 1.1 32.5 0.0443
Staphylococcus aureus 11.2 0.11 30.9 0.0443
Proteusbacillus vulgaris 11.2 — 29.6 —
Aerobacter aerogenes 16.0 1.1 28.2 0.0443

—: no inhibitory effect
Inhibition of Ophiurasaponin from Ophiopholis Mirabilis on Fungi

Ophiurasaponin Ophiurasaponin
Species Crude (mm) MIC (mg/mL) Refined (mm) MIC (mg/mL)
Mucor circinelloides 11.5 1.1 13.0 4.4
Saccharomyces 11.2 1.1 — —
cerevisiae

—: no inhibitory effect
3.3 FAMILY: OPHIOTRICHIDAE (ORDER: AMPHILEPIDIDA)

Ophiocnemis marmorata (Lamarck, 1816)
Common name(s): Marbled brittle star

Global distribution: Indo-West Pacific: Thailand, China, and Hong Kong
Ecology: This subtropical, demersal species is quite common in soft bottom habitats of
shallow waters with a depth range of 0–100 m.
Biology
Description: The disc of this species has dark and light green markings. Arms are banded
dorsally with green markings on a white background and ventrally pale coloured. It is a
six-armed species. Aboral disc scales are distinct, with scattered spinelets mainly on the
margin of the disc. Radial shields are large, contiguous at the distal end. The dorsal arm
plate is elliptical with more or less distinct small medium distal lobe mostly with a pair of
dark spots on each side, giving a trilobed appearance to the plate. The ventral disc is cov-
ered with scales, and the ventral arm plate is almost octagonal. Oral shields are large and
oval. Adoral shields are larger and are not meeting proximally.
Food and feeding: It is a filter-feeding ophiuroid species.
Associated species: This species is often found in association with the rhizostome medusa
Rhopilema hispidum. This brittle star possibly seeks for food supply, shelter, and protec-
tion through this association (Kanagaraj et al., 2008).

Antimicrobial and haemolytic activities: The ethanol extract of this species contain-
ing steroidal compounds showed both antimicrobial and haemolytic activities. Maximum
zone of inhibition was recorded against Staphylococcus aureus (7.0 mm), followed by
5.0 mm inhibition against Escherichia coli and 4 mm against Vibrio parahaemolyticus and
Staphylococcus typhi. Further, the haemolytic activity was high in goat blood (128 HU) in
methanolic extracts (Prabhu and Bragadeeswaran, 2013b).
Ophiothrix fragilis (Abildgaard in O.F. Müller, 1789)
Common name(s): Common brittle star, hairy brittle star

Global distribution: This species is found in Britain and Ireland and in the North Sea. It
ranges southwards from the Lofoten Islands and Iceland to the Mediterranean Sea and the
Azores, and along the west coast of Africa south to the Cape of Good Hope.
Ecology: It is a coastal species and is most common on tide-swept rock and on coarse
sediments, preferring hard substrates including sand and shingle. It is often found in empty
shells or under stones. Depth range is from the littoral zone down to 350 m.
Biology
Description: The central disc is about 1 cm in diameter with the five arms, which are about five
times as long. The disc is clothed in five rays of spines radiating from a spiny centre. Between
these are five pairs of triangular plates; each pair forms a heart-shaped pattern. The slender
tapering arms are quite distinct from the disc and are covered with overlapping scales. The
dorsal arm plates are naked with a longitudinal keel. Each arm segment bears seven glassy,
toothed spines. The arms are extremely fragile and are easily shed either whole or in pieces.
This species is extremely variable in colouration, ranging from violet, purple, or red to yellow-
ish or pale grey, often spotted with red. The arms are usually white or grey with pink bands.
Food and feeding: The common brittle star is a scavenger, feeding on dead organisms.
It is also a suspension feeder, raising an arm and extending the tube feet in order to catch
particles floating by.
Reproduction: This brittle star usually spawns between May and January. Each individual
is thought to spawn just once during each breeding season. A week after spawning, the
planktonic larvae appear in the water column. They metamorphose into young brittle stars,
which drift in the plankton for about three weeks before settling. The smallest brittle stars
found have just two segments per arm and a disc diameter of 2 mm.

Antibiotic activity: The extracts isolated from the unidentified surface associated bacteria
of this species showed antibiotic activity (Marmann et al., 2014).
HIV-inhibitory activity: The sulphated sterol isolated from this species exhibited HIV-1
and HIV-2 inhibitory activity with IC50 values of >322 and >161 µM, respectively (McKee
et al., 1994).
3.4 FAMILY: GORGONOCEPHALIDAE (ORDER: EURYALIDA)

Astrotoma agassizii (Lyman, 1875)
Common name(s): Predatory snake star, Antarctic brittle star
Global distribution: Ross Sea at the Balleny Islands, Victoria Land and Terra Nova Bay,
Antarctica
Ecology: It lives at continental shelf depths from 90 to 700 m, also on continental slope
down below 2000 m.
Biology
Description: This disc diameter of this species can be up to 6 cm across, and its arms can
reach an impressive 70 cm long. Although beautiful, these long, wiggly arms are ringed
with rows of minute hooks, which enable them to grip on to the corals to gain height up off
the seafloor and loop around and grab prey items. It has a life span of about 90 years.
Food and feeding: It preys mainly on small crustaceans (copepods, mysids, and euphau-
sids) and arrow worms (chaetognaths), but it has been found to eat a variety of other ani-
mals from the plankton too.

Antiviral activity: Three sulphated sterols (1–3) found in the Antarctic brittle star Astrotoma
agassizii (Roccatagliata et al., 1998) were tested against three pathogenic viruses of humans,
HSV-2, JV, 341 and PV-3. Compounds 2 and 3 exhibited marked HSV-2 inhibitory activity
(IC50 values of 18.4 and 24.3 µg/mL), while compound 1 was active at the highest tested
concentration (80 µg/mL) with an IC50 of 51.5 µg/mL. However, only compound 3 pre-
sented inhibitory activity (IC50 of 67.4 µg/mL) toward JV virus, while compound 2 was
only active against PV-3 at 80 µg/mL (IC50 of 58.5 µg/mL) (Comin et al., 1999).
Others: Sulfated steroidal polyols, viz. (20R)-cholesta-5,24-diene-2β,3α,21-triol

2,21-disulfate, (20R)-5α-cholest-24-ene-2β,3α,21-triol 3,21-disulfate, and (20R)-cholesta-
5,24-diene-2α,3α,4β,21-tetrol 3,21-disulfate have been isolated from this species
(Roccatagliata et al., 1998).
Gorgonocephalus chilensis (Philippi, 1858) (= Astrophyton chilense)
Common name(s): Antarctic basket star, Chilean basket star

Global distribution: Temperate to polar climates; Southern Ocean
Ecology: This benthic species is found on exposed sites, rocks, and other organisms such
as hydrocorals, gorgonians, and mussels; depth range is 4–900 m.
Biology
Description: The disc of this species is moderately deep, but it is narrowly excavated inter-
radially with a thick ring of peripheral plats. Radial shields are very conspicuous, nar-
row, parallel-sided, and are tapering only near the disc centre. Arms are branched at least
10 times. The dorsal arm surface is covered with domed or conical tubercles. Underarm
surfaces are with small and evenly distributed granules. Colour of the disc is pale brown;
radial shields, tubercles, and arms are cream coloured.
Food and feeding: It feeds on sponges.

Predator: Some bony fishes feed on this species.

Disulfated steroids and a mixture of monosulfated steroids have been isolated from the
ethanolic extract of this species (Maier et al., 2000).
3.5 FAMILY: OPHIOCOMIDAE (ORDER: OPHIACANTHIDA)

Breviturma dentata (Müller and Troschel, 1842)

Global distribution: It has a tropical, Indo-West Pacific distribution (with the exception of
the Red Sea and northwestern Indian Ocean).
Ecology: This species frequents the sub-littoral zone, under boulders or associated with
coral and coral debris on a sand or rubble substratum.
Biology
Description: In this species, oral shields are round, as long as wide, with a marbled pattern.
Dorsal arm plates are beige to brown, with a whitish-grey patch surrounded by a dark brown
border on the median distal side. Lateral arm plates are lighter with several spots. Ventral arm
plates are light with the same spots. Sometimes a dark-coloured patch is present centrally. Arm
spines are white to beige and are broadly and irregularly banded once or twice with light brown.
Upper arm spines are thick, blunt, somewhat flattened, and slightly shorter than the lower ones.
There are two tentacle scales. Colour of the individuals may be brown, white, and beige. Small,
dark brown spots are present both dorsally and ventrally. Disc diameter is 28 mm.
HIV-Inhibitory activity: The sulfated sterol isolated from this species exhibited HIV-1
et al., 1994).
Ophiocoma erinaceus (Müller and Troschel, 1842)
Common name(s): Spiny brittle star, Persian Gulf brittle star

Global distribution: Entire Indo-Pacific except for Pakistan and Western India
Ecology: This tropical species inhabits coral reefs, subtidal areas and in rubble areas;
Biology
Description: This species is black during the day and displays grey blotches at night. Disc
diameter is 2.0–2.5 cm. with red tube feet and two tentacle scales. Individuals attain an
overall diameter of 20 cm.
Food and feeding: It feeds on detritus.
Anticancer/cytotoxic activity: The crude saponin extracted from this species showed
cytotoxicity against Hela-S3 cervical carcinoma with an IC50 value of 23.4 µg/mL. In
addition, the crude saponin increased sub-G1 peak in flow cytometry histogram of treated
cells, ROS generation, and caspase-3 and -9 activity (IC50 of 11.10, 11.27 µg/mL) (Amini
et al., 2017).
Anticancer potential: The polysaccharide of this species has been reported to suppress the
proliferation of HeLa cells in a dose and time-dependent manner. Additionally, polysaccharide
extracted induced intrinsic apoptosis via upregulation of caspase-3, caspase-9, and Bax
along with down-regulation of Bcl-2 in HeLa cells (Baharara and Amini, 2015).
Haemolytic and cytotoxic activity: Sumithaa et al. (2017a) reported that the saponins
ethanolic fractions of this species exhibited haemolytic and cytotoxic activity.
Cytotoxicity: The dichloromethane extract of this species revealed significant cytotoxic
effect on B16F10 melanoma cells with IC50 = 31 µg/mL, which is stronger than the inhibi-
tory effect of methanol extract on melanoma cell growth. In addition, the dichloromethane
extract recruited apoptotic pathway in the response of 31 µg/mL treatment (Baharara et al.,
2015a).
The saponin compounds present in the 80% ethanol extract of this species showed cytotoxic
activity. Haemolytic assay revealed HD50 value was 500 µg/mL. MTT assay exhibited
that saponin extracted in IC50 value of 25 µg/mL induced potent cytotoxic activity against
HeLa cells in 24 hours and 12.5 µg/mL in 48 hours (Amini et al., 2014).
The dichloromethane extract of this species had a cytotoxic effect on lymphoma cell line.
While lower concentrations of 31 µg/mL inhibited cell proliferation, higher concentrations
of 31 µg/mL caused lysis of cells and the IC50 concentration was 31 µg/mL. The caspase
colorimetric assay showed that apoptosis induced by these extracts was caspase-dependent
via intrinsic pathway on lymphoma cell line (Afzali et al., 2015).
Antiproliferative and antiangiogenic activities: Extracts from this species have been
reported to possess antiproliferative activity against A2780cp cells, and antiangiogenic
activity in the fertilized Ross egg model44 (Blagodatski et al., 2017).
Antineoplastic efficacy: Wound healing migration assay method showed that extract of
this species has antineoplastic efficacy by inhibiting cell migration. Caspase assay and
RT-PCR analysis revealed that its methanol extract induced caspase-dependent apoptosis
in HeLa cells through upregulation of caspase-3 followed by upregulation of Bax gene
(Baharara et al., 2016).
Antioxidant and anti-inflammatory properties: The saponins, phenol, and fla-
vonoids compounds present in the extract of this species have shown DPPH
(1,1-diphenyl-2-picrylhydrazyl) and ABTS (azino-bis-3-ethylbenzothiazoline-6-sulfonic
acid) antioxidant activity in a dose-dependent manner. Further, the above extract also
inhibited TGF-β expression, which indicate anti-inflammatory properties of this species
(Baharara and Amini, 2015b).
Hepatoprotective activity: The organic extract of this species has been reported to exert
protective effect of the star extract on liver damages induced by carbon tetrachloride in
adult male Wistar rats. Treatment with the extract increased liver weight, reduced body
weight, and significantly altered other induced changes by carbon tetrachloride on liver
structure such as hepatocytes number, Kupffer cells, and arteritis, which indicated the
improvement of damaged liver tissue (Soheili et al., 2015).
Anti-vasculogenic activity: The polysaccharide extracted from this species was able to
decrease the viability of the human umbilical vein endothelial cells to suppress angiogenesis
and possibly act as a natural antiangiogenic and antimetastatic marine organic compound
against angiogenesis-related pathologies (Baharara et al., 2017).
Ophiocoma schoenleinii (Müller and Troschel, 1842)
Common name(s): Schoenlein’s brittle star

Global distribution: Tropical, east Indo-west-central Pacific Ocean: Gilbert Islands; Bay of
Bengal, East Indies, north Australia, China, south Japan and South Pacific Island; Australia
Ecology: This benthic species is found in coral reefs, mangroves, and on rocky substrates
in inshore, littoral waters; depth range is 0–20 m.
Biology
Food and feeding: Not available
Antibacterial activity: The methanol, chloroform, and hexane extracts of this species
showed antibacterial activity against E. coli, and the values of minimum inhibitory
concentration (MIC, µg/mL) and minimum bactericidal concentration (MBC, µg/mL)
recorded after the exposure of 8 hours were found to be 42, 46, and 20 µg/mL; and 3, 1305
and <1 µg/mL, respectively (Layson et al., 2014).
Ophiocoma scolopendrina (Lamarck, 1816)
Common name(s): Lagoon brittle star, millipede brittle star

Global distribution: Indo-Pacific: Red Sea to Hawaii and Japan
Ecology: This tropical species inhabits reef areas in crevices and under rocks in calcareous
algae areas; depth range is 0–13 m.
Biology
Description: It has long, thin arms that emanate from a small disc-shaped body. Dorsal
disc and dorsal arm plates vary from black, variegated black to pale brown. Arms are
irregularly banded. The individuals can reach a length of 15 cm, while the disc diameter
reaches 20 mm.
Food and feeding: It may also be a deposit feeder or suspension feeder feeding on detritus
and neuston.
Aquarium values: It is suitable for reef or fish-only aquariums.

Anticancer activity: The extract of this species containing the compound curacin E inhibited
the proliferation of mouse leukaemia P388 cells (Ueoka et al., 2016).
Cytotoxic activity: The tetrameric phenylpropanoids, viz. ophiodilactones A, B isolated

from this species showed cytotoxic activity against P388 murine leukemic acells (Atta-ur-
Rahman, 2016).
Ophiomastix brocki (O’Hara et al., 2018) (= Ophiomastix elegans)

Global distribution: It has a worldwide distribution.
Ecology: It inhabits coral reefs.
Biology
Description: In this species, the entire disc is covered by large projecting as well as scarce
and very irregular thorns or short spines. The oral shields are approximately two times as
long as broad. Upper arm plates are fan-shaped with rounded-off aboral edge, two times
broader than long. Lower arm plates are broadly hexagonal. It is dark grey-brown on the
disc with the arms white, and spines are lightly ringed with grey-brown. While its disc
diameter is 16 mm, arms have a length of about 80 mm.

HIV-inhibitory activity: The sulfated sterol isolated from this species exhibited HIV-1
et al., 1994).
Ophiomastix mixta (Lütken, 1869)

Global distribution: Western Central Pacific
Ecology: This benthic, tropical species is found from reef flat to fore reef. It is typically
found under rocks; the depth range is 0–54 m.
Biology
Description: It is a well-defined and strikingly pigmented species. In this species, upper arm
spines are well developed and are often modified in a clavifurcate, swollen, and/or elongated
condition. The disc is covered by few to many spinules, which are cone shaped. The colour of
the specimen is reddish with dark pigmentation. The species has a disc diameter of 15–28 mm.
Cytotoxicity: The butenolide and an acyclic polyhalogenated monoterpene isolated from
this species have shown mild to significant cytotoxicity against a panel of five human solid
tumour cell lines (Lee et al., 2007).
3.6 FAMILY: OPHIOMYXIDAE (ORDER: OPHIACANTHIDA)

Ophiarachna incrassata (Lamarck, 1816)
Common name(s): Green brittle star

Global distribution: Tropical Indo-West Pacific: East Africa to New Guinea and the Philippines
Ecology: It is a benthic dweller, found mainly on reefs and reef crests, usually under large
submerged coral rubble during the day and feeding at night. Depth range is 1–20 m.
Biology
Description: It is a large brittle star characterized by its green-patterned central disc, general
size, and striped spines found on all legs. Maximum length of the specimen is 5.5 cm.
Food and feeding: It is a carnivore feeding on small invertebrates.
HIV-inhibitory activity: The sulfated sterol isolated from this species exhibited HIV-1
and HIV-2 inhibitory activity with IC50 values of >313 and >157 µM, respectively
3.7 FAMILY: OPHIODERMATIDAE (ORDER: OPHIACANTHIDA)

Ophioderma longicauda (Bruzelius, 1805)
Common name(s): Smooth Brittle Star, snake brittle star

Global distribution: Tropical and subtropical; Eastern Atlantic and the Mediterranean
Ecology: This benthic species has a depth range of 0–70 m. It is shade tolerant and photo-
phobic; therefore, it hides below rocks by day.
Biology
Description: It is a large ophiure whose central disc measures 3 cm in diameter. Its five arms
reach four to five times the diameter of the disc. The disc is more pentagonal than circular
and is slightly convex, and it has a leathery texture. Its surface is finely granular to the touch.
The colour of the individuals is usually chocolate brown but may also be red-orange, dark
brown, or black. The arms often have lighter, greenish bands. Ventral side is clearer. There is
no anus in this species, and the undigested food leaves the animal through the mouth.
Food and feeding: It is an active and very voracious nocturnal predator. It feeds on inver-
tebrates buried in the sediment, such as worms and bivalves.
Reproduction: The sexes are separated and reproduction is sexual in this species. Fertilization
is external. The pelagic larvae, which are called ophioplutei, settle after several weeks. This
species also has great regenerative power and easily rebuilds an amputated arm.

Cytotoxic activity: Two sulphated polyhydroxysteroids—cholest-5-ene-2α,3α,4β,
21-tetraol 3, 21-disulphate (1) and cholest-5-ene-2β,3α, 21-triol 2, 21-disulphate (2)—and
a tetrol disulphated polyhydroxysteroid (3) have been isolated form this species. Among
these compounds, compounds 1 and 2 displayed potent cytotoxic activity and cytoprotec-
tive activity against HIV-1, respectively (D’auria et al., 1993b).
HIV-inhibitory activity: The two sulfated sterols isolated from this species exhibited HIV-1
and HIV-2 inhibitory activity with IC50 values ranging from >152–312 to 152–156 µM,
respectively (McKee et al., 1994).
Antibacterial activity: Two disulfated sterols isolated from this species showed inhibitory
activity against S. aureus at 20 µg/disc (Andersson et al., 1989).
Others: Two steroidal glycoside sulfates longicaudoside A and longicaudoside B have been
isolated from this species (Riccio et al., 1986b).
3.8 FAMILY: HEMIEURYALIDAE (ORDER: OPHIURIDA)

Ophioplocus japonicus (H.L. Clark, 1911)
Common name(s): Japanese smooth brittle star

Global distribution: Northwest Pacific: Hong Kong
Ecology: This benthic, subtropical species is found under stones in the intertidal zone;
Biology
Description: The whole body of this species is dark green-grey, and there are somewhat
irregular stripes in the arms.
Cytotoxicity and antibacterial activity: Ten terpenes (1: Pacifenol; 2: Johnstonol; 3:
Prepacifenol epoxide; 4: 2,10-dichloro-3-chlorochamigran-7-en-9-ol; 5: Laurinterol; 6:
10-acetoxy-18-hydroxy-2,7-dolabelladiene; 7: Dihydroxycrenulide; 8: Dictyolactone; 9:
Pachydictyol A; 10: Dictyol E); two sterols (11: Cholesterol sulfate; 12: (20R)-5α-
chlostane-3α,21-didyl disulfate); and two unusual phenylpropanoids (13: Maculalactone A;
and 14: Maculalactone E) have been isolated from this species. Among these compounds,
compounds 6–10 and 14 displayed moderate to significant cytotoxicity against a small
panel of human solid tumour cell lines. Further, the compound 5 exhibited antibacterial
activities against three Streptococcus and three Staphylococcus strains (Wang et al., 2004b).
3.9 FAMILY: OPHIURIDAE (ORDER: OPHIURIDA)

Ophiura albida (Forbes, 1839)
Common name(s): Serpent’s table brittle star

Global distribution: Northeastern Atlantic Ocean and in the Mediterranean Sea
Ecology: It occurs on the seabed on soft substrates including coarse sand, fine sand, gravel,
and muddy sand; depth range is 250–850 m.
Biology
Description: It has a central disc reaching a diameter of 1.5 cm and five arms, which are
up to 6 cm long. The disc is flattened and the upper surface is covered in small plates.
The disc is mostly brick red, but the plates at the edge of the disc close to where the arms
are attached are white. The arms are slender and fragile, segmented, and are gradually
tapering. The plates on the upper and lower sides have convex edges. Each segment has
three short spines. The underside of the disc is pale in colour and it has a central mouth.
Food and feeding: It is a predator and scavenger feeding on such small invertebrates as
polychaete worms, crustaceans and bivalve molluscs.
Predators: In the Irish Sea, it is eaten along with the common brittle star (Ophiothrix
fragilis) by the fast-moving seven-armed starfish (Luidia ciliaris).
Reproduction: Individuals of this species are either male or female. Fertilization is exter-
nal and the larvae are planktonic. It is a fast-growing brittle star and is not thought to live
for more than three years.

53126, Pseudomonas aeruginosa CIP 82118, and Salmonella enterica CIP 8039 with
MIC values of 12.5, 3.1, 6.3, 3.1, and 3.1 mg/mL, respectively (Marmouzi et al., 2018).
Ophiura (Ophiuroglypha) irrorata irrorata (Lyman, 1878) (= Ophiura irrorata)

Global distribution: Pacific Ocean and Antarctic Indian Ocean
Ecology: This is an epibenthic, polar species found at depths of 405–7199 m.
Biology
Description: The disc of this species is flat with 8 mm in diameter. The radial shields are
triangular with rounded corners. The dorsal surface is covered by plates and is without
spines/granules. The ventral interradial surface is plated. The oral shields are exposed,
pentagonal (some slightly notched), longer than wide. The adoral shields are exposed,
proximal to the lateral edge of the oral shield, separated radially, meeting interradially.
It has five arms that are unbranched, moniliform, or basally constricted, and are with
3–5 times the disc diameter. Dorsal arm plates are contiguous and are becoming sepa-
rate, without spines/granules. Ventral arm plates of the first free segments are separated,
squashed teardrop, and are 0.5–0.7 times long as wide.
Anti-Wnt activity (anticancer activity): Wnt signalling pathway is one of the key factors
in oncogenic transformation, growth, and metastasis in different cancers, including the
devastating triple-negative breast cancer. The extracts of this species containing anti-Wnt
compound(s) have been reported to inhibit the Wnt pathway in triple-negative breast cancer
(TNBC) cells (Blagodatski et al., 2017).
Ophiura kinbergi (Ljungman, 1866)

Global distribution: Indo-Pacific
Ecology: This subtropical, benthic species is found in soft sediments; depth range is
0–500 m. It is an indicator species for fine sandy mud substrata.
Biology
Description: The disc of this species is flat or tumid with 4 mm diameter. The radial
shields are round or triangular with rounded corners. The dorsal surface is covered by
plates and without spines/granules. The ventral interradial surface is plated. The oral
shields are exposed, arrowhead shaped (triangular pointed proximally, rounded distally),
and are longer than wide. The adoral shields are exposed, proximal to the lateral edge of
the oral shield, and are separated radially. It has five arms that are unbranched and basally
constricted. Dorsal arm plates are contiguous and without spines/granules. Ventral arm
plates of the first free segments are separated and are fan shaped. There are 2–3 arm spines
on the first ventral segment and 3 on the first free segments.
Food and feeding: It is a predator.

Antioxidant and antibacterial activities: The polysaccharide extracted from this species
had good scavenging effect on hydroxyl radical(·OH) and superoxide anion radical(O2-),
and the clearance rate showed a dose-effect relationship with the polysaccharide solution.
The maximum clearance rates of OH and of O2 polysaccharide were 45.68% and 40.47%,
respectively. Further, its polysaccharide also had good antibacterial activity on E. coli,
S. aureus and S. cerevisiae. The diameters of antibacterial circle were 13.46, 14.12, and
14.73 mm, respectively (Zhen et al., 2014, 2015).
Ophiura leptoctenia (H.L. Clark, 1911)

Global distribution: Northeast Pacific: The Sea of Okhotsk
Ecology: This temperate species has a depth range of 25–3239 m.
Biology
Description: Its aboral disc is bumpy with very large scales. Radial shield is in the form
of spaced buttons. Oral disc has large scales. Arms get inserted into the disc. Oral papillae
are in the form of spikes.
Shubina et al. (1998) reported on the isolation of (20R)-5acholestane-3a,21-diol disulfate
and (20R)-cholest-5-ene3a,4b,21-triol 3-sulfate from this species. Bioactivity of this spe-
cies is not known.
Ophiura sarsii (Lütken, 1855)
Common name(s): Notched Brittle Star

Global distribution: In the North Sea, this species is found as far south as Helgoland. The
species is circumpolar. At the Atlantic east coast, it goes down as far as south Ireland. In
the Pacific, it is found as far south as Japan and California.
Ecology: The species is found in soft sediments to about 2000 m. It lives on soft sediments,
the surface of sand or muddy sand. It occasionally burrows shallowly and moves rapidly by
a swimming action upon being disturbed; depth range is 10–3000 m.
Biology
Description: The body of this species has a central disc that is up to 35 mm in diameter. Its
five arms are about 150 mm in length. The scales of the disc are rather coarse, and the primary
plates are generally distinct. The radial shields are about half the length of the disc radius
and are separated or contiguous at their broadest part. The innermost dorsal arm plates are
usually rectangular or triangular. The free dorsal arm plates in the proximal part of the arm are
4–5 times as broad as long, with a straight or concave outer edge and are contiguous throughout
their whole breadth. The ventral arm plates are twice as broad as long, with a convex outer edge.
Colour of the specimens varies from dull grey-brown to sandy orange.
Food and feeding: It is a deposit feeder.

Haemolytic activity: The steroidal disulfate (disodium salt) Cholest-5-ene-3α,4β,21-triol
3,21-disulfate isolated from this species has shown haemolytic activity to mouse erythro-
cytes (Atta-ur-Rahman, 1995).
Others: Two sulfated steroid polyols, viz. cholest-5-ene-3α,4β,21-triol 3,21-di(sodium

sulfate) and cholest-5-ene-2β,3α,21-triol tri(sodium sulfate) have been isolated from this
species (Levina et al., 1990).
3.10 FAMILY: OPHIOPYRGIDAE (ORDER: OPHIURIDA)
Ophiosparte gigas (Koehler, 1922)
Common name(s): Snake star

Global distribution: Not reported
Ecology: The snake star crawls over the seabed with its flexible, snake-like arms.
Biology
Food and feeding: It is an active predator and scavenger whose diet consists of sponges,
ophiuroids, bivalves, polychaetes, and crustaceans.
Reproduction: Asexual reproduction is common in this species. The snake-like arms of
this species break off easily, but they grow again automatically.

Others: Atta-ur-Rahman (1995) reported on the occurrence of two polyhydroxysteroids
from this species.
HIV-inhibitory activity: The two sulfated sterols isolated from this species exhibited
anti-HIV activity. While the first compound inhibited both HIV-1 and HIV-2 with IC 50
values of >161 and >161 respectively, the second compound showed activity against HIV-1
only with IC 50 values of <0.02 µM (McKee et al., 1994).
4 Biology and Ecology
of Pharmaceutical Sea
Urchins and Sand Dollars
(Class: Echinoidea)
4.1 FAMILY: ARBACIIDAE (ORDER: ARBACIOIDA)

Arbacia lixula (Linnaeus, 1758)
Common name(s): Black sea urchin

Global distribution: Tropical and subtropical; coast of the Mediterranean Sea and
Macaronesian Islands (Azores, Madeira, Canary Islands), and less commonly, on the
Atlantic coast of Western Africa and the Brazilian coast
Ecology: This benthic species is found typically in the shallow rocky reefs at depths from 0
to 50 m. It has a good resistance to hydrodynamism due to good attachment strength to rocks.
Biology
Description: It is a medium-sized sea urchin, characterized by its deep black colour and
its hemispherical shape. All of its spines are roughly the same size (no “secondary spines”)
and worn erected (never dishevelled when in the water). The anus on the top is surrounded
by four plates forming an anal valve. The oral face is nearly naked, the mouth being sur-
rounded by soft, dark greenish skin.
Food and feeding: It feeds mainly on crustose red algae and small filamentous algae.

Neuroprotective activity: The sea urchins are valuable sources of bioactive compounds.
The gonads of some sea urchin species contain antioxidants including carotenoids and
polyhydroxylated naphthoquinones (PHNQs), such as echinochrome Arbacia lixula has a
197
high concentration of the carotenoid pigment, astaxanthin (27.0 μg/mg), which is known

to have particular bioactivity for the prevention of neurodegenerative diseases. Further,
methanol crude extracts obtained from eggs of farmed and wild specimens of this species
revealed a higher bioactivity (Cirino et al., 2017).
Lysozyme activity: The coelomic fluid and coelomocyte lysates of this species showed
lysosome activity with a mean diameter of lysis of 7.7 mm (Stabili et al., 2018).
Antioxidant Activity: This species also showed the highest antioxidant activity (by TEAC
assay) of 1792.75 nmolTE/mL sample (Stabili et al., 2018).
4.2 FAMILY: ECHINIDAE (ORDER: CAMARODONTA)

Echinus esculentus (Do, 2012)
Common name(s): European edible sea urchin, common sea urchin
Global distribution: It is found off the coasts of Portugal, Spain, France, Belgium,
Netherlands, Denmark, Norway, Sweden, the United Kingdom, and Ireland.
Ecology: It is common in all coastal areas with hard substrates, down to a depth of 1200 m.
Biology
Description: It is almost spherical but slightly flattened at both poles. It is reddish or pur-
plish with white tubercles. Its brittle, limy test is rigid and is divided into five ambulacral
areas separated by five interambulacral areas. There are two rows of plates in each of these
areas, making twenty rows of plates in total. The test is covered in spines, each articulating
with a tubercle. There is also a dense covering of secondary spines and a smaller number
of longer, primary spines. The spines are blunt ended and usually white with purplish tips.
There is a radially symmetrical pattern of holes in the ambulacral areas through which the
tube feet emerge. It typically reaches a diameter of 15–18 cm.
Food and feeding: It feeds on worms, barnacles, hydroids, tunicates, bryozoans, algae
(such as Laminaria spp.), sludge, and detritus.
Reproduction: Spawning in this species mainly occurs in the spring, and a large female
may release about 20 million eggs into the water column. The larvae become part of the
plankton, the development of which is complex and takes between 45 and 60 days in
captivity. It includes a blastula, gastrula, and a four-armed echinopluteus stage that forms
an important part of the zooplankton. Settlement mostly occurs in autumn and winter, and
the largest number of juvenile urchins was found at a greater depth than the kelp zone.
Biology and Ecology of Pharmaceutical Sea Urchins and Sand Dollars (Class: Echinoidea) 199
Associated species: The polychaete worm Adyte assimilis and the copepod
Pseudoanthessius liber are often found living as commensal organisms among its spines,
and the parasitic copepod Asterocheres echinola often infests its gut.
Edible values: The roe of this sea urchin is used as food around the world. It is not actually
the eggs that are eaten but the gonads, both male and female.

Anti-inflammatory activity: The HPLC fraction 27 of the organic gonad extract of this
species exhibited anti-inflammatory activity on U937 cell lines (Do, 2012).
Immunostimulatory activity: The intestines extract of this species displayed a weak
immunostimulatory activity (Do, 2012).
Antimicrobial activity: Three antimicrobial peptides—viz. two heterodimeric peptides
(EeCentrocin 1 and 2) and a cysteine-rich peptide (EeStrongylocin 2)—have been isolated
from the haemocytes of this species. Of these compounds, EeCentrocins showed antimi-
crobial activity against gram-negative bacteria, gram-positive bacteria, and against fungi
(Solstad et al., 2016; López et al., 2018).
López et al. (2018) also reported that the quinona pigments—viz. echinochrome A and spi-
nochrome D of this species—exhibited activity against gram-positive and negative bacteria.
4.3 FAMILY: ECHINOMETRIDAE (ORDER: CAMARODONTA)

Echinometra lucunter (Linnaeus, 1758)
Common name(s): Rock boring urchin

Global distribution: It is common throughout the Caribbean Sea and also occurs in
Florida, Bermuda, and the South American coast as far south as Brazil.
Ecology: This tropical species occurs on very shallow, rocky areas and on coral reefs usu-
ally; and among seagrasses at depths of 2 m or less but occasionally in deeper water down
to about 45 m. It sometimes occurs in large numbers and causes considerable damage to
coral reefs through its boring activities.
Applied ecology: An overabundance of this species occurs in areas with few predators,
and this species may therefore be used as an indicator species for environmentally stressed
or overfished coral reefs.
Biology
Description: It has an elliptical rather than a round test (shell). It can grow to a diameter of
8 cm and grows larger at the extreme north and south ends of its range than it does in the
centre. It has moderately short spines with wide bases and sharp tips. The colour of the test
varies from black to deep brownish-red. The spines are usually black.
Food and feeding: It has a specialized feeding apparatus (Aristotle’s lantern) with five
teeth. It grazes on the nonliving reef structure as well as on live coral surfaces; 95% of its
diet consists of algae such as Laurencia papillosa and the other 2% are invertebrates (such
as the sponge Darwinella sp.). It is also known to consume some seagrasses.
Predator: The king helmet shell (Cassis tuberosa) feeds on it.
Associated species: Several species of small goby conceal themselves underneath its test.
Behaviour: Echinometra lucunter uses the teeth that surround its mouth to grind away at
the rock underneath it so as to make a hemispherical depression in which it takes refuge
during the day. It emerges at night to graze on algae growing within a few centimetres of
its home. It defends this hole against other sea urchins of its own species.
Reproduction: Males of this species reach maturity from 18 mm and females from 21 mm
in diameter. This species spawns generally once or twice a year. Mating is polygamous, in
that a single female is able to mate with several males, because eggs and sperms are both
released into the water column. The male E. lucunter releases his spermatocytes first into
the column. This in turn stimulates the female to release her oocytes from the gonad.

Anticoagulant and antithrombotic activities: The polysaccharide, 2-sulfated, 3-linked
α-l-galactan (which is generally abundant in marine algae) isolated from the egg jelly
layer of this species showed potent anticoagulant and antithrombotic activities (Pereira
et al., 2002).
Ethnomedicine: This species has been reported to be a marine-based ethnomedicine for
the treatment of asthma (Anon., Marine-based ethnomedicine—Asthma-Ethnic group/
Georeference, Recife, Pernambuco, Brazil; Narchi, https://getd.libs.uga.edu/pdfs/narchi_
nemer_e_201105_phd.pdf).
Others: The MeOH/chloroform extracts of this species yielded a compound, sterol deriva-
tive (B) (Kaihil et al., 2016).
Ould et al. (2012) reported on the isolation of two lipids (sterol derivatives), viz. lucunter-
peracetate and peroxylucunterine from the CHCl3–MeOH 1:1 (v/v) extract of this species.
Echinometra mathaei (Blainville, 1825)
Common name(s): Rock—burrowing urchin

Global distribution: It is found in the Indo-Pacific and Western Central Atlantic regions;
its range extends from Madagascar, the East African coast, and the Red Sea to Hawaii.
Ecology: This tropical, benthic species is found subtidally; it inhabits fore reefs and hard
substrates by boring on basaltic and calcareous rocks in shallow waters; depth range is 0–139 m.
Biology
Description: It grows to a test diameter of 8 cm. The colour is quite variable, but the test
is usually a dark colour. The spines are sometimes green and purple with purple tips or
entirely green with purple tips. This sea urchin can be distinguished from other related
species by a characteristic pale ring at the base of each spine.
Behaviour: It uses its spines and teeth (part of the mouthparts known as Aristotle’s lan-
tern) to dig itself into the basaltic and calcareous rock where it lives. It emerges from these
hollows at night to graze on algae.
Food and feeding: It feeds on algae and small invertebrates.
Reproduction: As with other sea urchins, breeding in this species involves releasing
gametes into the water column. Fertilization is external and the echinopluteus larvae are
planktonic. When these settle on the seabed, they undergo metamorphosis into juvenile sea
urchins. In the Gulf of Suez, in the most northerly part of its range, it spawns in the sum-
mer and autumn but in warmer waters, breeding takes place at any time of year.
Predators: Its natural predators are mostly finfish, but there is also some predation by
brittle stars and gastropod molluscs.
Associated species: Commensal shrimp species, Athanas areteformi and Athanas indica
live among the spines of this sea urchin and its appearance is mimicked by the mantis
shrimp, Echinosquilla guerinii, which conceals itself in a hole in the rock with only its
spiny telson visible.

Antioxidant and amylase inhibitory properties: With respect to antioxidant properties,
the methanol extract of the shell of this species possessed the maximum reducing power
(OD = 0.48) at 40 μg/mL concentration and, the ethyl acetate extract of Aristotle’s lantern
at 1 mg/mL concentration presented the highest TAC (OD = 0.44). Further, the n-hexane
extract of gonad at a concentration of 1 mg/mL revealed the highest α-amylase inhibition
(84.7%) (Soleimani et al., 2017).
Anti-inflammatory property: The ethyl acetate extract of Aristotle’s lantern of this spe-
cies possessed the highest anti-inflammatory property (69.78 μg/mL) at 20 μg/mL concen-
tration (Soleimani et al., 2017).
Antibacterial, antioxidant, and cytotoxic activities: The spinochromes, viz. Spinochrome
B, two isomers of Spinochrome D, Spinochrome E, one isomer of Spinochrome A,
Echinochrome A, and Spinochrome C of this species showed antibacterial, antioxidant,
and cytotoxic activities. The crude extracts of this species showed a strong antibacterial
activity against all bacteria but appeared to be more efficient against E. coli and B. subtilis
(EC50 < 800 μg/mL) than against the other strains (EC50 > 1100 μg/mL). In the 2,2-diphenyl-
1-picrylhydrazyl antioxidant assays, spinochromes showed an enhanced antioxidant activity
compared to the positive control. These spinochromes also presented a pro-inflammatory
effect on lipopolysaccharide-stimulated macrophages, highlighting their possible implication
in the sea urchin immune system. Further, these spinochromes showed a weak cytotoxicity
against human cells. All these results confirm the implication of the spinochromes of this
species reveal their potential for pharmacological industries (Brasseur et al., 2017).
Antibacterial and haemolytic effects: The organic extracts (ethanol and acetonitrile) of
the gonadal tissue and test of this species displayed antibacterial effects against strepto-
coccal species, Streptococcus mutans and Streptococcus sobrinus. Further, these extracts
(80%) also had haemolytic activity (Kazemi et al., 2016).
Heliocidaris crassispina (A. Agassiz, 1864) (= Anthocidaris crassispina)
Common name(s): Purple sea urchin

Global distribution: Northwest Pacific: Hong Kong
Ecology: It is found in the rocky shores and rock pools at the low tides.
Biology
Description: It has a globular body consisting of pentamerous radial plates with movable
spines. Body and spines are black. It has a size range of 50–70 mm.
Food and feeding: It feeds on algae, sessile animals on rocks, and dead animals.
Reproduction: It has a life span of nine years. Juveniles of this species settle at a size of
between 0.4 and 0.7 mm test diameter and maximum growth rates occur in the years 2–4.

Anti-inflammatory activity: The gonad polysaccharide of this species showed the most
potent anti-inflammatory activity. The NO inhibition rate of its shell polysaccharides was
found to be 70% (Jiao et al., 2015).
Neuroprotective activity: Ganglioside, viz. NeuGc α 2-6Glc β 1-1Cer (M5 ganglioside)
and HSO3-8NeuGc α 2-6Glc β 1-1Cer (T1 ganglioside) have been reported from the
eggs of this species. These compounds have a function in memory, and disturbances in
ganglioside synthesis may lead to neurodegenerative disorders (Kubo et al., 1990).
Antioxidant property: The naphthoquinone pigments as well as known antioxidant,
α-tocopherol, of the shell of this species exhibited 1,1-diphenyl 2-picryhydrazyl (DPPH)
radical scavenging ability. Superoxide anion radical and hydrogen peroxide were also
scavenged. These findings suggest that the shells of this species that are still regarded
as food waste would be a new bioresource for obtaining natural antioxidant (Kuwahara
et al., 2009).
Osteogenesis and adipogenesis: The acetone extracts of the spindle and gonad of this
species have been reported to stimulate both osteogenesis and adipogenesis in mouse
mesenchymal stem cells (MSCs). These findings suggest that the spindle and gonad from
this species may be beneficial in the prevention of some bone-related diseases and insulin
resistance via its potency to stimulate osteogenesis and adipogenesis of MSCs (Li et al.,
2010c).
Others: A sulfonoglycolipid, which is a 96: 4 mixture of 1′-O-palmitoyl-3′-O-(6-sulfo-α-
d-quinovopyranosyl) glycerol, has been isolated from the shell of this species. The role of
this compound in pharmacology is to be known (Kitagawa et al., 1979).
Heliocidaris erythrogramma (Valenciennes, 1846)

Global distribution: It is endemic to Australia; southern Queensland, New South Wales,
Victoria, Tasmania, South Australia, and Western Australia.
Ecology: These organisms inhabit coastal waters up to 35 m in depth but are most com-
mon in waters shallower than 10 m. As benthic animals, they are often found attached to
rocky reefs, stones, seagrass beds, in crevices and burrows, and on sandy mud bottoms.
The species seems to prefer areas not fully exposed to wave motion. It moves mostly at
night and shows little fidelity to individual crevices and shelters.
Biology
Description: This species has a hard exoskeleton with numerous sharp, solid spines that
taper to a point. Dermis colour of this species may be red, purple or white. Its tube feet
and the mouth are located on the underside of the body. Purple sea urchins can reach up to
14 cm in test (external skeleton surrounding the body) diameter.
Food and feeding: Purple urchins feed by actively grazing rock surfaces and by trapping
algae or seagrasses.
Reproduction: It produces negatively buoyant, 95 µm-diam. eggs and the sperm have
a head region of 4 µm in length. It has a nine-month breeding period characterized by
the continual presence of nutrient reserves and vitellogenic oocytes that rapidly replaced
spawned ova (Laegdsgaard et al., 1991).

Antioxidant activity: The aqueous extract of male gonadal tissue of this species showed
antioxidant activity by scavenging the reactive oxygen species (ROS), hydrogen peroxide
(H2O2) (Sheean et al., 2007).
Anti-inflammatory activity: The dichloromethane, methanol, and aqueous extracts of
male gonadal tissue of this species displayed various levels of anti-inflammatory action
in connection with inhibition of cyclooxygenase (ovine COX-1 and COX-2). Further, the
dichloromethane and aqueous extracts also showed anti-inflammatory activity in rats
developing adjuvant-induced polyarthritis (Sheean et al., 2007).
Arthritis inhibition: The dichloromethane extract of this species showed 75% inhibitory
of arthritis symptoms, and the aqueous extract was 55% inhibitory (Sheean et al., 2007).
Antileukaemic activity: The dichloromethane extract of this species displayed the
suppression of growth of the mouse leukaemic lymphoblastic (P388) cell line (Sheean
et al., 2007).
Cytotoxicity: The dichloromethane extract of this species was also able to exert a cyto-
toxic effect on the tumour cell line (Sheean et al., 2007). Kilimnik et al. (2016) reported
that the acetylenic carotenoids diatoxanthin and alloxanthin present in the gonads of this
species showed cytotoxicity.
Heliocidaris tuberculata (Lamarck, 1816)
Common name(s): Red tipped urchin, tuberculate urchin

Global distribution: It is found in the Western Pacific. This species is common throughout
the Solitary Islands Marine Park. It is found on the east coast of Australia from Caloundra,
Queensland, to Eden, New South Wales, and including Lord Howe Island, Kermadec
Island, and New Zealand.
Ecology: This benthic, subtropical species can be readily seen in the pools around rock
platforms and in deeper water on reefs of the lagoon; depth range is 0–10 m.
Biology
Description: It is a large brown (orange red) sea urchin with a test size of 115 mm in diam-
eter. Its spines are also brown or tinged with orange red, and they are 5–8 cm long. These
spines are blunt ended and have an oval cross section.
Associated species: Small, brown shrimps of the species Athanas dorsalis are found asso-
ciated with this species of sea urchin. These shrimps live permanently in this spiny home,
safe from predators. It is possible that they provide some function that serves the interest
of the urchin, such as cleaning parasites from the body.
Reproduction: It produces buoyant, 450 µm-diam. eggs and the sperm have a head region
of 10 µm in length. It had a seasonal reproductive pattern with a three-month summer
spawning period (Laegdsgaard et al., 1991).

Cytotoxicity: The acetylenic carotenoids, diatoxanthin, and alloxanthin present in the
gonads of this species showed cytotoxicity (Kilimnik et al., 2016).
4.4 FAMILY: PARECHINIDAE (ORDER: CAMARODONTA)

Paracentrotus lividus (Lamarck, 1816)
Common name(s): Purple sea urchin, rock sea urchin, stony sea urchin
Global distribution: This subtropical, tropical, and temperate species is found through-
out the Mediterranean Sea and in the eastern Atlantic Ocean from western Scotland and
Ireland to the Azores, Canary Islands, and Morocco. It is most common in the western
Mediterranean, the coasts of Portugal, and the Bay of Biscay.
Ecology: This benthic species occurs on rocks and boulders and in seagrass meadows of
Zostera marina and Posidonia oceanica; depth range is 0–90 m.
Biology
Description: It has a circular, flattened, greenish test with a diameter of up to 7 cm. The
test is densely clothed in long and sharply pointed spines, which are usually purple but are
occasionally with other colours including dark brown, light brown, and olive green. There
are five or six pairs of pores on each ambulacral plate. The tube feet are in groups of five
or six, arranged in small arcs.
Food and feeding: It is an herbivore, mainly eating a range of red, green, and brown algae
in addition to seagrass; occasionally it feeds on animal materials.
Predators: The main predators of this species are the spider crab (Maja crispata), the fish
Diplodus sargus, Diplodus vulgaris, Labrus merula, and Coris julis, and the gastropod,
Hexaplex trunculus. The spiny starfish (Marthasterias glacialis) is a main predator
elsewhere.
Reproduction: Individual P. lividus are either male or female although hermaphrodit-
ism has been observed. They aggregate for spawning and release gametes into the water
column. The larvae form part of the zooplankton for 28 days before settling and undergo-
ing metamorphosis.
Edible values: The gonads of this species are considered a delicacy in Lebanon, France,
Italy, Spain, Malta, and parts of Croatia, most notably on the island of Korčula, and are
also eaten to a lesser extent in Greece. The urchins have been harvested for export over a
wider area including Croatia, Portugal, and Ireland.
Others: P. lividus is also an excellent animal model for toxicology, physiology, and biology
investigations, having been used for more than a century as a model for embryological
studies with synchronously developing embryos that are easy to manipulate and analyse
for morphological aberrations.

Hypoglycemic, hypolipidemic, and antioxidant properties: The organic extract of this
species has shown hypoglycemic, hypolipidemic, and antioxidant properties. Based on the
studies already made, the above extract may be useful as therapy against oxidative stress
and liver damage in diabetic rats (induced by high fat diet/streptozotocin) with both type 1
diabetes mellitus and type 2 diabetes mellitus (Soliman, 2016).
Apoptosis regulation: The metallothionein (protein) content of this species plays an
important role in the regulation of apoptosis (Ragusa et al., 2017). Further, this compound,
which is capable of binding heavy metals, plays a role in the homeostasis of essential metals
such as zinc and copper, detoxification of toxic metals such as cadmium, and protection
against oxidative stress.
Antibacterial activity: An antimicrobial peptide, paracentrin 1 isolated from the
coelomocyte cytosol of this species showed antibacterial activity against different strains
of Staphylococcus aureus and Pseudomonas aeruginosa ATCC 15442 at concentrations
ranging from 12.5 to 6.2 mg/mL. The MIC values recorded for these bacterial strains are
given below (Schillaci et al., 2014).
MIC Values of SP1 (Synthetic Paracentrin 1) Tested Against Bacterial Strains

Bacterial Strain MIC (mg/mL)
S. aureus ATCC 25923 12.5
S. epidermidis RP62A 6.2
S. aureus 100 12.5
S. aureus 657 6.2
S. aureus 700 12.5
S. aureus 702 12.5
P. aeruginosa ATCC15442 12.5
Source: Schillaci, D. et al., AMB Express, 4, 78, 2014.
A bactericidal protein isolated from the coelomocytes of this species has shown antibacte-
rial activity (Petzelt, 2005).
Stabili and Canicattì (1994) reported that the seminal plasma of this species exerted an
inhibitory action on the growth of bacterial colonies.
Antitumour activity: The acetylenic lipid compound, asterinic acid
(7,8-Didehydroastaxanthin), isolated from this species showed antitumour activity
(Kilimnik et al., 2016).
Others: Atta-ur-Rahman (1995) reported on the occurrence of redox-reactive 4-thiohistiidne

compounds from the eggs of this species.
4.5 FAMILY: STRONGYLOCENTROTIDAE (ORDER: CAMARODONTA)

Mesocentrotus nudus (A. Agassiz, 1864) (= Strongylocentrotus nudus)

Global distribution: Northwest Pacific: Japan
Ecology: It lives in a rocky tidal zone. Yearling sea urchins inhabited the subtidal zone
throughout the year.
Biology
Description: In this species, the spines have a large meshwork core in which radial wedges
are not apparent. It reaches a maximum size of 76 mm over the spines.
Food and feeding: It feeds on algae and epibenthic animals belonging to the phyla, viz.
Porifera, Cnidaria, Annelida, Mollusca, and Arthropoda. Yearling sea urchins, however,
showed their preference for attached diatoms or small algae.

Antioxidant activity: The aminonaphthoquinone, spinamine E, isolated from this species
showed scavenging activity of the DPPH radical and inhibited lipid peroxidation. All the
investigated naphthoquinones of this species showed a high antiradical activity, which was up
to 1.5 times higher than that of α-tocopherol. In a lipid peroxidation inhibition testing model,
spinamine E showed the highest inhibitory effect (Vasileva et al., 2016; Blunt et al., 2017).
Anti-obesity and hepatoprotective activity: Mice fed with this species exhibited lower
body, liver, and visceral fat weights; lower plasma levels of aspartate aminotransferase
and alanine aminotransferase; and lower hepatic triacylglycerol levels than those fed
carboxymethyl cellulose (CMC). Additionally, sea urchin consumption resulted in enhanced
levels of arachidonic acid and docosahexaenoic acid within mouse livers. Lastly, mice fed
with high-fat diet (HFD) with sea urchin (500 mg/kg) exhibited increased mRNA expression
of uncoupling protein-1 within brown adipose tissue, compared with those fed HFD with
CMC. These findings suggest that the consumption of sea urchin might provide a protective
effect against the development of obesity and/or nonalcoholic fatty liver disease (Yamamoto
et al., 2018).
Strongylocentrotus droebachiensis (O. F. Müller, 1776)
Common name(s): Green sea urchin

Global distribution: Widely distributed circumpolar species found in the boreal regions
of the Atlantic and Pacific Oceans
Ecology: It is found in rocky substratum in the intertidal and up to depths of 1150 m. It is
a euryhaline species and can survive in waters of low salinity.
Biology
Description: This species is in the shape of a slightly flattened globe (dorsoventrally).
The oral side rests against the substratum, and the aboral side (the side with the anus) is in
the opposite direction. It has pentameric symmetry, which is visible in the five paired rows
of podia (tube feet) that run from the anus to the mouth. It has a characteristic green colour.
This is a relatively fast-growing sea urchin and its maximum diameter is 87 mm.
Food and feeding: It feeds on algae, preferring species such as Sargassum muticum
and Mazzaella japonica over Saccharina latissima, Ulva sp., and Chondracanthus
exasperatus.
Predators: This species is eaten by a variety of predators, including sea stars, crabs, large
fish, mammals, birds, and humans.
Associated species: Snails of the families Melanellidae and Stiliferidae live on the surface
of the test and adhere their own eggs to the base of the spines as protection.
Others: It is a species of aquaculture.

Antineoplastic (antitumour) activities: The glycoproteins isolated from this species have
shown antineoplastic activities (Bhakuni and Rawat, 2005).
Pettit et al. (1981) reported that the presence of antineoplastic glycoproteins, viz. stron-
gylostatin 1 strongylostatin 2 in this species. Treatment of the murine P388 lymphocytic
leukaemia with strongylostatin 2 resulted in 39%–42% life extension at a dose of 4.5 mg/kg.
Antibacterial and haemolytic activities: The coelomocyte and body wall extracts of this
species have shown antibacterial activity. Further, its body wall extracts have also dis-
played haemolytic activity (Haug et al., 2002).
Antibacterial activity: The antibacterial peptides named strongylocins 1 and 2 from the
coelomocyte extracts of this species displayed potent activities against gram-negative and
gram-positive bacteria (Li et al., 2008).
Strongylocentrotus intermedius (A. Agassiz, 1864)

Common name(s): Short-spined sea urchin, Japanese common sea urchin

Global distribution: Japan
Ecology: It is found commonly on the rocky bottom in shallow waters.
Biology
Description: The somatic growth of this species is largely influenced by the kind and
quantity of marine algae it eats. Its size has been reported as 69 mm over spines.
Food and feeding: The principal food of this species is marine algae.
Edible values: The roe flesh of this species is eaten mainly as sashimi and sushi and is
served in a bowl with rice. Salted raw urchin is a familiar food in Hokkaido.

Antitumour effect: Four glycolipids, viz. 3′-sulphonoquinovosyl-1′, 2′-diacylglyceride
(A-4), 3′-sulphonoquinovosyl-1′-monoacylglyceride (2′-lyso A-4, A-5), NeuGc(α)2-6Glc(β)1-
1ceramide (A-6), and HSO3-8NeuGc(α)2-6Glc(β)1-1ceramide (A-7), have been isolated from
the intestine extracts of this species. Among these compounds, the compound, A-5 showed
significant activity against tumour cells in vitro. Further, the compound A-5 also exhibited
antitumour effects on nude mice bearing solid tumours of a human lung adenocarcinoma
cell line A-549. Pathologically, the solid tumours showed haemorrhagic necrosis areas after
treatment with A-5. These findings suggest that sulphonoquinovosyl-lysoglyceride (A-5) of this
species could be a useful drug for cancer chemotherapy (Sahara et al., 1997).
Antioxidant, antibacterial, antifungal, and antitumour activities: The quinone pigments
of this species, viz. spinochromes have been reported to possess antioxidant, antibacterial,
antifungal, and antitumour activities (Ageenko et al., 2014).
Strongylocentrotus pallidus (Sars G. O., 1872)
Common name(s): White sea urchin, pale urchin

Global distribution: It is found distributed in Pan-Arctic and subarctic regions.
Ecology: This polar species occurs on both sandy/shell hash bottoms and in rocky areas. It
is strictly subtidal and is usually found in areas deeper than 30 m.
Biology
Description: Its shorter spines are whitish or pale yellow-green with very light-coloured
tube feet. Its globiferous pedicellariae are smaller and are pigmented along edges of the
jaw. Trident pedicellariae are prominent, white, and particularly on upper test. Life span
of this species has been reported to be more than 40 years in high Arctic regions. It has a
diameter range of 30–90 mm.
Food and feeding: It is a deposit feeder and grazer.
Associated species: This deep-living species may harbour the intestinal rhabdocoel flat-
worm Syndisyrinx (= Syndesmis) franciscanus. We find this species on sandy/shell hash
bottoms at about 100–120 m depth.

Antioxidant activity: The aminonaphthoquinone, spinamine E, isolated from this species
showed radical-scavenging activity (Blunt et al., 2017). Vasileva et al. (2016) reported that
this compound showed a high antiradical activity, which was up to 1.5 times higher than
that of α-tocopherol. In a lipid peroxidation inhibition-testing model, spinamine E showed
the highest inhibitory effect.
Strongylocentrotus purpuratus (Stimpson, 1857)


Global distribution: It is found distributed along the eastern edge of the Pacific Ocean
extending from Ensenada, Mexico, to British Columbia, Canada.
Ecology: It occurs as a lower intertidal and nearshore subtidal community. Further, this
purple sea urchin, along with sea otters and abalones, is a prominent member of the kelp
forest community.
Biology
Description: This sea urchin species is deep purple in colour, and its eggs are orange when
secreted in water. It normally grows to a diameter of about 10 cm and may live as long as
70 years.
Food and feeding: It mainly feeds on the giant kelp, Macrocystis pyrifera.
Reproduction: January, February, and March function as the typical active reproductive
months for this species. Sexual maturity is reached around two years.
Edible values: This species was used for food by the indigenous peoples of California.
They ate the yellow egg mass raw.

Antimicrobial activity: The homologous putative proteins called SpStrongylocins 1 and 2
of this species showed antimicrobial activity against both gram-negative and gram-positive
bacteria. The results of membrane integrity assays against cytoplasmic membranes of E.
coli suggested that both these compounds conduct their antibacterial activity by intracel-
lular killing mechanisms (Li et al., 2010b).
Susceptibility of Bacterial Strains to the Recombinant Antibacterial Peptide

SpStrongylocins 1 and 2
Minimal Inhibitory Concentration (µM)
Peptide L. anguillarum E. coli C. glutamicum S. aureus
Recombinant SpStrongylocin 1 15.0 7.5 7.5 15.0
Recombinant SpStrongylocin 2 15.0 7.5 3.8 15.0
Source: Li et al., 2010b.
Antioxidant activity: A highly reducing antioxidant mercaptohistidine, ovothiol A

(N1-methyl-4-mercaptohistidine), isolated from the eggs of this species has been reported
to scavenge hydrogen peroxide released during fertilization (Atta-ur-Rahman, 1995).
.
4.6 FAMILY: TEMNOPLEURIDAE (ORDER: CAMARODONTA)

Temnopleurus alexandri (Bell, 1884)
Common name(s): Alexanders sea urchin

Global distribution: Western Central Pacific: South China Sea; Australia
Ecology: This tropical, benthic species inhabits coarse substrates such as rocky reefs as
well as seaweed beds; depth range is 9–73 m. It usually camouflages itself with debris and
weed fragments.
Biology
Description: This species exhibits a variety of test and spine colours, including shades of
green and purple. It has a size of 70 × 43 mm.
Behaviour: These individuals display collecting behaviour, covering themselves with vari-
ous materials available in the water column. They do this using their advanced tube feet,
which are involved in a number of other processes, such as locomotion, feeding, and sen-
sory perception.
Food and feeding: It feeds on various plant matter and loose benthic materials.

Antibacterial activity: The hexane extract of this species displayed antibacterial activity
against the gram-positive bacterial species, viz. Staphylococcus aureus ATCC 25923,
Bacillus subtilis MTCC 441, and Enterococcus faecalis ATCC 29212; and gram-negative
species, viz. Escherichia coli ATCC 25922, Pseudomonas aeruginosa ATCC 27853,
and Proteus vulgaris MTCC 1771. Lowest (2.5 ppm) MIC was noted for B. subtilis and
P. aeruginosa (Uma and Parvathavarthini, 2010).
Angiogenesis activity: The compound, pentadecanoic acid, 14–methyl–methyl ester
isolated from the hexane crude extract of this species showed angiogenesis activity
(Parvathavarthini and Uma, 2016).
Temnopleurus toreumaticus (Leske, 1778)
Common name(s): Black sea urchin, striped spine sea urchin

Global distribution: Indo-West Pacific
Ecology: This benthic, subtropical, and abyssal species inhabits sandy areas near sea-
grasses, in coral rubble and rocky shores, and under jetties. Depth range is 0–82 m.
Biology
Description: Body diameter of this species is 4–5 cm. It has short, slender spines (1–2 cm)
with long, translucent tube feet that may extend past the spines. The spines on the upper
side are black and pointed. Spines on the underside are flattened and may be banded.
Some individuals may have light-coloured zigzag lines radiating from the centre around
the body. This sea urchin appears to “carry” shells and other debris. This behaviour may
help camouflage it or shield it from sunlight.
Food and feeding: It feeds on the Rhodophyta, Corallina sp.

Cytotoxic and antimicrobial activities: Aqueous extract of this species possessed
cytotoxic activity with an LC50 value of 0.12 mgmL−1. Further, this extract showed
antimicrobial activity against K. oxytoca (maximum zone inhibition dia., 12.26 mm).
The chloroform extract was against Mucor sp. with inhibition zone dia. value of 3.33 mm
(Bragadeeswara et al., 2013).
Others: Two glycosphingolipids, temnosides A and B, have been isolated from the MeOH
extract of this species (Babu et al., 1997).
4.7 FAMILY: TOXOPNEUSTIDAE (ORDER: CAMARODONTA)

Lytechinus variegatus (Lamarck, 1816)
Common name(s): Green sea urchin, variegated sea urchin

Global distribution: Warm tropical waters of the western Atlantic Ocean and Caribbean
Sea
Ecology: It is found on rocky reefs, on or under rocks, on sandy or muddy substrates and
in seagrass meadows; depth range is 1–17 m.
Biology
Description: This green sea urchin has a globular test (shell) densely covered in spines, and
it can reach a diameter of 11 cm. The test may be purple, green, or dull red, blotched with
white. The majority of the spines are short, but there are a few longer primary spines. The
spines vary in colour; the base of the spine may be in one colour and its tip may have a dif-
ferent colour. The most common varieties in this species are green test with green spines or
green test with white spines. In between the spines are the pedicellaria (pincer-like struc-
tures), which are white. The body of this species may be up to 7 cm and spines up to 2 cm.
Food and feeding: It is largely herbivorous, feeding on the seagrass, Thalassia sp. Its tube
feet and spines also play a role in feeding, catching, and holding bits of debris that float
past. Its Aristotle’s lantern surrounds its mouth on its oral (under) surface, and its five teeth
help in the feeding.
Reproduction: In this species, breeding takes place at various times of the year in
different parts of its range. In Bermuda, the spawning period is short and seems to be
related to the phase of the moon. Eggs and sperm are liberated into the water column
and fertilization is external. The larvae are planktonic and are known as pluteus larvae.
They pass through several developmental stages before undergoing metamorphosis into
juvenile urchins.

Antineoplastic (antitumour) activity: The glycoproteins isolated from this species
yielded antineoplastic properties (Bhakuni and Rawat, 2005).
Antimicrobial activity: The coelomic fluid of this species has a wide variety of peptides
with antimicrobial activity. Among the nineteen fractions, thirteen fractions were against
Micrococus luteus, four against Pseudomonas aeruginosa, and three against both bacteria
and the fungus Cladosporium herbarum (Figueiredo et al., 2016).
Sphaerechinus granularis (Lamarck, 1816)
Common name(s): Purple spiny urchin

Global distribution: Eastern Atlantic and the Mediterranean Sea: from Ireland to the
English Channel (French coast) to Spain, into the entire Mediterranean, to Morocco and
Gulf of Guinea, including the islands of Azores, Madeira, Canary, and Cape Verde
Ecology: This temperate and subtropical, benthic species favours sheltered locations and
lives on rocks covered with seaweed (meadows of the seagrass Posidonia oceanica) or
gravelly substrates. It is usually found in depths of 2–130 m in more exposed locations.
S. granularis often covers itself with morsels of algae and shell fragments, which are held
in place by the tube feet and by the claw-like structures known as pedicellaria.
Biology
Description: It is a large sea urchin that is somewhat flattened dorsally. There are two dis-
tinct colour forms. The test is purple in both, but one has purple spines and the other white.
The spines are short and blunt, all the same length, and are arranged neatly in rows. This
species may grow up to 15 cm in diameter.
Food and feeding: It grazes on algae, especially encrusting coralline algae, seagrass
blades, and their epiphytic organisms and detritus.
Reproduction: Spawning in this species takes place at any time of year, but the peak
period is spring and early summer. Eggs and sperm are liberated into the water column,
where fertilization takes place. The larvae are planktonic. After several moults, the echi-
nopluteus larva settles and undergoes metamorphosis before developing into a juvenile.
Associated species: This species is found living in association with two other species of
sea urchins, Centrostephanus longispinus and Paracentrotus lividus.
Predators: It is preyed upon by the starfish Marthasterias glacialis and Luidia ciliaris.
Edible values: The gonads of this species are considered a delicacy in Italy, Provence, and
Catalonia.

Cell vitality and lysozyme and antioxidant activities: The coelomic fluid and coelomo-
cyte lysates of this species exhibited cell vitality and lysozyme and antioxidant activities.
While the coelomic fluid showed a cell vitality of 90.2%, the coelomocyte lysates showed
Lysozyme activity: (diameter of lysis = 9.4 ± 0.5 mm). Further, both coelomic fluid and
coelomocyte lysates showed antioxidant activity as detailed below (Stabili et al., 2018).
Antioxidant Activity of Coelomic Fluid and Coelomocyte Lysates

TEAC (nmol TEa/mL sample) ORAC (nmol TEa/mL sample)
Coelomocyte lysates 435.8 671.2
Coelomic fluid 64.4 46.1
Source: Stabili, L. et al., Evid. Based Compl. Alt. Med., Article ID 7891748, 8, 2018.
a TE = Trolox equivalent.
Antitumour activity: The acetylenic lipid compound, asterinic acid, isolated from this
species showed antitumour activity (Kilimnik et al., 2016).
Toxopneustes pileolus (Lamarck, 1816)
Common name(s): Flower urchin
Global distribution: Tropical Indo-West Pacific; north from Okinawa, Japan, to Tasmania,
Australia in the south; and west from the Red Sea and the East African coast, to Raratonga
in the Cook Islands in the east
Ecology: It inhabits coral reefs, seagrass beds, and rocky or sandy environments at depths
of up to 90 m. It may sometimes partially bury itself on the substrate.
Biology
Description: The body of adult flower urchins is equally divided into identical
segments around a central axis in multiples of five (pentaradial symmetry). Its rigid
shell (test) has five interambulacral segments separated from each other by five ambu-
lacral segments. The test is variegated in colouration, usually deep red and grey,
though there are rare instances of green and pale purple. Each ambulacral segment
is ornamented by a large purple zigzag pattern running along its length. Two rows of
tube feet emerge from the grooves on either side of each of the ambulacral segments
(for a total of ten rows). The mouth is centrally located in the bottom (oral) surface of
the test. It is surrounded by a ring of small plates overlaid by softer tissue known as
the peristome. These urchins are relatively large and can reach a maximum diameter
of about 15–20 cm.
Food and feeding: It feeds on algae, bryozoans, and organic detritus.
Predators: Flower urchins have few predators. They are known to be toxic to fish. One of
the few organisms capable of consuming flower urchins with no apparent adverse effects is
the predatory corallimorph Paracorynactis hoplites.
Reproduction: Flower urchins are dioecious (having separate male and female individu-
als), but it is almost impossible to determine the sex of an individual by external charac-
teristics alone. In males, the genital pores (gonopores) are generally short, cone-shaped,
and extrude above the body surface; in females they are usually sunken. All other external
features such as shape and size of the tests or colour of the spines are indistinguishable
between the two sexes.
Associated species: The commensal alpheid shrimp Athanas areteformis can sometimes
be found living among the spines of flower urchins. The intestines of these urchins can
also serve as habitats for the commensal flatworm Syndesmis longicanalis. Flower urchins
are also common hosts of the zebra crab, Zebrida adamsii. These tiny crabs are obligate
symbionts of sea urchins.
Others: This venomous species is considered highly dangerous, as it is capable of delivering
extremely painful and medically significant stings when touched.

Antibacterial activity: Spinochromes, viz. spinochrome B, two isomers of spinochrome
D, spinochrome E, one isomer of spinochrome A, echinochrome A, and spinochrome C
have been isolated from this species. The crude extracts of tests/spines of this species have
shown antibacterial activity against different bacterial species as detailed below (Brasseur
et al., 2017).
Antibacterial activity
EC50 and MIC—Bacterial Growth
E. coli B. subtilis V. aestuarianusa C. marinaa S. oneidensisa
Tests/Spines Crude Extracts (μg/mL)

>2000 512.60 >2000 >2000 >2000 EC50
>2000 >2000 >2000 >2000 >2000 MIC
a Vibrio aestuarianus; Cobetia marina; Shewanella oneidensis
Antioxidant activity: The crude extracts of this species showed DPPH antioxidant activity
with an EC50 value of more than 400 μg/mL (Brasseur et al., 2017).
Mitogenic and chemotactic activity: A lectin with biological activities such as mitogenic and
chemotactic characteristics has been described in the venom of the pedicellariae (Petzelt, 2005).
Mitogenic and cytotoxic activities: Two d-galactose-binding lectins (SUL-I and SUL-II)
have been purified from the globiferous pedicellariae of this species. Among them, SUL-I
had mitogenic activity and cytotoxic activity (Satoh et al., 2002).
Others: Suzuki-Nishimura et al. (2001) reported on the presence of lectin (which is a
causative factor for agglutination of RBCs) in the pedicellariae of this species. This lectin
has also been reported to inhibit histamine (which serves as an important role in our body’s
immune response) release dose dependently.
Sakai et al. (2013) reported that the sea urchin lectin-III isolated from the pedicellariae of
this species agglutinated the rabbit erythrocytes besides inducing mitogenic stimulation on
murine splenocytes.
Takei et al. (1991) reported that the toxic substance produced by this species induced
histamine release from rat peritoneal mast cells.
Two active toxins, viz. contractin A and peditoxin, have been purified from the pedicellarial
venom of this species. The first toxin was found to interfere with the transmission of signals
at nerve endings as well as cause hemagglutination (clumping of the red blood cells). At
low doses to mice, the second toxin, pedoxin, was found to result in markedly lower body
temperatures, muscle relaxation, sedation, and anaesthetic coma. At higher doses, it resulted in
convulsions and death. In addition to these toxins, lectins, SUL-I, SUL-II, SUL-IA, and SUL-
III (SUL stands for “sea urchin lectin”) have also been isolated from this species. These lectins
may be valuable as research tools for investigating the functions of cell processes Edo (2014).
Tripneustes gratilla (Linnaeus, 1758)

Common name(s): Collector urchin

Global distribution: Indo-Pacific, Hawaii, the Red Sea, and the Bahamas
Ecology: Mature collector urchins prefer open sea bottoms with some cover, but the young
prefer rocky areas for concealment; depth range is 2–30 m.
Biology
Description: These urchins are dark in colour, usually bluish purple with white spines.
The pedicles are also white, with a dark or black base. The spines of this species may be
orange, orange-tipped, or white. Some specimens are wholly orange, while those of others
are only orange-tipped or completely white. Collector urchins reach up to 15 cm in size.
Food and feeding: The diet of this species includes algae, periphyton, and seagrass. Most
collector urchins prefer seagrass species, viz. Thalassodendron ciliatum and Syringodim
isoetifolium. Unlike some other sea urchins, collector urchins graze continually, day and
night. They graze near the substrate.
Edible/commercial values: Collector urchins are economically important in some parts of
the world. They are edible and sometimes exploited by humans. The species has ecological
value and prospects as a biological control agent. It is considered as the commercially
traded sea urchin (Anon, 2018).
Others: Ciguatera poisoning has been reported following the consumption of this species in
Anaho Bay (Nuku Hiva Island, Marquesas archipelago, French Polynesia). The ciguatoxin
(CTX) bioaccumulation in this species as a cause of ciguatera-like poisoning is due to the
abundance of toxic dinoflagellate, Gambierdiscus sp. in this region (Darius et al., 2018).

have been isolated form this species (Brasseur et al., 2017). The crude extracts of tests/
spines of this species have shown antibacterial activity against different bacterial species as
detailed below (Brasseur et al., 2017).
Antibacterial Activity
E. coli B. subtilis V. aestuarianus C. marina S. oneidensis
>2000 1227.00 >2000 543.80 >2000 EC50
>2000 >2000 >2000 >2000 >2000 MIC
Source: Brasseur, L. et al., Mar. Drugs, 15, 179, 2017.
Methanolic extract of the guts and gonad of this species showed antibacterial activity
(Ambag et al., 2016).
Antimicrobial activity: Abubakar et al. (2012) reported that the methanol and chloroform
extracts of the gonad and gut of this species showed antimicrobial activity against
Salmonella typhi, Escherichia coli, Shigella sonnei, Pseudomonas aeruginosa, and
Penicillium spp. The values of zone of inhibition (dia., mm) for the different species of
microbes in both the said extracts are detailed below.
Antimicrobial Activity of the Sea Urchin (T. gratilla) Methanol Extracts

Zone of Inhibition (mm)
Organ/Tissue Extract E. coli S. aureus S. sonnei S. typhi P. aeruginosa P. sppa
Gonads 16 13 11 18 18 22
Guts 16 14 13 18 20 20
Source: Abubakar, L., et al., Afr. J. Pharm. Therap., 1, 19–23, 2012.

a Penicillium spp.
Antimicrobial Activity of the Sea Urchin (T. gratilla) Chloroform Extracts

Zone of Inhibition (mm)
Organ/Tissue Extract E. coli S. aureus S. sonnei S. typhi P. aeruginosa P. sppa
Gonads 8 9 11 9 8 8
Guts 8 7 8 9 7 6
Source: Abubakar, L., et al., Afr. J. Pharm. Therap., 1, 19–23, 2012.

a Penicillium spp.
Antioxidant activity: The crude extracts of this species showed DPPH antioxidant activity
with an EC50 value of more than 400 μg/mL (Brasseur et al., 2017).
Cytotoxic activity: The compound epidioxysterol isolated from this species showed mild
cytotoxicity against three human tumour cell lines, viz. SGC-7901, HepG2, and HeLa cells
with IC50 values of 99, 65, and 94 mg/mL, respectively (Liu et al., 2011).
Tripneustes ventricosus (Lamarck, 1816)
Common name(s): West Indian sea egg, white sea urchin

Global distribution: It is found in the western Atlantic Ocean, the Caribbean Sea, and
Gulf of Mexico; its range extends from Bermuda, the Carolinas, and Florida to Belize,
Venezuela, and Brazil and also includes the west coast of Africa and Ascension Island.
Ecology: It is found at depths of less than 10 m in seagrass meadows, in rubble areas, and
on shallow rocky reefs. Young sea urchins conceal themselves in crevices and under rocks
during the day, but larger individuals stay out in the open.
Biology
Description: The test of this species is dark, black, dark purple, or reddish brown and has
white spines of 1–2 cm long. It is often covered with pieces of seagrass, fragments of shell,
and other debris. These decorations are held in place by tube feet among the spines and are
believed to provide protection from the intense sunlight that penetrates the shallow water.
The test can reach 10–15 cm in diameter.
Food and feeding: It feeds on algae but tends to avoid the crustose, highly calcified
coralline algae.
Reproduction: Ripe gonads are found in urchins at any time of year, but breeding prob-
ably takes place mostly in the summer. Male and female urchins liberate gametes into the
sea where fertilization takes place. The eggs soon hatch into larvae that are planktonic.
These develop through a number of larval stages over the course of about one month before
settling on the seabed and undergoing metamorphosis into juveniles.
Predator: The queen triggerfish, (Balistes vetula), is the main predator.

against Plasmodium falciparum FcB1 strain, and the recorded IC50 value was 84.8 µg/mL
Anticancer activity: The clarified extracts of this species showed anticancer activity
against 3LL and PC3 cancer cell lines with IC50 values of 39.9 and 77.0 µg/mL, respectively
4.8 FAMILY: SCUTELLIDAE (ORDER: CLYPEASTEROIDA)

Scaphechinus mirabilis (A. Agassiz, 1864)
Common name(s): Sand dollar

Global distribution: Northwest of the Pacific Ocean: From southern Japan to the Aleutian
Islands
Ecology: In the intertidal zone, it is seen in the sandy bottom of shallow coasts of less than
25 m in depth.
Biology
Description: It is disc shaped with about 8 cm in diameter. On the back of the shell there
are five radiant flower patterns covered with dark purple, short spines. The groove of the
abdominal surface extends radially around the mouth branches into a vein. There are many
variations in the shape of the trailing edge of the shell.
Food and feeding: It largely feeds on the diatoms.
Reproduction: Under laboratory conditions, the development of larvae of this spe-
cies took 28.5–29 days from fertilization to settling and the end of metamorphosis at a
temperature 20°C and salinity 32.2–32.6‰. The free-swimming ciliary blastula appears
from the egg membrane and attains pluteus I stage with one pair of arms after about
40 hours. In 4.5–5 days, the pluteus II stage occurs with three pairs of arms, and in
9 days the pluteus III stage with four pairs of arms are formed. On days 28-29 of develop-
ment, the larvae settle. Most of the larvae complete their metamorphosis in 5 hours after
settling.

Antioxidant, antibacterial, antifungal, and antitumour activities: quinone pigments,
viz. echinochrome A and spinochrome E produced by this species, displayed effective
antioxidant, antibacterial, antifungal, and antitumour activities (Ageenko et al., 2014).
Cardioprotective effects: The naphthoquinoid pigment, echinochrome A (Ech A) of

this species, is already known for its antioxidant, antimicrobial, anti-inflammatory,
and chelating abilities. Now, it has been reported that this compound also possesses
cardioprotective effects against toxic agents that induce death of rat cardiac myoblast
H9c2 cells. Co-treatment with Ech A prevented the decrease in mitochondrial membrane
potential and increase in ROS level. Co-treatment of Ech A also reduced the effects of these
cardiotoxic agents on mitochondrial oxidative phosphorylation and adenosine triphosphate
level. These findings indicate the therapeutic potential of Ech A for reducing cardiotoxic
agent-induced damage (Jeong et al., 2014).
Mischenko et al. (2005) reported on the occurrence of two spinochromes, echinamines
A and B, from this species. Pokhilo et al. (2006) described the total synthesis of two marine
aminated hydroxynaphthazarins, echinamines A (3-amino-7-ethyl-2,5,6,8-tetrahydroxy-
1,4-naphthoquinone) and B (2-amino-7-ethyl-3,5,6,8-tetrahydroxy-1,4-naphthoquinone), and
amino analogues of spinazarin and spinochrome D produced by this species.
A new sulfolipid, sulfoquinovosyldiacylglycerin (SQDG), has been isolated form this

species (Logvinov et al., 2012).
Acetylcholinesterase (AChE) inhibiting and antioxidant activities: Echinochrome
A (EchA), a dark-red pigment of the polyhydroxynaphthoquinone class isolated from this
species, inhibited Acetylcholinesterase (AChE) with an irreversible and uncompetitive
mode. In addition, EchA showed reactive oxygen species scavenging activity, particularly
with nitric oxide. These findings indicate new therapeutic potential for EchA in treating
reduced acetylcholine-related diseases including Alzheimer’s disease, and they provide an
insight into developing new AChE inhibitors (Lee et al., 2014).
Others: Pelageev and Anufriev (2016) described the synthesis of mirabiquinone
A (2,3,5,6,8,10,11,13-octahydroxy-7-methyl-1H-dibenzo[b, h]xanthene-1,4,9,12(7H)-tetra-
one), and methyl analogues of mirabiquinone A produced by this species.
Isai et al. (2007) reported on the isolation of eicosapentaenoic acid (20:5) from the scalp
lipids of this species.
4.9 FAMILY: DIADEMATIDAE (ORDER: DIADEMATOIDA)

Diadema savignyi (Audouin, 1829)
Common name(s): Long-spined sea urchin, black longspine urchin, banded diadema
Global distribution: It is native to the east coast of Africa, Red Sea, Indian Ocean, and
western Pacific Ocean.
Ecology: It is typically found on mixed sandy, rocky, and coral substrates, especially in
reefs and in shallow lagoons disturbed by storms or by other natural causes. Its depth range
is from the surface down to about 70 m.
Biology
Description: It has a spherical, slightly flattened test that is up to 9 cm in diameter. Its brit-
tle, thin, hollow spines grow in tufts and can be as long as 25 cm. Individuals of this spe-
cies are usually black but can also be grey, dark brown, or purple. They may be banded
with lighter and darker shades in juveniles. This species can be distinguished from other
related species by its iridescent green or blue lines in the interambulacral areas and around
the periproct, a cone-shaped region surrounding the anus. In a small number of individuals
there are pale coloured spots at the aboral (upper) ends of the interambulacrals.
Food and feeding: It feeds on the algal mat that grows over the surface of seabeds.
Associated species: Certain small fish such as cardinal fish, flatworms, and shrimps some-
times seek protection from predators among the long spines.
Predators: It is preyed on by pufferfish (Tetraodontidae) and porcupinefish (Diodontidae),
lobsters, and snails. It reacts to a shadow falling on it by angling its spines towards the pos-
sible attacker.
Reproduction: Breeding of this species occurs mainly during the northeast monsoon
period, peaking in May. This species spawns just after the full moon, on days seventeen to
eighteen of the lunar cycle.
Ecological importance: Diadema savignyi is nocturnal and tends to hide in crevices or
under boulders during the day, or several individuals may huddle together in the open. The
urchins disperse at dusk to feed. In the course of tearing up the mat, the urchin also abrades
the underlying surface, causing bioerosion. Its activities help control the algae, which oth-
erwise might overwhelm the corals.

have been isolated from this species (Brasseur et al., 2017). The crude extracts of tests/
spines of this species have shown antibacterial activity against different bacterial species
as detailed below (Brasseur et al., 2017).
Antibacterial Activity
E. coli B. subtilis V. aestuarianus C. marina S. oneidensis
58.70 252.40 534.80 681.00 391.00 EC50
406.80 1219.00 >2000 >2000 >2000 MIC
Source: Brasseur, L. et al., Mar. Drugs, 15, 179, 2017.
Antioxidant Activity: The crude extracts of this species showed DPPH antioxidant activ-
ity with an EC50 value of 34.5 μg/mL (Brasseur et al., 2017).
Others: The fatty acids from the body and egg of this species have been reported to possess
positive effect on human health. The essential lipid ingredients include wax (H + W), tria-
cylglycerol (TG), monodiacylglycerol MDAG, free fatty acid (FFA), sterol (ST), and polar
lipid (PL). Palmitic acid (16:0) accounted for the largest proportion of saturated fatty acids.
Beside this, eicosenoic (20:1n-9) is a dominant fatty acid in the monounsaturated fatty acid
group (MUFA) group. In the polyunsaturated fatty acid group (PUFA), arachidonic acid
(20:4n-6) made up the highest proportion, accounting for over 50%. Furthermore, four
omega-3 fatty acids including eicosapentaenoic fatty acid (C20:5n-3, EPA), a very valuable
fatty acid, are also present in the body and egg of this species. The PUFA/SFA ratio and the
n3/n6 ratio present in the total lipid contents of the egg and the body of this species have
been found to meet the WHO standards for healthy food (Kim et al., 2018).
Antimicrobial and antioxidant properties: The methanolic shell extract of this species
showed promising inhibitory effect against two gram-negative bacteria, Escherichia coli
(inhibition zone dia. value, 16.3 mm) and Acinetobacter spp. (inhibition zone dia. value
11.1 mm). On the other hand, the methanolic gonadal extract of this species inhibited
gram-positive bacteria, Staphylococcus aureus, with an inhibition zone value of 17.1 mm.
Further, the non-freeze-dried methanolic shell extract of this species revealed excellent
DPPH scavenging activity with an IC50 value of 3.8 µg/mL (Tee et al., 2017a).
Diadema setosum (Leske, 1778) [1]
Common name(s): Long-spined sea urchin

Global distribution: Throughout the Indo-Pacific region, from Australia and Africa to
Japan and the Red Sea
Ecology: It is commonly associated with rocky coral reefs but is also found on sand flats
and in seagrass beds.
Biology
Description: The test of this species is spherical and black in colour. However, the body
is not perfectly spherical. It is a typical sea urchin, with extremely long, hollow spines that
are mildly venomous. These spines are often black but sometimes they are brown-banded.
This species differs from other Diadema with five characteristic white dots that can be
found on the animal’s test, strategically located between the urchin’s ambulacral grooves.
Food and feeding: It is a prolific grazer and is known to feed on a variety of algal species
common on tropical coral reefs.
Predator: The primary predator of this species is the blackspot tuskfish (Cheorodon
schoenleinii).
Reproduction: This species has been known to spawn both seasonally and year-round
depending on the location of the spawning population. It has been suggested that its
populations are temperature-dependent in their spawning seasonalities. This species has
been found to trigger spawning events in concordance with the appearance of a full moon.
Others: Like other venomous sea urchins, the venom of this species is only mild and not at
all fatal to humans. The toxin mostly causes swelling and pain, and it gradually diffuses
over several hours. More danger is presented by the delivery system—the urchin’s spines.

Antibacterial activity: The methanol extracts of the ovary of this species have shown
potential antimicrobial properties against the gram-negative bacteria (Salmonella typhi,
Salmonella typhimurium, Shigella flexneri, Pseudomonas aeruginosa, Aeromonas
hydrophila, Acinetobacter sp., Citrobacter freundii, and Klebsiella pneumoniae) and
gram-positive bacteria (Bacillus subtilis, Staphylococcus epidermidis, and Staphylococcus
aureus) (Marimuthu et al., 2015).
Antibacterial and antioxidant activities: The organic extracts of this species displayed
antibacterial activity against Escherichia coli and Staphylococcus aureus with inhibition
zone values of 16.3 and 17.1 mm, respectively. Further, these extracts also showed DPPH
antioxidant activity with half maximal inhibitory concentration of 3.8 µg/mL (Tee et al.,
2017b).
Immune-nutrients: The extracts of the gonad of this species were found to have steroid,
amino acids, and antioxidant compounds as well as vitamin E, which serve as immune nutri-
ents. Owing to this, this species may be a potent, alternative food source (Salma et al., 2016).
Cytotoxic activity: The compounds, 5,8-Epidioxycholest-6-en-3-ol (1), cholesterol (2), glyc-
erol 1-palmitate (3), and glycerol 1,3-dioleate-2-stearate (4) have been isolated from the meth-
anol extract of this species. Among these compounds, compound 1 was found to have strong
cytotoxic effect against various cancer cell lines, such as KB (human epidermoid carcinoma),
(IC50, 2.0 µ/mL), FL (fibrillary sarcoma of the uterus), (ICso, 3.93 µ/mL), and Hep-2 (human
hepatocellular carcinoma cells (IC50, 2.4 µ/mL) by in vitro assay (Minh et al., 2004).
Echinothrix diadema (Linnaeus, 1758) [1]
Common name(s): Diadema urchin, blue-black urchin, horned sea urchin, long-spined urchin
Global distribution: Throughout all the Indo-Pacific coral reefs, from the Red Sea to Hawaii
Ecology: This tropical, reef-associated species occurs in shallow coral and coral rubble areas
at depths of 1–40 m. It is active at night, hiding in crevices or under rocks during the day.
Biology
Description: It is a long-spined urchin. With its spines, its typical diameter is 10–20 cm.
It is generally black or blue-black in colour and always dark (spines show a blue sheen in
the light). The spines, which are not banded, are closed at the tip.
Associated species: It hosts commensal species such as the shrimp Stegopontonia
commensalis. Saron marmoratus stays close for protection, like many fish of the families
Apogonidae (cardinal fish) and Centriscidae (razorfish and relatives).
Food and feeding: It is known to graze on organic material, and adults may also feed on
live hard corals. It is a generalist herbivore, displaying nocturnal feeding behaviour.

Cytotoxic and contraceptive properties: Pokhilo et al. (2014, 2015) described the synthesis
of pyranonaphthazarin pigments, 6,8,9-Trihydroxy-2-methyl-2H-naphtho[2,3-b]pyran-5,10-
dione, and its analogues 6,9-dihydroxy-2-methyl-2H-naphtho[2,3-b]pyran-5,10-dione and
6,9-dihydroxy-2-methyl-7,8-dichloro-2H-naphtho[2,3-b]pyran-5,10-dione produced by this
species. These compounds displayed cytotoxic and contraceptive properties.
Others: Moore et al. (1968) reported on the presence of the polyhydroxynaphthoquinone
pigment, 2-methyl-8-hydroxy-2H-pyrano [3,2-g] naphthazarin from this species.
4.10 FAMILY: LOVENIIDAE (ORDER: SPATANGOIDA)

Echinocardium cordatum (Pennant, 1777)
Common name(s): Sea potato

Global distribution: This species has a discontinuous cosmopolitan distribution. It is
found in temperate seas in the Adriatic Sea, the North Atlantic Ocean, the west Pacific
Ocean, around Australia, New Zealand, South Africa, and the Gulf of California.
Ecology: It is found in the subtidal regions in temperate seas around the world and lives
buried in the sandy seafloor at depths of down to 230 m.
Biology
Description: The sea potato is a heart-shaped urchin clothed in a dense mat of furrowed
yellowish spines that grow from tubercles and mostly point backwards. The upper surface
is flattened and there is an indentation near the front. This urchin has a fawn colour but its
test (without spines) may be white. During life, these spines are said to trap air, which helps
prevent asphyxiation for the buried urchin. The ambulacrum forms a broad furrow in a star
shape extending down the sides of the test. There are two series each of two rows of tube
feet. The test reaches a size of 6–9 cm in length.
Food and feeding: This species feeds mainly on organic debris.
Behaviour: The sea potato buries itself in sand to a depth of 10–15 cm. It makes a respira-
tory channel leading to the surface and two sanitary channels behind itself, all lined by a
mucus secretion. The location of burrows can be recognized by a conical depression on the
surface in which detritus collects.
Reproduction: In this species, the sexes are separate and the males and females both lib-
erate gametes into the water table in the spring. The echinoplutei larvae that develop after
fertilization have four pairs of arms and are laterally flattened. The larvae are pelagic and
form part of the zooplankton. Metamorphosis takes place in 39 days after fertilization,
with the larvae settling out and burrowing into the substrate. The life span of this species
is thought to be about 10 years.
Associated species: In the sandy seabed, this species is often found in association with
the bivalve molluscs Tellina fibula, Ensis ensis and Venus striatula.
The bivalve Tellimya ferruginosa is often found living inside the sea potato’s burrow as
a commensal organism. Another species that makes use of the burrow is the amphipod
crustacean, Urothoe marina

Antitumour and antibacterial activities: Alkanesulfonic acid and its derivatives of this
species have shown antitumour activity and antibacterial activity against Rhodospirillum
solexigens SCR 113 (Kim, 2015).
Others: Thiosulfonic acids, viz. hedathiosulfonic acids A and B, have been isolated from
this species. Their activities are, however, to be known (Kita et al., 2002).
4.11 FAMILY: SCHIZASTERIDAE (ORDER: SPATANGOIDA)

Brisaster latifrons (Agassiz, 1898)
Common name(s): Northern heart urchin, wide heart sea urchin
Global distribution: Eastern Pacific: Alaska to the United States.
Ecology: This temperate, benthic species lives in deeper waters; depth range is 20–1800 m.
Biology
Description: Test of this species is cordiform, with deep anterior sulcus. Its posterior face
is truncated. It has the dimensions of 41 × 41 × 25 mm.
Associated species: This species serves as a host for the commensal epibiont Waldo
arthuri, a galeommatid clam.

Anti-inflammatory effects: Tanzawaic acid derivative, 2E,4Z-tanzawaic acid D isolated
from the associated Penicillium sp. SF-6013 of this species showed anti-inflammatory
effects in lipopolysaccharide (LPS)-activated microglial BV-2 cells by inhibiting the pro-
duction of nitric oxide (NO) with an IC50 value of 37.8 μM (Jin et al., 2016).
4.12 FAMILY: GLYPTOCIDARIDAE (ORDER: STOMOPNEUSTOIDA)

Glyptocidaris crenularis (A. Agassiz, 1864)
Global distribution: Japan
Ecology: It lives in crevices in tropical rocky shores and coral reef habitats.
Biology
Description: Test of this species has a subconical in profile. Apical disc is small and hemi-
cyclic. Periproct is oval. Ambulacra are straight.
Food and feeding: It feeds on marine algae, especially Caloglossa leprieurii and Sargassum
pallidum.

Zhou et al. (2010a) reported on the isolation of compounds, viz. homarine, halicerebroside A,
cholesterol sulfate, pyropheophorbide-a, saropeptate, aurantiamide acetate, and cholesterol
from this species. Activities of these compounds are, however, to be known.
Zhou et al. (2010b) also reported on the presence of compounds, viz. N-Acyl taurine and
5α,8α-epidioxycholest-6-en-3β-ol, hypoxanthine, 1-(β-d-ribofuranosyl)-1,2,4-triazole
from this species.
Castro et al. (2009) reported on the presence of a unique 2-Sulfated β-Galactan from the
egg jelly of this species.
4.13 FAMILY: STOMOPNEUSTIDAE (ORDER: STOMOPNEUSTOIDA)

Stomopneustes variolaris (Lamarck, 1816)
Common name(s): Black sea urchin, long-spined sea urchin

Global distribution: Throughout the tropical Indo-Pacific; from the east African coast to
the Philippines; particularly abundant in Sri Lanka
Ecology: This species shows a remarkably patchy distribution. It is found on rock and
damaged reefs, most often in shallow waters but never too close to wave action.
Biology
Description: It is a rather big and strong sea urchin. Its spines are robust and sharp. Colour
of the spines is black with sometimes a blue-greenish tinge (depending on the light).
The individuals can be recognized by their five grey and zigzag sutures on the upper face.
The oral face is clearer. The juveniles of this species may be black or pale brown and they
often show strikingly asymmetrical spines, due to their habit to use them for digging hid-
ing holes in soft rocks.

Antiandrogenic, antioxidant, hypocholesterlemic, and nephroprotective properties:
The crude ethanolic extract of this species containing the ethyl ester compound, hexadeca-
noic acid, exhibited antiandrogenic, antioxidant, hypocholesterlemic, and nephroprotec-
tive properties (Devi and Selvaraj, 2015).
Hypocholesterolemic and antioxidant properties: The 12 hydroxy-, methyl ester
compound, 9-Octadecenoic acid, and the bis (trimethylsilyl) ester compound propane-
phosphonic acid have shown hypocholesterolemic and antioxidant properties. The latter
compound may find use in the treatment of atherosclerosis (Devi and Selvaraj, 2015).
Others: Devi and Selvaraj (2015) also reported on the presence of compounds, viz. dimethyl
sulfoxide, tetradecanoic acid, and 2,3-Dihydroxy propyl elaidate from this species.

The acetylenic lipid compounds, viz. pectenolone and 4-keto-cynthiaxanthin isolated from
certain unidentified species of sea urchins, displayed cytotoxicity (Kilimnik et al., 2016).
5 Biology and Ecology
of Pharmaceutical Sea
Cucumbers (Class:
Holothuroidea)
5.1 FAMILY: SYNAPTIDAE (CLASS: HOLOTHUROIDEA; ORDER: APODIDA)

Synapta maculata (Chamisso & Eysenhardt, 1821)
Common name(s): Spotted worm sea cucumber, snake sea cucumber, Vietnamese sea
cucumber
Global distribution: Tropical Western Indo-Pacific Ocean
Ecology: This shallow water species occurs at depths down to about 20 m on reefs and on soft
sediments on the seabed among seagrasses and seaweeds. It can also bury itself under rubble.
Biology
Description: It is one of the longest sea cucumbers in the world with a size of about 3 m. It
is a long, slender sea cucumber with 15 tentacles. Its colouring is variable, being some shade
of yellowish brown with wide longitudinal stripes and patches of darker colour. The spicules
243
(microscopic calcareous spike-like structures that support the body wall) of this species are
large and shaped like anchors and are used in locomotion; they measure 2 mm long.
Food and feeding: It mainly feeds on seagrass. The tentacles of this species surround
the mouth and are used in surface feeding. The tentacles are in continuous motion; they
flatten themselves against the substrate or seagrass leaf blades and collect food particles by
adhesion, then bend inwards until the tips are in the mouth, where the food is scraped off
by the buccal sphincter muscle.

Anticancer activity: Among the triterpene holostane glycosides, synaptosides A and A1,
isolated from this species, synaptoside A showed moderate cytotoxic activity (IC50 8.6 µg/
mL) against HeLa tumour cells (Avilov et al., 2008; Mondol et al., 2017).
5.2 FAMILY: CUCUMARIIDAE (ORDER: DENDROCHIROTIDA)

Actinocucumis typica (Ludwig, 1875)
Global distribution: Tropical Indo-West Pacific
Ecology: This benthic, reef-associated species occurs at depths of 3.5–15 m.
Biology
Description: This dendrochirotid species has uniform brown colour or yellowish brown
with some red patches and fine black flecking, or pale brown to cream with fine brown
flecking on the body and larger brown patches on the tube feet. The body of this species is
pentagonal in section with raised radial ridges and five thick oral valves. There are about
20 dendritic tentacles, which are variable in arrangement and size. The ventral pair is
the smallest. Tube feet are confined to radii. Ossicles of the body wall are abundant and
crowded. Spicules of the body wall are tables and buttons. Buttons look like figures of
eight-shaped fenestrated ellipsoids and measure 0.4–0.5 mm. Tables are irregular.
Biology and Ecology of Pharmaceutical Sea Cucumbers (Class: Holothuroidea) 245

Antifungal, haemolytic, and cytotoxic activities: Minor triterpene glycosides—
typicosides A1 (1), A2 (2), B1 (3), C1 (4), and C2 (5)—along with two known glycosides,
intercedenside A and holothurin B3, have been isolated from this species. The first five
glycosides showed antifungal activity against Aspergillus niger, Fusarium oxysporum, and
Candida albicans as detailed below. Further, glycosides 2–5 displayed cytotoxic activity
against mouse spleen lymphocytes and mouse Ehrlich carcinoma cells (ascite form), as
well as haemolytic activities against mouse erythrocytes (Silchenko et al., 2013).
Antifungal Activity at Concentration of 100 µg/mL (Zone of Inhibition, mm)

Aspergillus niger Fusarium oxysporum Candida albicans
Typicoside A1 (1) 6.2 4.0 0
Typicoside A2 (2) 8.0 6.0 2.3
Typicoside B1 (3) 5.0 4.0 1.2
Typicoside C1 (4) 0 0 0
Typicoside C2 (5) 1.8 0 0
Source: Silchenko et al., 2013a.
Cytotoxic activity against mouse spleen lymphocytes and ascites form of mouse Ehrlich
carcinoma and haemolytic activity against mouse erythrocytes for typicosides (1–5)
Haemolytic Activity
(ED50, µg/mL) Cytotoxic Activity (ED50, µg/mL)
pH 6.0 pH 7.4 Mouse Spleen Lymphocytes Ehrlich Carcinoma Cells
Typicoside A1 (1) 0.12 0.25 1.7 3.0
Typicoside A2 (2) 0.15 0.29 1.2 1.3
Typicoside B1 (3) 0.11 0.33 3.0 4.5
Typicoside C1 (4) 4.80 6.25 48.0 72.0
Typicoside C2 (5) 0.09 0.18 2.6 1.7
Source: Silchenko et al., 2013.
Lysosomal activity and ROS formation: The triterpene glycosides—typicosides A1, A2,
B1, C1, and C2—have been reported to stimulate lysosomal activity and ROS formation in
mouse peritoneal macrophages (Pislyagin et al., 2014).
Immunomodulatory activity: The typicoside C1 of this species demonstrated a strong
immunostimulatory effect on mouse peritoneal macrophages in vitro (Pislyagin et al., 2014).
Aslia lefevrei (Barrois, 1882) (= Cucumaria lefevrii)
Common name(s): Brown sea cucumber

Global distribution: Mediterranean, Atlantic, English Channel, and North Sea,
English Channel (western and southern coasts of Great Britain, Ireland, French coasts);
Northwest Atlantic from Brittany to Portugal; and Mediterranean
Ecology: It is found on rocky bottoms, in crevices, from the infralittoral, from the first
meters to about 50 m of depth.
Biology
Description: The body of this species is shaped like a gherkin. It is brownish to black and
leather-like with a length of 15 cm. Often hidden in the crevices of the rock, this species reveals
only its 10 tentacles (8 long and 2 short) which are ramified; they are brown orange to grey
black and 10 cm long. These can retract inside the body of the animal when it is disturbed.
Food and feeding: It is a suspensivore filter. It captures food particles suspended in water.
The tentacles are licked one after the other in a definite order, specific to the individual.
Reproduction: Reproduction is sexual. The sexes are separated. Fertilization is external.

The egg laying takes place from late February to early April. Larvae are barrel shaped,
with short pelagic life, and settle in late March to early April.

The compounds lefevreiosides A1, A2, C, and D have been isolated from the aqueous-
alcoholic extracts of this species (Rodriguez and Riguera, 1989; Mondol et al., 2017). The
activities of these compounds are to be known.
Bahrami and Franco (2016) reported on the presence of lefevreiosides A1, A2, B, and C
from this species.
Athyonidium chilensis (Semper, 1868)
Common name(s): Burrowing shaggy sea cucumber

Global distribution: Southeast Pacific coast
Ecology: This shallow-water species occurs in intertidal boulder beaches and in the
Macrocystis zone.
Biology
Description: These are large forms (25 cm long) with thick, soft skin and numerous stout
feet. Tentacles consist of five large external pairs and five small inner pairs. There is a large
dorsal stone canal, which is often branched. Further, there are one or two tufts of smaller stone
canals with minute heads. It is the most economically important holothuroid. Colour of the
individuals is greyish mottled to almost black. While the ventrum is paler, tentacles are dark.
Food and feeding: These individuals are capable of active feeding on microalgae associ-
ated with the sediment.
Reproduction: This species has shown continuous gametogenesis, and spawning individuals
could be found throughout the year. However, spring was the main reproductive time. Males
reached sexual maturity at a smaller size than females (males: 21.2 g, females: 43.7 g eviscerated
weight), and mature females showed a high mean absolute fecundity (Peters-Didier et al., 2018).
Fisheries and aquaculture: It seems that this species has a low commercial value,
even though the interest in monospecific fisheries in cold waters has increased steadily.
Nevertheless, its aquaculture potential is interesting due to the fact that it could be the
source of a natural product, or, in great abundance, could be used as a biological filter of
organic particles in the discharges from fish farms.
Antibacterial, antifungal, haemolytic, and cytotoxic activities: Two saponins (includ-
ing holothurinoside D) have been isolated from this species. The methanolic extract of this
species showed antibacterial, antifungal, and haemolytic activities. The cytotoxic activity
of this extract was on a neuroblastoma cell line, N2A. A dose-depending reduction in the
cell viability was detected in this regard with an IC50 of 77.3 μg mL−1. The antimicrobial
activity of this extract was against gram-positive bacteria such as S. aureus and B. subtilis,
with a zone of inhibition of 22 and 19 mm, respectively. Antifungal activity of the extract
was with Aspergillus sp., Botrytis sp., and C. albicans, with a zone of inhibition of 24.5,
13.5, and 22 mm, respectively (Sottorff et al., 2013).
Anticoagulant activity: This species synthesized a fucosylated chondroitin sulphate,

which showed anticoagulant activity similar to heparin (Matsuhiro et al., 2012).
Fatty acids: Phospholipids were the major lipid contents of the ethanolic extracts of
tubules, internal organs, and body wall of this species. Saturated fatty acids predomi-
nated in tubule phospholipids (40.69%), while in internal organs and body wall phos-
pholipids, the monounsaturated fatty acids were in higher amounts (41.99% and 37.94%,
respectively). The main polyunsaturated fatty acids in phospholipids were C20: 2ω-6,
arachidonic (C20: 4ω-6), and eicosapentaenoic (C20: 5ω-3) acids. These results demon-
strate that A. chilensis is a valuable food for human consumption in terms of fatty acids
(Careaga et al., 2013).
Cercodemas anceps Selenka, 1867 (= Colochirus anceps)
Common name(s): Red box sea cucumber, pink warty sea cucumber, spiny sea cucumber,
pink sea cucumber, pink, and green sea cucumber
Global distribution: Indo-West Pacific Ocean: East Africa and Madagascar to Malaysia
to Northern Australia
Ecology: This tropical, benthic species lives among seagrasses, clinging to tubeworm tubes
or other hard surfaces; sometimes it is half buried in sediments. Its depth range is 0–50 m.
Biology
Description: The body of this species is short and somewhat angular in cross section. It
has an underside with three rows of tiny tube feet. The upper side has pink warty bumps
on a pink or yellow background. It has length of 6–8 cm.
Food and feeding: It is a suspension feeder, feeding largely on phytoplankton.

Cytotoxic activity: Seven holostane-type triterpene saponins including five new compounds,
namely cercodemasoides A-E, have been isolated from this species. All these compounds
showed potent cytotoxicity on five human cancer cell lines including Hep-G2 (hepatoma
cancer), KB (epidermoid carcinoma), LNCaP (prostate cancer), MCF7 (breast cancer), and
SK-Mel2 (melanoma), with IC50 values ranging from 0.03 to 7.36 μM (Cuong et al., 2015).
Antiangiogenic and cytotoxic activities: The organic extracts of this species exhibited
strong cytotoxicity against human tumour cancer cells and potent antiangiogenic activity
(Sima and Vetvicka, 2011).
Antitumour activities: The triterpene glycoside, colochiroside A, of this species exhibited

significant cytotoxic activity against six types of cultured tumour cell lines of p388, HL60,
A-549, SpC-A4, MKN-28, and SGC-7901 with the mean IC50 value of 3.6 mg × L(-1)
(Zhang and Yi, 2011).
Colochirus quadrangularis (Troschel, 1846) (= Pentacta quadrangularis)
Common name(s): Thorny sea cucumber
Global distribution: It is found in tropical parts of the Indo-West Pacific; its range extends from
East Africa and Madagascar to Malaysia and Northern Australia, Singapore, and Australia.
Ecology: This benthic species is usually found on the seabed of shallow seas where it clings
to seagrass, tube worm cases or other projections with its tube feet; depth range is 0–115 m
Biology
Description: It is a moderate-sized sea cucumber growing to about 10 cm in length. Its roughly
cylindrical body has four longitudinal ridges, giving it a square cross section and a flat base. It
has irregular, thorn-like, soft projections called papillae that lie along these ridges. The leathery
body wall is reinforced by calcareous spike-like structures that include basket-shaped spicules
and perforated ellipsoids. There is a ring of large, branched feeding tentacles around the mouth.
There are three rows of red tube feet on the underside, and the body tapers at the posterior end.
The anus is surrounded by five tooth-like projections. The colour of this species is mainly grey
often with pink on the ridges and thorns, and with yellow or reddish tentacles.
Food and feeding: It is a suspension feeder, rearing up its anterior end and spreading its
feathery tentacles to catch phytoplankton and other organic particles. The tentacles are
then retracted one by one and the mouthparts scrape off the food particles.
Associated species: It often has transparent, and almost invisible, commensal shrimps
(Periclimenes sp.) living among its tentacles and on its body wall.
Uses: It is popular in the aquarium trade, but it is difficult to keep in a reef aquarium
because of its specialist feeding requirements.

Cytotoxicity and antiangiogenic activity: The triterpenoids, philinopgeneins, and their ana-
logues philinopsides of this species have shown strong cytotoxicity against human tumour can-
cer cells. Some of them also exert potent antiangiogenic activity (Sima and Vetvicka, 2011).
Mondol et al. (2017) reported that the compounds Philinopsides A, B showed cyto-
toxic activity against human tumour cell lines with IC50 values of 1.70–3.50 µg/mL and
0.75–3.0 µg/mL, respectively.
Antitumour activity: A sulfated triterpene glycoside, philinopside E, of this species
showed a significant cytotoxicity (IC50 = 0.75–3.50 µg/mL) against ten tumour cell lines
(mouse lymphocytic leukaemia cells-P388, HL60, A549, lung adenocarcinoma cells-
SPC-A4, gastric carcinoma cells—MKN28, gastric carcinoma cells-SGC7901, BEL7402,
human ovarian carcinoma—HO8901, human fetal lung fibroblasts-W138, and human epi-
thelial carcinoma cells-A431) (Li et al., 2013).
Anticancer activity: Triterpene glycosides, philinopsides A, B, E, and F, as well as pen-
tactasides I, II, and III, isolated from this species revealed significant cytotoxicities in vitro
against such tumour cell lines as U87MG, A-549, P-388, MCF-7, HCT-116, and MKN-28
with IC50 in the range of 0.60–3.95 µM (Aminin et al., 2015).
Colochirus robustus (Östergren, 1898)
Common name(s): Robust sea cucumber, yellow sea cucumber

Global distribution: It is found in tropical parts of the central Indo-Pacific region; its
range includes the Philippines and Indonesia.
Ecology: This shallow-water species is found on rocks and reefs in places with moderate
to rapid water flow; depths are down to about 25 m.
Biology
Description: It is roughly cylindrical with five shallow longitudinal ribs, and it grows to
7 cm in length. It has an angular appearance and finger-like or thorn-like protuberances on
the ribs. At the anterior end, there is a ring of about eight large, feathery feeding tentacles.
There is a slight transverse indentation near the rounded posterior end. There are three rows
of tube feet on the underside. The colour of the individuals is a vivid yellow, sometimes
with grey between the ridges.
Food and feeding: It feeds on zooplankton and other organic particles. It clings to the sub-
strate with its tube feet, spreading its feathery tentacles to catch its food materials as they float
past. The tentacles are then retracted to the mouth where the food particles are scraped off.
Reproduction: It can reproduce both sexually and asexually. In the latter case, fission can
take place with a transverse crack developing halfway along the body and gradually wid-
ening until the two halves split apart. The posterior end then grows a new anus while the
anterior end develops a mouth and new tentacles.
Aquarium values: It is suitable for reef aquaria.

Edible/medicinal values: It has long been used in East and Southeast Asia as nutritious
food as well as for certain medicinal purposes.

Haemolytic activity: The compound colochiroside E, an unusual non-holostane triterpene
sulfated trioside, showed weak haemolytic activity (Blunt et al., 2017).
Immune function: Oral administration of peptides of this species (0, 25, 50, and 75 mg/kg
body weight) in C57BL/6 mice enhanced lymphocyte proliferation, serum albumin (ALB)
levels, and the natural killer (NK) cell activity. Moreover, these peptides promoted func-
tions of helper T cells (Th) as indicated by increased production of Th1 type cytokines of
Interleukin (IL)-1β, IL-2, Interferon (IFN)-γ, and TNF-α and Th2 type cytokines (IL-4,
IL-6 and IL-10) (Du et al., 2017).
Cucumaria conicospermium (Levin & Stepanov, 2002)
Global distribution: Sea of Japan

The triterpene glycosides, cucumariosides A2-5, A3-2, A3-3, and isokoreoside A, along
with the previously isolated koreoside A, have been isolated from this species. Activities
of these compounds are to be known (Avilov et al., 2003; Mondol et al., 2017).
Silchenko et al. (2005) reported on the isolation of three new minor monosulfated triter-
pene glycosides, viz. frondosides A2-4, A2-7, and A2-8; and disulfated cucumariosides,
A3-2 and A3-3; and trisulfated koreoside A and isokoreoside A.
Cucumaria fallax (Ludwig, 1875)
Common name(s): Pale sea football
Global distribution: Aleutian Islands, Alaska

No other information is available for other species.

Blunt et al. (2017) reported on the isolation of trisulfated non-holostane triterpene glycoside,
viz. fallaxosides D4, D5, D6, and D7; and unusual oligosulfated triterpene glycosides, viz.
fallaxosides C1, C2, D1, and D2. Their activities are to be known.
Silchenko et al. (2017) reported on the occurrence of two unprecedented triterpene glyco-
sides, fallaxosides B1 and D3.
Cucumaria frondosa (Gunnerus, 1767)
Common name(s): Orange-footed sea cucumber
Global distribution: It is found in the North Atlantic Ocean and the Barents Sea (Russia);
it is most abundant along the eastern coast of North America.
Ecology: Its habitat is rocks, crevices, or low-tide waters. It is known to cover vast areas of
the substrate at depths of less than 30 m.
Biology
Description: These sea cucumbers reach about 20 cm in length and have ten branched oral
tentacles that range in colour from orange to black. This species has a football shape with a
leathery skin ranging in colour from yellowish white to dark brownish black and is covered
with five rows of retractile tube feet. The young are about 1–6 mm long and are translucent
orange and pink. Adults of this species have a reduced number of spicules (skeletal struc-
tures), which are shaped like rounded plates with many holes. In this species, the sexes can
be identified by the conspicuous tube-shaped (female) or heart-shaped (male) gonopore
located under the crown of oral tentacles.
Food and feeding: It is a suspension feeding organism and catches available particles on
its tentacles.

Cytotoxicity: A new non-holostane triterpene glycoside, frondoside C, has been iso-
lated from this species. Its desulfated derivative presented an intense cytotoxic activity
(IC50 = 1 μg/mL) in mice and human tumour cell lines (Avilov et al., 1998).
Anticancer activity: A novel triterpenoid glycoside frondoside A was isolated from this
species (Girard et al., 1990). This compound exhibited anticancer activity in a human
pancreatic mouse xenograft model when given intraperitoneally. Further, it also exhibited
activity towards prostate cancer cell lines in vitro and in vivo (Blunt et al., 2017).
Antitumour activity: In animal trials, frondoside A inhibited the tumour growth of PC-3
and DU145 cells with a notable reduction in lung metastasis and in circulating tumour cells
in the peripheral blood (Ruiz-Torres et al., 2017).
Ruiz-Torres et al. (2017) also reported that frondoside A showed inhibition of migration
and invasion at 0.1–1 µM in the human estrogen receptor-negative breast cancer cell line
MDA-MB-231 in a wound-healing model assay. In addition, in an in vivo assay, frondoside
A (100 µg/kg/day i.p. for 24 days) strongly decreased the growth of MDA-MB-231 tumour
xenografts in athymic mice without any manifest toxic side effects.
Others: Mondol et al. (2017) reported on the presence of glycosides, frondosides B, A2-2,
A2-4, A7-1, A7-2, A, A2-1, A2-3 from this species.
Silchenko et al. (2005) recorded the presence of frondosides A2-1, A2-2, A2-4, A2-7, and
A2-8 from this species.
Cucumaria frondosa japonica (Semper, 1868) (= Cucumaria japonica)
Common name(s): Japanese sea cucumber
Global distribution: Tropics; far eastern seas of Russia

Ecology: The adults of this species prefer to inhabit sites off the open coast with rocky or
muddy substrate at depths of 5–300 m and in temperatures ranging from −1.8°C to 18.0°C.
The young ones prefer to inhabit kelp beds and shallow-water habitats warmed thoroughly
in summertime. These individuals are found distributed randomly on the bottom but some-
times congregate in aggregations of up to several hundreds.
Biology
Description: It is a fairly large sea cucumber, and its body length is up to 20 cm. Its live
weight is up to 1.5 kg, and the weight of the body wall is about 20% of the total weight.
The body is roundish and smooth, with five rows of tube feet. Its colour is greyish purple
or white.
Food and feeding: Its diet comprised an abundance of phytoplanktonic cells (Coscinodiscus
centralis, Chaetoceros debilis, Skeletonema costatum, and Thalassiosira gravida), with
occasional ingestion of small crustaceans and a variety of eggs and larvae.
Reproduction: It is gonochoric (separated sexes). It can produce up to 300,000 float-
ing eggs that are green, very large (500 µm), and go up to the surface during spawning.
Embryonic and larval development is observed in the upper-water layer and is probably
short. The spawning seems to occur twice a year, in April–June and September–October.
Edible values: It is the most commercially harvestable species. In Russia, all harvested
individuals are boiled, sliced in small pieces, and sold in local shops as a salad with the
addition of the seaweed Laminaria japonica (“sea cabbage”) and various spices.

Immunomodulatory and antibacterial properties: A complex of monosulfated
cucumariosides with cholesterol isolated from this species demonstrated immunomodulatory
properties in C57Bl6 mice. When administered in low doses, this complex displayed more
than twofold stimulation of lysosomal activity on mouse macrophages. In addition, this
complex significantly increased the animals’ resistance against bacterial infections elicited
by Y. pseudotuberculosis or S. aureus. Furthermore, it increased phagocytosis and ROS
formation, and stimulation of IL6 and TNF-α production was observed in lymphocytes
(Aminin et al., 2001; Janakiram et al., 2015).
Antifungal activity: The compounds cucumariosides I and II isolated from this
species exerted antifungal activity against C. albicans and C. tropicalis. The MIC of
cucumarioside I for these organisms was 30 and 40 µg/mL, respectively (Batrakov et al.,
1980; Bahrami, 2015).
Others: Mondol et al. (2017) reported on the presence of cucumariosides A0-1, A0-2,
A0-3; A1-2; A2-2, A2-3, A2-4; A4-2; and A7-1, A7-3 from this species.
Cucumaria koreaensis (Östergren, 1898)
Global distribution: Northern parts of Pacific and Atlantic Oceans

No other information is available for this species.

Mondol et al. (2017) reported on the isolation of a nonholostane triterpene glycoside, kore-
oside A, from this species.
Cucumaria okhotensis (Levin & Stepanov, 2003)
Global distribution: This is a tropical species.
Ecology: This is a benthic species.
Biology
Description: This is a large holothurian with a body length of 190 mm. The body is in the
form of a cucumber, elongated; the posterior part of the body is expanded, with an acute
extremity. The madreporic body is oval shaped, yellowish white, sometimes very large, up to
3.5 mm at a body length of 100 mm. The stone canal is straight. The animals fixed in alco-
hol are usually dark brown from the dorsal side and light brown from the ventral side. Podia
are yellowish white and conspicuous against the background of the body. Tentacles are light
coloured, grey, brownish, or yellowish, and their trunks are almost white.
Cytotoxicity: Triterpene oligoglycosides, okhotosides B1 (1), B2 (2), and B3 (3); and fron-
doside A (4), frondoside A1, cucumarioside A2-5, and koreoside A have been isolated from
this species. Among these compounds, compounds 1–3 were found to be moderately toxic
against HeLa tumour cells. Frondoside A (4) showed more potent cytotoxicity against THP-1
and HeLa tumour cell lines (with IC50 values of 4.5 and 2.1 µg/mL, respectively) (Silchenko
et al., 2008).
Three monosulfated triterpene glycosides, viz. okhotoside A1-1 and okhotoside A2-1, and
a pentaoside cucumarioside, A0-1, have been isolated from this species (Silchenko et al.,
2007; Mondol et al., 2017).
Hemioedema spectabilis (Ludwig, 1883)
Common name(s): Patagonian sea cucumber
Global distribution: Argentina Sea
Biology
Description: Tube feet are scattered all over the body surface. Deposits rods.
Cytotoxic and antiproliferative activities: The triterpene glycosides, hemoiedemosides

A and B, isolated from this species exhibited in vitro cytotoxic and antiproliferative
activities of compounds against A549 and HeLa cell lines as detailed below (Careagaa
et al., 2014).
A549 HeLa
IC50 (µM) IC50 (µM)
Hemoiedemoside A 7.43 9.95
Hemoiedemoside B 3.16 2.15
Source: Careaga, V.P. et al., Nat. Prod. Res., 28,

213–220, 2014.
Antifungal activity: The triterpene glycosides, hemoiedemosides A (1) and B (2), exhibited
considerable antifungal activity against the phytopathogenic fungus Cladosporium
cucumerinum. While the compound 1 showed inhibition zones of 8–33 mm at the tested
concentrations (1.5–50 µg/spot), compound 2 was found to be less active than compound 1
(Chludil et al., 2002b).
Mensamaria intercedens (Lampert, 1885)
Common name(s): Orange sea cucumber
Global distribution: Tropical Western Central Pacific: South China Sea
Ecology: This coastal species is found on intertidal flats in “U-shaped” burrows or

entwined in rhizomes of seagrasses. It has a depth range of 0–183 m.
Biology
Description: It is a blue-black sea cucumber with bright red ambulacra. It is a small (<115 mm),
cylindrical species, tapering at both ends, with 30 tentacles. Tube feet are concentrated along
ambulacra. The body is thin and pliable, and the tegument is smooth. Spicules are in the
form of tables with a high spinous spire and a pillar with two bridges; the disc is formed
with four large and four small holes.
Food and feeding: It is a suspension feeder.

Anticancer and antineoplastic activities: Three new triterpene glycosides, intercedensides
A, B, and C (1-3), isolated from this species showed anticancer activity against human tumour
cell lines, viz. A549, MCF-7, IA9, and CAKI-1; human glioblastoma cells, U-87-MG, PC-3,
KB, and KB-VIN; and human skin melanoma cells, SK-MEL-2 and HCT-8. Interestingly,
all these compounds showed a significant cytotoxicity against all tumour cell lines within
the IC50 value range of 0.7–4 µg/mL, and the compounds 1 and 3 showed similar potencies,
while compound 2 was generally more potent in all cell lines. Furthermore, compound 1
also exhibited significant in vivo antineoplastic activity against mouse Lewis lung cancer and
mouse S180 sarcoma, with 48.39% and 57.48% tumour reduction levels (Li et al., 2013).
Cytotoxicity: Six new triterpene glycosides, intercedensides D–I (1-6), have been isolated
from this species. Among these compounds, intercedensides D–H (1-5) showed significant
cytotoxicity (ED(50) 0.96–5.0 µg/mL) against 10 human tumour cell lines (Zou et al., 2005).
Cytotoxicity and antiangiogenic activity: Sima and Vetvicka (2011) reported that the
compounds, intercedensides, of this species showed strong cytotoxicity against human
tumour cancer cells. Further, they also exerted potent antiangiogenic activity.
Pentactella leonina (Semper, 1867) (= Pseudocnus dubiosus leoninus)
Global distribution: Southern Peru in the west of South America to the Rio de la Plata in
the east, and the Falkland Islands
Ecology: It occurs from the intertidal zone to approximately 100 m.
Biology
Description: It is medium sized with length up to 100 mm. Tube feet are in five bands
and are numerous in the dorsal ambulacra. Tentacles are bushy and are of unequal size.
Calcareous ring is simple. The colour of this species varies between white and pink.

Cytotoxic and antiproliferative activities: The triterpene glycoside, pseudocnoside A,
isolated from this species showed in vitro cytotoxic and antiproliferative activities against
A549 and HeLa cell lines with IC50 values of 14.5 and 57.4 µM, respectively (Careaga
et al., 2014; Mondol et al., 2017).
Plesiocolochirus australis (Ludwig, 1875) (= Pentacta australis)
Common name(s): No common names found
Global distribution: Tropical west Pacific, East Indies, Australia
Ecology: P. australis may be found on reefs and in coastal areas, concealed among rubble,
on reef flats and rocky shores.
Biology
Description: It is a grey to orange sea cucumber with small papillae. These papillae are
present along the corners of its somewhat square, firm body (length 50–65 mm). The tegu-
ment is rough, and the spicules look like homogeneous-sized baskets.

Two non-holostane-type triterpenoid oligoglycosides, DS-penaustrosides A [1] and B [2],
and two holostane-type glycosides, DS-penaustrosides C [3] and D [4], have been isolated
from this species. Activities of these compounds are to be known (Miyamoto et al., 1992).
Pseudocolochirus violaceus (Théel, 1886)
Common name(s): Sea apple sea cucumber
Global distribution: Western Central Pacific Ocean: Singapore
Ecology: This tropical, benthic species is found in shores among seagrasses or attached to
rubble.
Biology
Description: The body of this species is long (10–20 cm) or spherical with five rows of
yellow tube feet. But the two rows on the upper side of the body are usually less distinct.
The organism is usually red on the upper side, shading to lilac and white to the underside.
The mouth is ringed with blue. During low tide, it retracts its colourful feeding tentacles.
When relaxed, its normal shape is short and sausage like. When stressed, however, it may
inflate itself into a large round ball.
Food and feeding: Sea apples are filter feeders with tentacles, ovate bodies, and tube-like
feet. They can release their internal organs or a toxin into the water when stressed.
Aquarium values: Unfortunately, these beautiful sea cucumbers are harvested for the
aquarium trade. Ironically, they do not make good aquarium specimens, as they are often
toxic to their tank mates.

Cytotoxicity: The sulfated triterpene glycoside, intercedenside B, isolated from this
exhibited significant cytotoxicity against cancer cell lines MKN-45 (human gastric
adenocarcinoma) and HCT-116 with IC50 values in the range of 0.052–0.442 μM, and
both compounds showed significantly higher activity against HCT-116 compared to the
positive control HCP (Li et al., 2013).
Zhang et al. (2006b) reported that the two new bioactive triterpene glycosides, viz. vio-
laceusides A and B isolated from this species, exhibited significant cytotoxicity against
HL-60 and BEL-7402 cancer cell lines.
Zhang et al. (2006c) reported on the isolation of three new sulfated triterpene glycosides—
i.e., violaceusides I, II, and III—from the BuOH extract of this species. All these glycosides
showed significant in vitro cytotoxicities against human gastric cancer MKN-45 and
human colon cancer HCT-116 cells.
Silchenko et al. (2014) reported on the isolation of four triterpene glycosides, violaceusosides C
(1), D (2), E (3), and G (4) from this species. While compounds 1 and 2 demonstrated moderate
cytotoxic and haemolytic effects, violaceusosides E (3) and G (4) had more powerful activities.
Pseudocnus californicus (Semper, 1868)
Global distribution: Bay of Panama, eastern Pacific
Ecology: It is found subtidally in shallow water in crevices.
Biology
Description: It has two smaller ventral tentacles and its tube feet are radial. The body wall
lacks knobbed buttons.

Cytostatic (growth inhibition) and cytotoxic effects: The whole body extracts of this
species showed a strong cytostatic (growth inhibition) and cytotoxic effect against two
human cell lines, lung carcinoma A-549 and colon carcinoma HT-29 (Petzelt, 2005).
Pseudocnus echinatus (von Marenzeller, 1881) (= Cucumaria echinata)
Global distribution: Indo-Pacific: from the Red Sea and Bay of Bengal to China and Japan
Biology
Description: The body of this species is columnar. The skin is thin. It has a brown spot.
There are ten tentacles. The tube foot strikes in two rows on the pedestrian zone. The body
has reddish skin colour. Length of the specimens varies from 17 to 22 mm.
Reproduction: It is a dioecious species, with males and females being indistinguishable
externally. Spawning takes place in this species between mid-June and early August, usually
in the late afternoon. The gametes are liberated into the water column and fertilization
takes place at once. The larvae are planktonic at first before settling on the seabed, growing
feeding tentacles and metamorphosing into juvenile sea cucumbers.

Haemolytic/antimalarial activity: The galactose-specific lectins present in this species
showed direct haemolytic activity. These lectins damage the cell membrane, leading to
cell lysis (Petzelt, 2005). Further, this lectin has the ability to block the development of
Plasmodium, the causal agent of malaria, when it is expressed in genetically modified
Anopheles mosquitoes.
Haemolytic and hemagglutination activities: The C-type lectin CEL-III of this
species showed strong haemolytic activity toward human and rat erythrocytes at low
concentrations (IC50 = 0.3 and 0.8 µg/mL), respectively. Similarly, CEL-III exhibited
strong hemagglutination activity towards human and rat erythrocytes (Yoshida et al.,
2007).
Neuritogenic activity: Five cerebrosides, CE‐1‐1, CE‐1‐2, CE‐1‐3, CE‐3‐1, and CE‐3‐2,
and a ganglioside molecular species CG‐1, have been obtained from the less polar and
polar fractions, respectively, of the chloroform/methanol extract of this species. Among
these compounds, the ganglioside CG‐1 exhibited neuritogenic activity towards the rat
pheochromocytoma cell line, PC‐12 cells (Yamada et al., 1998).
Antifungal activity: Triterpene glycosides—cucumechinosides A, B, C, D, E, and F—of

this species showed antifungal activity against Candida albicans, Trichomonas foetus,
Trichophyton mentagrophytes, Aspergillus niger, Mucor hiemalis, and Penicillum chrys
ogenum (Bahrami, 2015).
Staurocucumis liouvillei (Vaney, 1914) (Ekman, 1927)
Common name(s): Antarctic sea cucumber
Global distribution: Antarctic
Ecology: This polar, benthic species has been recorded from a depth of 229 m.
Biology
Description: It has a thin body wall with an outer gelatinous brown layer that darkens with
age. There are ten equal, long, very branched tentacles that are always extended. Tube feet
are present on radii only and are more developed ventrally. A calcareous ring is normally
not evident. While ossicles are frequently not evident in larger specimens, oval bowl ossicles
with denticulate perforations are present. Individuals have a length of 27–80 mm.

Cytotoxic and virucidal activity: The trisulfated triterpene glycosides, liouvillosides A
and B, isolated from this species were found to have cytotoxic and virucidal activities.
While liouvillosides A was against herpes simplex virus type 1 (HSV-1) at concentrations
below 10 µm/mL, liouvillosides B was cytotoxic against human tumour cell lines with
IC50 values of 0.44–2.62 µg/mL (Maier et al., 2001; Mondol et al., 2017).
Others: Triterpene glycosides, liouvillosides A1 (1), A2 (2), A3 (3), B1 (4), and B2 (5), have
been recorded from this species (Antonov et al., 2008).
Antonov et al. (2011) reported on the presence of triterpene glycosides, liouvillosides A4

and A5 in this species.
Staurocucumis turqueti (Vaney, 1906)
Global distribution: Antarctic: Antarctica, Terre Adelie, and South Orkney
Ecology: This polar species lives in benthic habitats.
Biology

The triterpene holostane disulfated tetrasaccharide olygoglycoside, turquetoside A, has
been isolated from this species (Silchenko et al., 2013b).
Thyonidium kurilensis (Levin, 1984) (= Duasmodactyla kurilensis)
Common name(s): Kurile sea cucumber
Global distribution: Northeast Pacific and Northern Atlantic: US, Canada, and Alaska
Ecology: This tropical, demersal species has a depth range of 10–228 m.
Biology
Two new triterpene glycosides, kurilosides A and C, have been isolated from this species
(Avilov et al., 1991).
5.3 FAMILY: PHYLLOPHORIDAE (ORDER: DENDROCHIROTIDA)

Massinium magnum (Ludwig, 1882) (= Neothyonidium magnum)
Common name(s): Magnum sea cucumber, burrowing sea cucumber, Indo-West Pacific
sea cucumber, Vietnamese sea cucumber, sea basket sea cucumber
Global distribution: Widespread in Indo-Pacific region
Ecology: It is found singly buried in the sand of shallow waters on sand and rubble areas;
the depth range is 0–40 m.
Biology
Description: It grows to a length of 75 mm. Colour of the individual is yellowish brown;
tube feet and tentacles are darker brown. The mouth is encircled by well-developed
brownish-white peristomial membrane. There are twenty tentacles in two distinct circles;
the outer circle has five pairs of large (25–45 mm) tentacles, alternating with an inner
circle of five pairs of small (5–7 mm) tentacles. While the shaft of large tentacles are pale
brown with one dark banding proximally, tips of these tentacles are darker brown. The
shaft of small tentacles is uniformly pale brown and the tips are darker. Tube feet are
found distributed in two radial rows and are decreasing in size proximally, where they also
scatter over interradial areas.
Food and feeding: These animals feed on plankton; When feeding, they fan out their
“arms” and then one by one roll them into their mouths to feed.

Anticoagulant activity: The compound, fucosylated chondroitin sulfate, isolated
from this species exhibited anticoagulant activity. In the experiments with purified
proteins, this species effectively potentiated inhibition of thrombin and factor Xa by
ATIII. Besides, this species did not induce platelets aggregation in platelets-rich plasma
(Ustyuzhanina et al., 2017a).
Cytotoxic and haemolytic activities: Two new monosulfated triterpene glycosides,

magnumosides B3 and B4, and one new disulfated triterpene glycoside, magnumoside C3,
isolated from this species showed cytotoxic activity against mouse spleen lymphocytes
and ascite form of mouse Ehrlich carcinoma cells, human colorectal carcinoma DLD-1.
Further, magnumoside B4 was found to show haemolytic activity by greatly suppressing the
colony formation and decreasing the size of colonies of DLD-1 cancer cells at noncytotoxic
concentrations (Silchenko et al., 2017a).
The sulfated triterpene glycosides, magnumosides A1 (1), A2 (2), A3 (3), A4 (4), B1 (5), B2
(6), C1 (7), C2 (8), and C4 (9) as well as a known colochiroside B2 (10), isolated from this
species have shown cytotoxic and haemolytic activities against mouse spleen lymphocytes,
the ascites form of mouse Ehrlich carcinoma cells, and human colorectal carcinoma DLD-1
cells. Interestingly, the erythrocytes were more sensitive to the glycosides action than
spleenocytes and cancer cells tested. The compounds 3 and 7 significantly inhibited the
colony formation and decreased the size of colonies of DLD-1 cancer cells at noncytotoxic
concentrations. Moreover, the synergism of effects of radioactive irradiation and
compounds 3 and 7–9 at subtoxic doses on proliferation of DLD-1 cells was demonstrated
(Silchenko et al., 2017a).
Pentamera calcigera (Stimpson, 1851)
Common name(s): Stony sea cucumber
Global distribution: Northeast Pacific and West Atlantic: Gulf of St. Lawrence, southern
Gaspe waters, downstream part of middle St. Lawrence estuary, and lower St. Lawrence
estuary
Biology
Description: It is variable in shape and has a maximum size of 9 cm. Colouration of the
body is white.

Cytotoxicity: Three new monosulfated triterpene glycosides, viz. calcigerosides B, C(1),
and C(2) along with the known cucumarioside G(2), have been isolated from this species.
The first three compounds showed moderate cytotoxicity (IC(50) = 5 µg/mL) against a
selection of four human and mouse tumour cell lines (Avilov et al., 2000).
Others: Mondol et al. (2017) reported on the occurrence of glycosides, viz. calcigerosides
D2, C2, and E.
Avilov et al. (2000) reported on the occurrence of three new triterpene glycosides—
calcigerosides D(1), D(2), and E—from this species.
5.4 FAMILY: PSOLIDAE (ORDER: DENDROCHIROTIDA)
Psolus eximius (Savel’eva, 1941)

No information on the biology and ecology is available on this species.

The compound, eximisoside A, has been isolated from this species (Mondol et al., 2017).
Psolus fabricii (Düben & Koren, 1846)
Common name(s): Scarlet psolus
Global distribution: Arctic and Northwest Atlantic: from Arctic to Cape Cod
Ecology: This benthic, polar species lives from lower intertidal zone depths greater than
90 m, on hard bottoms.
Biology
Description: The bottom of this sea cucumber is like the sole of a shoe, flat and rimmed
with a marginal band of tube feet and a weaker row down the middle. Its top is domed, with
the anus on a mound at one end. The anus is also ringed with 5–6 granular scales. Scarlet
sea cucumbers are bright red and are with a size up to 20 cm.
Food and feeding: These animals are suspension feeders. They extend branched and
mucus covered tentacles into water to trap suspended particles including live plankton.
Tentacles then are pushed to the mouth.
Reproduction: In this species, the sexes are separated, with eggs being fertilized externally.
They also have a free-swimming larval stage. They are unique among echinoderms in
possessing a single gonad that opens the outside between two pairs of tentacles.

Activities relating to cellular processes: The acetylated triterpene glycosides, viz.
psolusosides A, B isolated from this species, have been reported to inhibit the membrane
transporter, Na+-K+-ATPase, an enzyme found in the plasma membrane. It is sodium-
potassium adenosine triphosphatase, also known as the “Na+/K+pump,” “sodium-
potassium pump,” or simply “sodium pump” that is important for the functioning of most
cellular processes. Further, it is integral in maintaining the acid-base balance as well as in
healthy kidney function (Bahrami et al., 2016).
Psolus patagonicus (Ekman, 1925) (= Psolus marcusi)
Common name(s): Patagonian rock sea cucumber
Global distribution: Southwest Atlantic: Argentina and Chile
Ecology: This subtropical, benthic species is found on rocks, Macrocystis fronds, and
holdfast; individuals are often found attached to the Patagonian scallop, Zygochlamys
patagonica. Depth range is intertidal to 308 m.
Biology
Description: In this species, the mouth and anus are dorsal in position and are covered by
five valves. There are ten tentacles that are white in colour with brown dots. Tube feet are
up to 0.35 mm in diameter and are present only on the ventral side. The calcareous ring is
simple. Respiratory trees are well extended up to the anterior part of the body. Colour of
the individual in life is light orange to white. It grows to a size of 26 mm.
Food and feeding: It is a deposit feeder on a variety of benthic organisms.
Reproduction: Spawning occurs between February and March. Eggs are brooded under
the mother’s sole until they develop into crawling juveniles within seven months. During
the brooding period (February–September), the number of brooded embryos decreased
while their size increased.

Antifungal activity: The triterpene glycoside, patagonicoside A(1) and its desulphated
derivative, ds-patagonicoside A (2), isolated from this species showed antifungal activity
against the pathogenic fungus Cladosporium cucumerinum. While compound 1 showed
high activity with inhibition zone values of 8–19 mm at concentrations of 1.5–50 µg/
spot, compound 2 was weakly active with inhibition zone values of 5–9 mm at highest
concentrations of 6–50 µg/spot (Murray et al., 2011).
Careaga et al. (2011) reported that the glycosides, patagonicosides A(1), B(2), and C(3) and
their desulfated analogues isolated from the ethanolic extract of this species, showed anti-
fungal activity against the phytopathogenic fungus Cladosporium cladosporoides in a dose-
dependent activity. Patagonicoside A (1) resulted to be considerably active, in a dose-dependent
activity, showing inhibitions zones of 10–22 mm at the tested concentrations (5–20 µg/spot).
Glycosides 2 and 3 were less active than 1 but more active than their desulfated analogues 2a
and 3a, which were inactive at the lowest concentration (5 µg/spot) and weakly active (inhibi-
tion zones of 6 and 7 mm, respectively) at the highest tested concentration (20 µg/spot).
Cytotoxic and antiproliferative activities: The compounds, patagonicosides B and C iso-
lated from this species, showed in vitro cytotoxic and antiproliferative activities against
A549 and HeLa cell lines as detailed below (Careaga et al., 2014; Mondol et al., 2017).
A549 HeLa
IC50 (µM) IC50 (µM)
Patagonicoside B 9.7 7.9
Patagonicoside C 5.6 3.6
Source: Careaga, V.P. et al., Nat. Prod. Res., 28,

213–220, 2014.
Psolus paradubiosus (Carriol & Féral, 1985)
Common name(s): Basket rock sea cucumber
Global distribution: Southeast Pacific and Antarctic
Ecology: This temperate to polar and benthic species is found on rock at depths of
10–567 m.
Biology
Antitumoural/cytotoxic activity: The aqueous and dichloromethane/methanol extracts
of this species showed antitumoural activity against HT-29 (colorectal adenocarcinoma),
A-549 (lung carcinoma), and MDA-MB,231 (breast adenocarcinoma) cell lines as detailed
below (Taboada et al., 2010).
Percentage Values of Cell Growth for the Active Samples Against Human Tumour Cell Lines
Extract HT-29 A-549 MDA-MB,231
A −90 −90 −87
DM −83 −85 −83
A, aqueous extract; DM, dichloromethane/methanol extract.

Negative values, in the range between 0 and 100, represent samples with cytotoxic activity.
Source: Taboada, S. et al., Antarct. Sci., 22, 494–507, 2010.
5.5 FAMILY: SCLERODACTYLIDAE (ORDER: DENDROCHIROTIDA)

Cladolabes schmeltzii (Ludwig, 1875)
Common name(s): Vietnamese sea cucumber, Indo-West Pacific sea cucumber
Global distribution: Madang’s Reef, Madang Province, Papua New Guinea
Ecology: It is a coral reef-associated species recorded from a depth of 3 m. It is found in

reefs and coastal waters, below rocks and rubble.
Biology
Description: It is a light to dark brown, small (9 mm when preserved), short and cylindri-
cal sea cucumber covered with tube feet, which are in rows along the ambulacra only, more
numerous ventrally than dorsally. Spicules are short, spiked rods. Only the outer circle of
10 tentacles is visible in this species.

Cytotoxic and haemolytic effects: The triterpene glycosides, cladolosides Al-A6, have
been isolated from this species. All the compounds except cladoloside A5 showed cyto-
toxic activities against mouse spleenocytes and haemolytic effects against mouse erythro-
cytes (Silchenko et al., 2014).
Silchenko et al. (2013b) also reported that the nonsulfated triterpene glycosides, viz.
cladolosides B1, B2, C, C1, C2, and D isolated from this species, demonstrated rather
strong cytotoxic and haemolytic effects.
Silchenko et al. (2015) reported that the nonsulfated triterpene glycosides, cladolosides
C3 (1), E1 (2), E2 (3), F1 (4), F2 (5), G (6), H1 (7), and H2 (8) of this species demonstrated
strong or moderate cytotoxic and haemolytic effects.
Cytotoxic activities: The triterpene olygoglycosides—cladolosides C4 (1), D1 (2), D2 (3),
M (4), M1 (5), M2 (6), N (7), and Q (8)—have been isolated from this species. The majority
of tested compounds, except for cladoloside D2 (3), showed cytotoxic activities against
human colorectal adenocarcinoma HT-29 cells by inhibiting their colony formation and
growth at noncytotoxic concentrations. The highest inhibitory activity was demonstrated
by cladoloside M1 (5). Moreover, synergism of effects of radioactive irradiation and
nontoxic dose of compounds 1–8 decreasing the number of colonies of HT-29 cells was
observed. Cladoloside N (7) was the most active and increased the inhibitory effect from
radiation by 75% (Silchenko et al., 2018).
Eupentacta fraudatrix (D’yakonov & Baranova in D’yakonov et al., 1958)
Common name(s): Far Eastern sea cucumber
Global distribution: Peter the Great Bay, East/Japan Sea
Ecology: This tropical, demersal species is found in the subtidal zone.
Biology
Associated species: The ectosymbiotic copepod, Vostoklaophonte eupenta, has been
found associated with this species (Yeom et al., 2018).

The following glycosides have been isolated from this species (Avilov et al., 1991; Silchenko
et al., 2013).
Cytotoxic and haemolytic activities: Seven new minor triterpene glycosides—cucumari-

osides A2 (1), A7 (2), A9 (3), A10 (4), A11 (5), A13 (6), and A14 (7)—have been isolated
from this species. Among these compounds, cucumariosides A2 (1), A8 (8), and A13 (6)
demonstrated high haemolytic activities against mouse erythrocytes. Glycosides 1, 4, and
6 showed moderate cytotoxic activity against mouse spleen lymphocytes and the cells of
the ascite form of mouse Ehrlich carcinoma (Silchenko et al., 2012a).
Four new triterpene glycosides cucumariosides H5 (1), H6 (2), H7 (3), and H8 (4) along
with the known cucumarioside H (5) have been isolated from this species. Among them,
glycosides 1–3, and 5 were found to be cytotoxic against mouse lymphocytes and haemolytic
against mouse erythrocytes. Glycoside 2 was, however, less active in comparison with
others (Silchenko et al., 2011).
Two new minor triterpene glycosides, cucumariosides B1 (1) and B2 (2) have been isolated
from this species. Among them, cucumarioside B2 (2) demonstrated moderate haemolytic
activity against mouse erythrocytes and low cytotoxic action against Ehrlich carcinoma
cells (Silchenko et al., 2012b).
Cytotoxicity and lysosomal activity: Triterpene glycosides, viz. cucumariosides I 2 (1),
H (2), A5 (3), A6 (4), B2 (5), and B1 (6), have been isolated from this species. While
glycosides 1 and 5 possessed low cytotoxicities against mouse Ehrlich carcinoma cells,
compounds 2, 3, and 4 possessed moderate cytotoxicities. Further, glycosides 1, 3, and
5 increased the lysosomal activity of macrophages on 15%–17% at doses of 1–5 μg/mL
(Silchenko et al., 2013).
Others: Ustyuzhanina et al. (2017b) reported on the presence of two fucosylated chondroi-
tin sulfates EF1 and EF2 from this species.
Sclerodactyla briareus (Lesueur, 1824) (= Holothuria briareus; Thyone briareus)
Common name(s): Hairy sea cucumber, hard-fingered sea cucumber
Global distribution: Western Atlantic: east coast of the United States from Massachusetts
to Florida, plus the Gulf of Mexico, the Caribbean Sea, and extending southward as far as
Venezuela
Ecology: This tropical, benthic species lives in shallow waters; depth range is 0–66 m.
This species burrows in soft substrates and is more tolerant than many other echinoderms
of water with low salinity levels or low oxygen levels. If threatened by predators, it can
eject its viscera in a sticky, white mass to confuse and deter the aggressor.
Biology
Description: It is an elongated oval or cigar-shaped sea cucumber and grows to about
15 cm. It often adopts a characteristic pose with both ends raised above the substrate. At
the anterior end, there is a mouth surrounded by a ring of ten, short, branched feeding
tentacles. These are modified tube feet that can be retracted back into the mouth when
the animal is not feeding. The surface of the body is divided into five ambulacral areas
with five shallow interambulacral grooves in between. Just behind the tentacles there is a
group of small ossicles, calcareous stiffening plates, forming a short tube surrounding the
pharynx. These provide a skeletal support for the muscles and internal organs. The skin
is thick and leathery, and the tube feet are scattered and are protruding as soft finger-like
projections from the body wall. The minute calcareous spicules of this species are square
or round and table shaped with spires formed of four pillars. The colour of the specimen is
dark greenish brown or charcoal grey with pale grey tentacles on blackish stems.
Food and feeding: It is a scavenger and filter feeder, collecting organic matter with its
feeding tentacles and thrusting the particles into its mouth.
Reproduction: It is gonochoristic with individuals being either male or female although
there is no outward difference in appearance. Fertilization is external, and the developing
larvae form part of the plankton.

Activity relating to cardiac rhythmicity: The smooth muscle of this species has been reported
to have the coexistence of excitatory nicotinic ACh receptor (nAChRs) and mAChRs, which
are of pharmacological importance. While the nAChRs possibly modulate tone, the muscarinic
ACh receptors (mAChRs) initiate and enhance cardiac rhythmicity (Devlin et al., 2000).
Wound healing property: Its polyunsaturated fatty acids, (arachidonic acid, eicosapentaenoic
acid, docosahexaenoic acid) have been reported to possess wound healing effects (Bordbar
et al., 2011).
5.6 FAMILY: ELPIDIIDAE (ORDER: ELASIPODIDA)

Kolga hyalina (Danielssen & Koren, 1879)
No information is available relating to the biology and ecology of this species.

Cytotoxic and haemolytic activities: Triterpene holostane nonsulfated pentaosides,
kolgaosides A (1) and B (2), and holothurinoside B (3) have been isolated from this species.
Among these compounds, kolgaosides A (1) and B (2) demonstrated low cytotoxic activity
against the cells of the ascite form of mouse Ehrlich carcinoma and moderate haemolytic
activity against mouse erythrocytes (Silchenko et al., 2014; Mondol et al., 2017).
Rhipidothuria racovitzai (Hérouard, 1901) (= Achlionice violaecuspidata)
Global distribution: Antarctic Indian Ocean: Eastern Antarctica; Western Antarctica,

Weddell Sea
Ecology: This polar, benthic species has a depth range of 225–785 m.
Biology
Description: Specimens of this species are up to 60 mm long. There are ten tentacles; the
anteriormost is the largest. There are seven pairs of dorsal papillae, distributed evenly along the
dorsum. Among its twelve pairs of tube feet, the posterior ones are the smallest and webbed.
Tentacles, mouth, papillae, tube feet, and anus are with residual violet brown colouration.

The triterpene glycosides, achlioniceosides A1, A2, and A3 have been isolated from
this species. But their bioactivity has not been reported (Antonov et al., 2009; Mondol
et al., 2017).
5.7 FAMILY: HOLOTHURIIDAE (ORDER: HOLOTHURIIDA)

Actinopyga agassizi (Selenka, 1867)
Common name(s): Five-toothed sea cucumber, West Indian Sea cucumber, Bahamian sea
cucumber
Global distribution: Tropical Western Atlantic, including the Gulf of Mexico, the
Caribbean Sea, the Southeast US Continental Shelf, the Bahamas, Bermuda, and
northern Cuba
Ecology: This is an epibenthic, subtidal, nocturnal species found in coral reefs

(including coral reef lagoons), seagrass, and rocky bottoms from shallow depths down
to 60 m.
Biology
Description: The ground colour of this species is white, with yellow brown spots. The
tegument (skin) is thick and leathery, with papillae (small projections) on the dorsal sur-
face. On the other hand, the ventral surface is flat with broad, knobby podia (tube feet). The
dorsal surface is rounded and the podia on the ventral surface have suckers at their tips.
They also have five calcareous teeth. The species is known to have a maximum length of
approximately 35 cm.
Food and feeding: It is a nocturnal feeder utilizing organic particles and detritus as its
food.
Reproduction: In Bimini in the Bahamas, this species is mostly found as large individu-
als while smaller individuals were rarely observed. This therefore implies their life cycles
may be long and their turnover rates are possibly low. The species is gonochoric and there-
fore an individual is either male or female. It has external spawning and fertilization. Its
life cycle consists of embryos that later develop into auricularia larvae and subsequently
develop into doliolaria larvae, a barrel-shaped stage, and then they metamorphose into
juveniles.
Associated species: These animals act as hosts for the pearlfish Carapus bermudensis,
one of the closest associations of a vertebrate and an invertebrate host.

Anticancer activity: The triterpene glycoside, holothurin A isolated from this species,
showed anticancer activity against sarcoma-180 (Lewis lung carcinoma) (Lakshmi et al.,
2014; Aminin et al., 2015; Mondol et al., 2017).
Actinopyga caerulea (Samyn, Vandenspiegel & Massin, 2006)
Common name(s): Blue sea cucumber
Global distribution: Tropical Indo-West Pacific: from Comoros in the west to New
Caledonia in the east
Ecology: This reef-associated, relatively shallow water species is usually found during the
day and night on sand or sand and rubble; depth range is 12–45 m.
Biology
Description: This is a rare but large species with a maximum length of 40 cm. Body is loaf
shaped with slight ventral flattening (more or less cylindrical with some distal tapering). Body
wall is smooth and is up to 14 mm thick. Mouth is ventral and is surrounded by 15–18 large,
peltate, uniformly bluish grey tentacles. Colour in life is bluish with patches of white.
Food and feeding: It is predominantly a detritus/deposit feeder on coral patches on the
outer slope of coral reefs. It forages actively during the day.
Edible values: This species is harvested for the production of “bêche-de-mer.”

Nutritional values: This species has been reported to contain higher amounts of SFA
and MUFA but lower contents of PUFA, and amino acid, glycine, which are essential in
tissue regeneration and inflammatory diseases besides serving as a valuable, functional
food (Bordbar et al., 2011; Omran, 2013).
Actinopyga crassa (Panning, 1944)
Global distribution: Eastern Africa and Madagascar; and Southeastern Arabia
Ecology: This species is found on sand and rubble slopes; depth range is 25–40 m.
Biology
Aquarium value: This species is used in the aquarium trade.

Antileishmanial activity (antiprotozoal activity): The dichloromethane/methanol
extracts of this species showed antileishmanial activity against Leishmania amazonensis
(Oliviera et al., 2016).
Actinopyga echinites (Jaeger, 1833)
Common name(s): Brown sea cucumber, spiny sea cucumber, brownfish, deepwater redfish
Global distribution: Indo-Pacific; throughout the Western Central Pacific, Asia and
Africa, and Indian Ocean regions
Ecology: This tropical, reef-associated species is found on shallow seagrass beds and rub-
ble reef flats; depth range is 0–30 m.
Biology
Description: It is a brown sea cucumber. Its ventral tegument is with a variety of small
rosettes and rods that are occasionally perforated. Dorsal tegument is with larger rosettes
and branched plates. Ventral podia are with some straight, spinous rods. Tentacles are with
rods that are large, straight, or curved, and are very spiny at their extremities. This species
grows to a maximum length of 36 cm.
Reproduction: This species reproduces sexually, usually during the warm season. It is a
species of high potential fecundity and early sexual maturity. Spawning occurs between
the months of January and February.

Antitumour activity: The saponin, echinoside A, isolated from this species showed
anticancer activity by inhibiting the growth of tumours in mouse models and human
prostate carcinoma xenografts in nude mouse models (Li et al., 2009).
Antibacterial activity: The alcoholic extract of this species showed antibacterial activity against
Staphylococcus aureus with an inhibition zone value of 1.25 mm (Abraham et al., 2002).
Antifungal activity: The oligoglycosides, echinoside A and B, isolated from this species
showed antifungal activity (Kitagawa et al., 1980).
Actinopyga flammea (Cherbonnier, 1979)

Global distribution: Tropical Western Central Pacific: New Caledonia, Borneo, and the
South China Sea
Ecology: This benthic species occurs under boulders and on slopes; depth range is
35–60 m.
Biology
Description: Its maximum length is 45 cm. No other information is available on this
aspect.

The triterpenoid saponins, viz. 24(S)-hydroxy-25-dehydro-echinoside A, 22 ξ-hydroxy-24-
dehydro-echinoside A, 22 ξ-acetoxy-echinoside A and 25-hydroxy-dehydroechinoside A;
and a sapogenin, 16-keto-holothurinogenin have been isolated by acid hydrolysis of crude
saponins of this species (Motuhi et al., 2016; Mondol et al., 2017).
Actinopyga lecanora (Jaeger, 1833)
Common name(s): Stonefish, white-rumped sea cucumber, white-bottomed sea cucumber
Global distribution: Tropical Indo-Pacific: Western Pacific Region; East Africa and
Central Indian Ocean region; and Asian region
Ecology: In the daytime, this shallow-water species is found under large stones, in gaps in
reef slopes or in sheltered areas; depth range is 20–23 m.
Biology
Description: This animal may be elongated into a long sausage-like shape, or contracted
into a more rounded loaf shape, or even into a more globular oval shape. Surface is some-
what smooth with long, thin tube feet that are sparsely distributed all over. Sometimes, this
sea cucumber is seen with many tube feet on the underside. The spicules are composed of
small rods. Colours of the specimens are variable, ranging from shades of brown to golden
yellow or white, sometimes with blotches, sometimes a uniform colour. It has a maximum
length of 25 cm.
Food and feeding: This species is often found grazing on live and dead coral.

Antibacterial activity: The hydrolysates of this species produced from proteolytic
enzymes, namely alcalase, papain, pepsin, trypsin, bromelain, and flavourzyme showed
antibacterial activity against pathogenic gram-positive bacteria (Bacillus subtilis and
Staphylococcus aureus) and gram-negative bacteria (Escherichia coli, Pseudomonas
aeruginosa, and Pseudomonas sp.). Papain hydrolysis showed the highest DH value
(89.44%), followed by alcalase hydrolysis (83.35%). After 1 hour and 7 hours of
hydrolysis, Bromelain hydrolysate exhibited the highest antibacterial activities against
Pseudomonas sp., P. aeruginosa and E. coli at 51.85%, 30.07%, and 30.45%, respectively,
compared to the other hydrolysates. Protein hydrolysate generated by papain after 8 hours
of hydrolysis showed maximum antibacterial activity against S. aureus at 20.19%. When
the potent hydrolysates were further fractionated, three fractions from the bromelain
hydrolysates exhibited inhibitory activities against Pseudomonas sp., P. aeruginosa, and
E. coli at 24%, 25.5%, and 27.1%, respectively, and one fraction of papain hydrolysate
showed antibacterial activity of 33.1% against S. aureus (Ghanbari et al., 2012, 2018).
Antifungal activity: The methanol extract c of this species showed compounds, viz.
triterpene glycoside, holothurin B, and holothurin A. Among these compounds, holothurin
B showed in vitro antifungal activity against twenty fungal test isolates including ATCC
strain and was found to be most effective against Trychophyton mentagrophytes and
Sporothrix schenckii. MIC was in the range of 1.56 µg/mL (Kumar et al., 2007).
Antioxidant property: The ethanolic and methanolic tissue extracts of this species
showed moderate antioxidant activity (39%–34% DPPH radical scavenging activity
and 23.95–22.30 mmol/100 mL FeSO 4 FRAP value) (Bordbar et al., 2013).
ACE inhibitory and antihypertensive effects: The proteolysate prepared from this species
exhibited ACE inhibitory potentials in vivo. The pre-fed rats with the said proteolysate
at various doses (200, 400, 800 mg/kg body weight) exhibited a significant (p ≤ 0.05)
suppression effect after inducing hypertension. The optimum effective dose of proteolysate
that will produce maximal reduction in blood pressure was also determined. The results
showed that the 800 mg/kg body weight dose significantly reduced blood pressure without
noticeable negative physiological effect. In addition, there were no observable changes in the
rats’ heart rate after oral administration of the proteolysate. These findings suggest that the
proteolysate of this species could potentially be used for the development of functional foods
and nutraceuticals for prevention and treatment of hypertension (Vishkaei et al., 2016).
Antiamoebic activity: Human amoebiasis due to Entamoeba histolytica infection has
been affecting the quality of life and pace of development in countries with warm climatic
conditions. The available drugs provide only symptomatic relief and are not devoid of
side effects. This has resulted in obtaining novel molecules from natural sources having
antiamoebic activity. The methanol extract of this species displayed antiamoebic activity
in vitro with an MIC value of 125 µg/mL. In experimented rats, it revealed 88% inhibition
of trophozoites at the dose of 900 mg/kg body weight against Entamoeba histolytica.
(Lakshmi and Ghosal, 2015).
Antileishmanial and antifungal activities: The methanol extract and n-butanol fraction
of this species exhibited excellent Leishmania donovani inhibition activity in vitro and
in vivo. Further, the methanol extract also showed promising antifungal activity in vitro
(Bordbar et al., 2011).
Antileishmanial activity (antiprotozoal activity): The crude methanol extract of this species
displayed antileishmanial activity by reducing 88.50% and 72.45% of the promastigotes and
amastigotes of Leishmania donovani population, respectively, at the concentration of 100 lg/
mL. When applied at the same concentration, the ethyl acetate soluble fraction was poorly
active, inhibiting less than 22.0% of both parasites forms. On the other hand, the butanol
soluble fraction reduced 98.5% and 76.4% of the promastigote and amastigote growth.
Two glycosides, namely holothurin A (1) and B (2) have been isolated from the n-butanol
fraction of this species. Compound 2 was found to be more active than compound 1, both
in vitro and in vivo, and when applied at the concentration of 50 lg/mL, both were able to
reduce the viability of intracellular amastigotes by 45% (1) and 57.65% (2). In L. donovani
infected hamsters, the application of molecule 2 at a dose of 100 mg/kg/day for 5 days
resulted in a reduction of the parasite burden in 71.5%, in contrast to compound 1, which
reduced approximately 50%. Even at a lower dose (50 mg/kg/day for 5 days), molecule 2 was
significantly more efficient than 1, reducing the parasite burden in 40%, comparatively to
compound 2, which only allowed a reduction of 20% (Oliviera et al., 2016).
Actinopyga mauritiana (Quoy & Gaimard, 1833)

Common name(s): Surf redfish
Global distribution: Throughout the central Pacific and Indian Oceans; from eastern
Africa and the Red Sea in the west, to Hawaii, and the Solomon Islands in the southeast;
and Madagascar
Ecology: This tropical, reef-associated species prefers outer reef flats and fringing reefs,
mostly in the surf zone between 0 and 20 m. It is usually found in subtidal and intertidal
sandy areas as well.
Biology
Description: The body of this species is elongated and is arched dorsally (bivium) and
flattened ventrally (trivium). Bivium is sometimes wrinkled, wider in the middle, and tapering
towards both ends. Papillae on bivium are long and slender. Podia are numerous on trivium,
and they are stout and crowded on the radii and interadii. The mouth is ventral, surrounded
by 25 short and stout tentacles. The anus is surrounded by 25 stout anal teeth. Calcareous ring
shows large radial pieces and interradials with a triangular top. Colour of bivium is variable—
usually dark brown, with more or less distinct whitish spots. Tentacles are with large rods
bearing small spines. The species has a maximum length of 25 cm and a weight of 1 kg.
Food and feeding: It is active both day and night, and it feeds on detritus lying on hard reef
substrates.
Reproduction: This species has a high fecundity and early sexual maturity. In Guam,
the peak reproductive activity of this species is between April and August, whilst in New
Caledonia, it is in December and January; in the Solomon Islands, it is from October until
January. In Egypt it reproduces annually between spring and summer.

Hypotensive, antimicrobial, anti-amoebic, and hypoglycaemic properties: The organic
extract of this species yielded the compound echinoside B, which has been reported to possess
hypotensive, antimicrobial, anti-amoebic, and hypoglycaemic properties (Anon, http://shodh-
ganga.inflibnet.ac.in/bitstream/10603/31706/15/15_chapter%206%20&%20section%202.pdf).

Nutritional values: It showed the highest percentage of total protein (48.27%) and lycine:
argignin ratio (3.6). Further, it is also rich in SFA, MUFA, and PUFA (Bordbar et al., 2011;
Omran, 2013). This species may therefore serve as a good functional food.
Actinopyga miliaris (Quoy & Gaimard, 1833)
Common name(s): Blackfish, harry blackfish
Global distribution: Tropical Indo-Pacific, including the Red Sea, but excluding the
Persian Gulf and Hawai
Ecology: This coastal species inhabits reef flats of fringing reefs and lagoon islet reefs,
rubble reefs, and compact flats with depths of 0–20 m. It is also common in sandy beds
and abundant in seagrass beds and on rubble reef flats. It prefers areas affected by a strong
wave action and is gregarious in nature.
Biology
Description: The body of this species is elongated, cylindrical, and slightly arched dor-
sally (bivium); it is flattened ventrally (trivium). Bivium is covered by mucus and fine
sediment. Papillae on bivium are numerous, long, and slender. Long and thick podia are
seen on trivium and are arranged more or less regularly in tight rows on the radii. The
mouth is ventral and is surround by 20 stout tentacles. The anus is surrounded by five
strong, triangular anal teeth. Calcareous ring has large radial pieces and narrow inter-
radials. Cuvierian tubules are absent. Colour of bivium is black, and trivium is lighter and
dark brown. Spicules on ventral tegument are provided with a variety of simple or more
complicated rosettes. Dorsal tegument is with rosettes. Ventral and dorsal podia have some
rosettes. Podia and papillae contain large spicules. Tentacles have large rods. The species
has a maximum length of 35 cm and weight of 2 kg.
Food and feeding: It feeds mainly on epiphytes and seagrass leaves.

Antimicrobial activity: The alcoholic extracts of this species showed antimicrobial
activities against Aeromonas hydrophila (inhibition zone, distance from the border of
paper disc to the edge of the clear zone, 2.5 mm); Enterococcus sp., Escherichia coli,
Klebsiella pneumoniae, and Pseudomonas aeruginosa (1.3 mm); Vibrio harveyi (2.8 mm);
and Aspergillus sp. (4.5 mm) (Abraham et al., 2002).
Others: Honey-Escandón et al. (2015) reported on the isolation of triterpenoid glycosides
holothurins A, B from this species.
Bohadschia argus (Jaeger, 1833)
Common name(s): Leopard sea cucumber
Global distribution: This species is found in the Western Indian Ocean and the Pacific
Ocean. Its range extends from Madagascar, the Seychelles, and Sri Lanka to Malaysia and
the South Pacific Islands.
Ecology: It lives on coral reefs and on exposed, sandy areas of the seabed at depths of
between 10.3 and 37 m.
Biology
Description: The body of this species is sausage shaped with a smooth, tough, leath-
ery skin. Its colour is greyish brown and paler below, with distinctive dark eyespots sur-
rounded by white haloes. There are several rows of tube feet on the underside. Surrounding
the mouth at the anterior end is a ring of paddle-shaped, black tentacles fringed with white.
Cuvierian tubules are situated at its base and are readily ejected as sticky threads if the
animal is disturbed or handled. These tubules contain toxins that deter predators and are
irritating to human skin. This species can grow to 60 cm in length.
Food and feeding: It is an omnivore. As it moves across the seabed, it sweeps sand grains
and detritus into its mouth using its sticky tentacles. It obtains some nourishment from the
biofilm that coats the grains.
Associated species: Fish of the species Carapus mourlani are sometimes found living in
the coelomic cavity of this species. This fish enters through the anus, either going in head
first or more frequently tail first. The small emperor shrimp (Periclimenes imperator) is
also often associated with this species and may help keep it clear of ectoparasites.

Cytotoxicity: A holostane-based triterpene glycoside, arguside A, extracted from the
tissues of this species exhibited cytotoxicity against several different types of human
tumour cells, being more active towards HCT‐116 cells (IC50 = 0.14 μM) (Liu et al., 2007).
Antifungal activities: The triterpenoid oligosaccharides, bivittoside C and D, have

been reported from this species. However, only bivittoside D was found to exhibit
significant antifungal activities against Aspergillus niger, A. oryzae, Penicillium citrinum,
P. chrysogenum, Mucor spinescens, Cladosporium herbarium, Rhodotorula rubra,
Trichophyton mentagrophytes, T. rubrum, Candida albicans, and C. utilis (Emily, http://
www.freepatentsonline.com/y2005/0208147.html).
Bohadschia bivittata (Mitsukuri, 1912)
Common name(s): Two-ribboned sea cucumber
Global distribution: Tropical Western Central Pacific: Micronesia, Caroline Island, and
Philippines
Ecology: This benthic species is found in shallow water, usually in sandy seagrass beds and
reef flats.
Biology
Description: It has a maximum length of 25 cm. No other information is available on this aspect.
Food and feeding: It is a deposit feeder. It has diurnal burrowing and nocturnal epibenthic
feeding behaviour.

Antifungal activities: The methanolic extract of the Cuvierian tubles of this species afforded
triterpenoidal oligoglycosides, viz. bivittoside A, B, C, and D. Among these four bivittosides,
only bivittoside D was found to exhibit significant antifungal activities against Aspergillus niger,
A. oryzae, Penicillium citrinum, P. chrysogenum, Mucor spinescens, Cladosporium herbarium,
Rhodotorula rubra, Trichophyton mentagrophytes, T. rubrum, Candida albicans, and C. utilis
(Kitagawa et al., 1981a; Emily, http://www.freepatentsonline.com/y2005/0208147.html).
Bohadschia cousteaui (Cherbonnier, 1954)
Global distribution: Tropical Western Indian Ocean: Red Sea, Madagascar, Comoros,
and Kenya
Ecology: This benthic species is found on rocky bottom habitats.
Biology
Description: This species is rare.
Reproduction: It is a gonochoric species with only one gonad. Spawning and fertilization
are both external and some individuals exhibit brooding. Embryos develop into planktotro-
phic larvae (auricularia), then into doliolaria (barrel-shaped stage), which later metamor-
phose into juvenile sea cucumbers.

Antifungal activity: A total of 21 lanostane-type non-sulphated triterpene glycosides
have been isolated from the methanol/methylene chloride extract of the body walls of this
species. These compounds are ten new saponins (including two pentasaccharide and eight
hexasaccharide saponins) called coustesides A–J and eleven known triterpene glycosides.
Most of the isolated compounds showed good antifungal activity against Candida albicans
as detailed below (Elbandy et al., 2014).
Antifungal Activity of the Tested Saponin Mixtures

Mean Inhibition Zone
Compounds (1 mg/mL) Diameter (mm)a after 48 hours
1, 11 15.7
2, 13 15.4
3, 15 14.9
4, 16 14.7
5, 17 11.5
6, 18 10.7
9, 10 16.3
12 11.0
14 18.0
a The diameter of the filter paper disc (6 mm) is included.

Source: Elbandy, M. et al., Eur. Food Res. Technol., 238, 937–955, 2014.

Bohadschia marmorata (Jaeger, 1833)
Common name(s): Brown sandfish, chalky cucumber, pale marbled sea cucumber
Global distribution: Tropical Indo-Pacific Ocean: Its range extends from the Red Sea and
east coast of Africa to Japan, the Philippines and Australia.
Ecology: This reef-associated species occurs in coarse coral sand, from the intertidal to a
depth of 20 m. It is a burrowing form and comes out of the mud during cloudy days and at
night, being a nocturnal species. In particular, it prefers reef flats and seagrass beds down
to about 36 m. Small individuals are usually buried in the sand while larger specimens are
covered with a thin layer of sand.
Biology
Description: It is cylindrical in shape. The body wall is tough and leathery and has a rough
texture due to its calcareous spicules. The anterior end of the body is somewhat narrowed and
has a mouth surrounded by a ring of retractile tentacles. The posterior end is rounded and has
an anal opening. Adjoining this is the cloaca through which defensive white sticky threads
(Cuvierian tubes) may be ejected when the animal is stressed. The cloaca is also connected
to the respiratory tree, into and out of which water is pumped for gas exchange. The body
colour is cream or orange, partially obscured by variable numbers of dark brown speckles and
blotches. On the underside is a wide, pale-coloured longitudinal stripe and many short, slender
tube feet with knobbly tips. It has a maximum length of 40 cm and weight of 2 kg.
Food and feeding: It is a detritivore. It uses its tube feet to move around and scoops sand,
mud, and debris into its mouth with its oral tentacles. Having processed this material in its
gut, it voids the remains, leaving a characteristic trail of sand on the seabed.
Antifungal activity: Holostan-type triterpene glycosides—identified as marmoratoside A,
17α-hydroxy impatienside A, marmoratoside B, and 25-acetoxy bivittoside D (4)—have
been isolated from this species. Among these compounds, compounds 1 and 2 showed
moderate antifungal activity (Gomes et al., 2014).
Antifungal and cytotoxic activities: The methanol extract of the body wall and
water-methanolic extract of Cuvierian organ of this species displayed antifungal activ-
ity against A. niger with minimum inhibitory concentration (MIC/disc) of <1 mg. On
the other hand, the methanol extract of the body wall and water-methanolic extract of
Cuvierian organ of this species showed significant antifungal activities against C. albi
cans (MIC/disc = 3 mg) Significant inhibitory effect was observed in cytotoxic assays,
which was highest in Cuvierian organ methanol extract (LC 50 = 12 µg/mL) (Mokhlesi
et al., 2011).
Among the triterpenoid oligosaccharides, Bivittosides C and D reported from this
species, only Bivittoside D was found to exhibit significant antifungal activities against
Aspergillus niger, A. oryzae, Penicillium citrinum, P. chrysogenum, Mucor spinescens,
Cladosporium herbarium, Rhodotorula rubra, Trichophyton mentagrophytes,
T. rubrum, Candida albicans, and C. utilis (Emily, http://www.freepatentsonline.com/
y2005/0208147.html).
Antinociceptive activity: The water extracts as well as the coelomic fluid of this species
showed antinociceptive effect in mice (Bordbar et al., 2011).
Bohadschia subrubra (Quoy & Gaimard, 1834)
Common name(s): Leopardfish
Global distribution: Tropical Western Indian Ocean: Kenya (Samyn 2000), Comoros,
Madagascar, Mauritius, Tanzania, and Mayotte
Ecology: This benthic species lives over rubble in sandy bottoms or among seagrass and
around shallow reefs between 0 and 30 m. It remains buried during the day. Individuals are
most often covered with coarse coral and shell debris, or even pieces of algae scattered over
the bivium.
Biology
Description: It is a midsized to large sea cucumber with a maximum length of 40 cm. Its
body is subcylindrical with a trivium (ventral side), which is flat. Both ends of the body
are rounded. The integument is relatively thin (2–5 mm thick). The pattern of colours is
quite variable. The background colour can range from yellowish to bright orange and dark
brown. Black spots are seen on the bivium (dorsal surface), sometimes covering more
surface than the background colour. They vary in size, number, and location. Some are
occasionally surrounded by white. The mouth is ventral and is surrounded by 18 whitish
tentacles. The anus is dorsal and when it is open, the cloacal orifice can take the form of a
five-pointed star.
Food and feeding: It is limnivorous, which means that it ingests the sediment and retains
the nutrients of plant and animal origin, waste and bacteria contained therein. It thus par-
ticipates in the balance of the coastal ecosystems both by elimination of organic debris and
by reworking of the substrate. It feeds at night.
Reproduction: Fertilization in this species takes place at the chance of encounters of the
gametes in the current. Larvae are pelagic. The larval stages (blastula, then auricularia,
then doliolaria) take place in open water. At the end of the last stage, the juvenile definitively
joins the substratum and evolves towards adulthood.
Parasite: This species can be a victim of external parasites, including gastropods of the
family Eulimidae.

Haemolytic activity: The saponins, viz. holothurinoside J1 and holothurinoside K1
isolated from the body wall of this species, exhibited weak haemolytic activity (Gomes
et al., 2014).
Others: Mondol et al. (2017) reported on the isolation of compounds, viz. holothurinoside
H, I, K1.
Bohadschia vitiensis (Semper, 1868) (= Bohadschia tenuissima; Holothuria bivittata )
Common name(s): Brownspotted sandfish, brown sandfish
Global distribution: It is widespread in the Indo Pacific. It occurs from East Africa
(Madagascar) to India, Ceylon, Bay of Bengal, East Indies, North Australia, Philippines,
China, South Japan, and South Pacific Islands.
Ecology: It lives in coastal lagoons and inner reef flats and among the seagrasses. It is
generally buried in sandy, muddy bottoms during the day and only emerges at dusk. Depth
range is 0–20 m.
Biology
Description: The body of this species is oval. It is usually yellowish with lots of little
brown spots and tiny, dark tube feet and has 20 short, yellowish feeding tentacles with
bushy tips. When disturbed, even slightly, it releases sticky, white cylindrical tubes called
Cuvierian tubules. It has a maximum length of 30 cm.
Edible values: It is considered a delicacy or a protein component in traditional diets.


Antifungal activity: The methanol extract of this species showed promising antifungal activity
in in vitro models against Candida albicans, Cryptococcus neoformans, Sporothrix schenckii,
Trichophyton mentagrophytes, Aspergillus fumigatus, and Candida parapsilosis. The aqueous
fraction with its compound bivittoside-D had antifungal activity against C. albicans, C.
neoformans, S. schenckii, T. mentagrophytes, and A. fumigates (Lakshmi et al., 2012).
Emily (http://www.freepatentsonline.com/y2005/0208147.html) reported on the presence
of triterpenoid oligosaccharides, Bivittosides C and D from this species. However, only
Bivittoside D was found to exhibit significant antifungal activities against Aspergillus niger, A.
oryzae, Penicillium citrinum, P. chrysogenum, Mucor spinescens, Cladosporium herbarium,
Rhodotorula rubra, Trichophyton mentagrophytes, T. rubrum, Candida albicans, and C. utilis.
Other uses: The compound bivittoside (saponin) of this species may be employed as
a local antifertility agent and a single sea cucumber drug in the form of contraceptive
cream, jelly, or water-soluble film to prevent pregnancy in women and as a fungicide
cream. The use of contraceptive creams, jellies, or water-soluble film containing a
spermicide provides an easy and convenient method of preventing pregnancy in women.
Most of the currently used spermicidal agents are either costly or cause undesirable
side effects e.g., irritation, eczema, dermatitis, or skin rash. Besides, the fungicidal
activity of the saponin, especially that against Candida albicans, can offer prophylactic
contraception as C. albicans causes a very common vaginal infection in the human
population. The saponin can also be utilized as a fungicide cream or jelly (Emily, http://
www.freepatentsonline.com/y2005/0208147.html).
Holothuria (Halodeima) atra (Jaeger, 1833)
Common name(s): Black sea cucumber, lollyfish
Global distribution: Tropical Indo-Pacific region: its range is extending from the Red Sea
and East Africa to Australia.
Ecology: It lives on the seabed, in shallow waters on reefs and sand flats and in seagrass
meadows at depths of up to 20 m.
Biology
Description: It has a smooth, pliable, entirely black skin that often has sand adhering
to it, especially in smaller individuals. The mouth is on the underside at one end and is
surrounded by a fringe of 20 black, branched tentacles. The anus is at the other end. It can
grow to a maximum length of 60 cm.
Food and feeding: It is an omnivore, sifting through the sediment with its tentacles and
feeding on detritus and other organic matter.
Behaviour: As a defence against predators, this species emits a toxic red fluid when its skin
is rubbed or damaged. When attacked, it does not eject Cuvierian tubules in the way that
some sea cucumbers do, but instead it extrudes its internal organs through its anus. Further,
it is very often camouflaged by a coating of sand, which may also serve to keep it cool by
protecting it from the sun’s rays.
Reproduction: In this species it is not possible to distinguish between male and female
externally. Maturity is reached at a body length of about 16 cm. Spawning mostly takes place
during the summer and autumn, although in equatorial waters it may take place all year round.
Holothuria atra is also fissiparous, meaning that it can reproduce by transverse fission.
Associated species: It is often found associated with the polychaete worm Gastrolepidia
clavigera, a black worm that crawls about over the sea cucumber’s skin. Lissocarcinus
orbicularis, a small crab, is known to live on this species in a commensal relationship.

activities against Aeromonas hydrophila (inhibition zone, distance from the border of paper
disc to the edge of the clear zone, 2.0 mm); Enterococcus sp., Klebsiella pneumoniae,
Pseudomonas aeruginosa, and Aspergillus sp. (1.25 mm); and Vibrio harveyi (1.0 mm)
(Abraham et al., 2002).
Antioxidant activity: Mixed extract of this species containing some phenolic components,
such as chlorogenic acid, pyrogallol, rutin, coumaric acid, catechin, and ascorbic acid
exhibited a high scavenging activity for NO∙ (93.42% at 600 mg/mL) and a low scavenging
activity for DPPH (16.8%–17.01%) at all tested concentrations. For iron chelating activity,
the extract showed moderate potential (57% at 600 mg/mL). Further, this extract also
showed a dose-dependent inhibition of lipid peroxidation (Esmat et al., 2013).
Antileishmanial activity (antiprotozoal activity): The dichloromethane/methanol extracts
of this species showed antileishmanial activity against Leishmania amazonensis (Oliviera
et al., 2016).
Holothuria (Microthele) fuscopunctata (Jaeger, 1833) (= Holothuria axiologa)
Common name(s): Elephant trunkfish

Global distribution: This species is found in the tropical Indo-Pacific. Is range extends
from Madagascar and the eastern coast of Africa to Japan, China, Australia and the
Mariana Islands, Palau, and New Caledonia.
Ecology: This shallow-water species inhabits sandy seabeds and seagrass meadows, reef
slopes, and on coral rubble. It is often exposed on deeper fine sediments. Its depth range is
down to a maximum of 30 m.
Biology
Description: The body wall of this species is thick and golden brown with darker spots and dark
brown wrinkles. The underside is whitish. Spicules are stout, square tables, and fenestrated
ellipsoid buttons. The mouth is surrounded by 20 thick, brown tentacles, and the anus is
surrounded by a black band. The cloaca is large and black, but there are no Cuvierian tubules.
It is a large sea cucumber reaching a maximum length of 70 cm and weight of 4.5 kg.
Food and feeding: It feeds on organic material that it grazes on the seabed.
Reproduction: It is slow to mature and has low fecundity. The sexes are separate and
breeding takes place in the warm season, between December and January.

Antibacterial activity: The methanolic, aqueous, choloroform, and hexane extracts of this
species showed antibacterial activity against E. coli as detailed below (Layson et al., 2014).
MIC (µg/mL) after 4/8/24 hours of Exposure

4 8 24
Methanolic extract 31 816 0.5
Aqueous extract 17 8072 11755
Choloroform extract 194 250 6952
Hexane extract 16 29 <0.01
Source: Layson, R.J. et al., J. Trop. Life Sci., 4, 175–181, 2014.
Antifungal activity: Among the compounds holothurin B, arguside F, impatienside B, and

pervicoside D isolated from this species, the compound impatienside B displayed antifun-
gal activity (Yuan et al., 2009; Gomes et al., 2014; Bahrami and Franco, 2016).
Holothuria (Panningothuria) forskali (Delle Chiaje, 1823)
Common name(s): Black sea cucumber, cotton-spinner
Global distribution: Atlantic coasts of northwest Europe, Canary Islands, Azores, and
Mediterranean Sea
Ecology: It is found on boulders and rocks, particularly vertical surfaces, from the intertidal
zone down to a depth of about 50 m.
Biology
Description: This sea cucumber has a cylindrical body and can grow to 30 cm long.
It is usually deep brown or black but sometimes has an underlying yellowish mottling,
especially on the underside. The skin is soft, coarse, and tough and it is covered with fleshy
papillae, which are often tipped with white. The underside has three rows of tube feet
while the upper side has two rows of rudimentary suckers. The anterior end has a bunch of
20 yellowish, short, retractable tentacles encircling the mouth. At the posterior, inside the
body cavity, there are a bundle of Cuvierian tubules or cotton glands that can be ejected as
a tangle of sticky white threads.
Food and feeding: The black sea cucumber is a detritivore and feeds mostly at night. When
feeding, it bends its body down towards the substrate, presses the ring of oral papillae against
the surface, and opens its mouth wide. It then extends and retracts its short tentacles repeatedly
and extracts the nutritious part and deposits the unmetabolized portion as droppings.
Reproduction: Adult black sea cucumbers are normally either male or female. The gonads
take a long time to mature, and gametes are released synchronously into the water column
in early spring. The larvae become part of the zooplankton. After several moults they grow
tentacles and settle out onto the seafloor.
Parasite: The parasitic copepod Asterocheres boecki is an endoparasite of this species.

Antitumour and antiviral activities: The triterpene glycosides, viz. holothurinosides A, B,
C, and D, desholothurin A, and a related saponin have been isolated from this species. Among
these compounds, holothurinoside A and desholothurin A displayed strong antitumour and
antiviral activities. The IC50 values recorded for antitumour activity of P388 cells for these
compounds were 0.46 and 0.38 mg/mL, respectively (Rodriguez et al., 1991).
Cytotoxic activity: Mondol et al. (2017) reported that the compounds holothurinosides A,
C of this species were found to be cytotoxic against human tumour cell lines with IC50
values of 0.33–0.71 µg/mL and 0.16–0.93 µg/mL, respectively.
Holothuria fuscocinerea (Jaeger, 1833)
Common name(s): Ashy pink sea cucumber

Global distribution: This species is found in tropical and subtropical Indo-Pacific; its
range extends from East Africa and the Red Sea to southern Japan, northern Australia, and
Central America.
Ecology: This shallow-water species is typically found on coral rubble and soft sediments,
and among seagrasses at depths of less than 40 m.
Biology
Description: This sea cucumber has an oval, cylindrical body that is flattened somewhat
on the underside and rounded at both ends. It can reach a maximum length of 70 cm and a
weight of 5.5 kg. The mouth is on the underside at the front and is surrounded by 20 short,
brown, branched feeding tentacles. The anus is dark brown or black and has no anal teeth,
but it is surrounded by five white-tipped papillae. Cuvierian tubules are present in this
species. The skin is soft, thick, and wrinkled, often with sand adhering to it. Spicules are
sometimes poorly formed and include low tables with a spinose disc and irregular buttons.
It varies in colour as its upper side is usually greyish or brownish, while the underside is a
dirty white. There is often a series of dark bars in the transverse wrinkles.
Food and feeding: It is a detritivore and feeds at night, sifting through the sediment with
its feeding tentacles, swallowing the sediment and extracting the bacteria and organic
material it contains, while the sand passes on through its gut.
Reproduction: In this species, breeding takes place in the hot season. The sexes are
separate. The individuals climb to an elevated location before releasing their gametes into
the water column. Females produce an average of around three million eggs. The larvae
are pelagic, drifting in the plankton. When sufficiently developed, these larvae settle on
the seabed and undergo metamorphosis into juveniles.

Cytotoxicity: The triterpene glycosides, fuscocineroside A, B, and C; pervicoside C; and
holothurin A isolated from this species showed potent cytotoxicity on human leukaemia
HL-60 and human hepatoma BEL-7402 cell lines. However, fuscocineroside C was found
to be the most potent (IC50 = 0.88, IC50 = 0.58 µg/mL) in HL-60 and BEL-7402 cell lines,
respectively (Li et al., 2013).
Zhang et al. (2006d) reported that the n-BuOH extract of this species showed the presence
of three triterpene glycosides, viz. fuscocinerosides A (1), B (2), and C (3), along with two
known glycosides, pervicoside C (4) and holothurin A (5). All these glycosides showed
in vitro cytotoxicity against two human tumour cell lines.
Mondol et al. (2017) reported that the compound, fuscocineroside C, of this species showed
cytotoxicity against two human tumour cell lines with IC50 values of 0.88 and 0.58 µg/mL,
respectively.
Holothuria (Microthele) fuscogilva (Cherbonnier, 1980)
Common name(s): Black teatfish, white teatfish
Global distribution: Africa and Indian Ocean: Madagascar, East Africa, Seychelles,
India, Sri Lanka, the Maldives, Comoros, and the Red Sea
Ecology: It is largely restricted to coral reef habitats. It also occurs in sandy hard substrate
and shallow seagrass beds, and it is generally solitary. Depth range of this shallow-water
species is from 0 to 40 m.
Biology
Description: The body of this species is massive and tubular in shape. Eight teat-like
projections are seen on each side of the body hence the common name teatfish or mammyflsh.
The body wall is 10 mm in thickness. Pedicels and papillae are indistinguishable. Dorsal
papillae are sparsely scattered than the ventral pedicels. The anus is surrounded by five
calcified papillae. Calcareous ring is massive with a distinctly scalloped anterior margin.
The radials and interradials are squarish. Radials are twice the length of interradials.
Tentacular ampullae are very large and Cuvierian tubules are absent. The colour in life
is yellowish white. The body surface is covered with a fine coating of coral sand. It has a
maximum length of 400 mm and a live weight of 2.5 kg.
Food and feeding: This species feeds on the fine layer of sediment or algae on reef surfaces.
Reproduction: This species has been reported to mature at four years. It is characterized
by a sexual reproduction that takes place during the cold season. The juveniles are very
rarely seen and have a different colour.

Holothuria (Halodeima) edulis (Lesson, 1830)
Common name(s): Pink and black sea cucumber, pink fish sea cucumber
Global distribution: This species is found in the tropical Indo-Pacific Ocean, from the
Red Sea and East African coast to Sri Lanka, Japan, China, Indonesia, the Philippines,
northern Australia, and various Pacific islands.
Ecology: This benthic, shallow-water species has different habitats including on sandy
or muddy substrates; coral rubble, outer reef flats, back reef slopes and in lagoons; and in
seagrass meadows at depths down to 20 m.
Biology
Description: It is a medium-sized sea cucumber reaching a maximum length of about
30 cm. It has a roughly cylindrical shape with rounded ends but can retract and expand its
body and adopt different shapes. It is usually soft and pliable with a smooth skin. But due
to its special characteristics in connective tissue, it can become firm and rigid. The body is
lined with longitudinal rows of small tube feet, which can be withdrawn into the body wall,
leaving small hollows. There are about 20 tube feet in a ring round the mouth, and these
organs are modified into feeding tentacles. This sea cucumber is usually a dark reddish-
black colour on its upper side and a pinkish mauve colour below.
Food and feeding: Holothuria edulis is mainly nocturnal and tends to hide during the day
under rocks or pieces of coral. It is a detritivore and feeds by ingesting sand and debris
accumulated on the seabed. Sand is pushed into the mouth, and any organic matter present
is digested as the sand passes through the gut. The indigestible matter is expelled from the
anus, leaving a sand ridge as the animal moves around. During its feeding activities, the
sea cucumber churns up the top few centimetres of seabed and aerates the sediment.
Reproduction: It has separate sexes and spawns at any time of year with gametes being
liberated into the water column. The larvae are planktonic. It can also reproduce asexually
by breaking into two parts, each of which then regrows the missing organs.
Edible values: It is edible and is dried and sold as “bêche-de-mer” or “trepang” in China
and Indonesia.
Aquarium values: It is common in the aquarium trade for its colours and its ability to eat
detritus. It needs a tank size of 50 gallons or more. It is easy to care for but may release a
toxin when stressed, although this only happens very rarely.

Antibacterial activity: The water extracts of this species have shown antibacterial activity
(Dobretsov et al., 2009).
Anticoagulant activity: The sulphated fucan, fucosylated chondrotin sulphate, isolated
from this species has shown anticoagulant activity (Bahrami, 2015).
Holothuria (Halodeima) grisea (Selenka, 1867) (= Ludwigothuria grisea)
Common name(s): Gray sea cucumber
Global distribution: Its range is from Florida (although not reported in the Florida Keys)
to the Bahamas, Puerto Rico, Jamaica, the Lesser Antilles to Curaςao, Panama, Colombia,
southern Brazil, and West Africa. This species may be a seasonal resident of coastal Texas.
Ecology: It inhabits seagrass beds and sand flats. These individuals are found intertidally
to a depth of less than 5 m.
Biology
Description: The body of this species is subcylindrical with a flattened sole covered in
cylindrical tube feet. In adults, six rows of papillae are found on the surface of the upper
body. Juveniles, on the other hand, have four rows. The papillae originate from large warts
and each one is surrounded by 5–10 small tube feet. The mouth is positioned downward
slightly and is encircled by 20–25 bushy peltate tentacles. The gray sea cucumber has
a more striking colour pattern. Base colours include variations of red or yellowish red,
marked with brown mottling. Papillae on the upper body are white with yellow tips; the
feet on the sole are tipped in yellow, and the tentacles are yellow. This species can attain a
maximum length of 25 cm.
Food and feeding: It is primarily a deposit feeder, gathering organic matter and sediments
with its branched feeding tentacles.

Cytotoxic activity: Two lanostane‐type nonsulfated pentasaccharide triterpene glyco-
sides, 17‐dehydroxyholothurinoside A and griseaside A isolated from this species, exhib-
ited significant cytotoxicity against HL‐60, BEL‐7402, Molt‐4, and A‐549 cancer cell lines
(Sun et al., 2008).
Antithrombotic and antitumour activities: The fucoidans—viz. fucan sulphate,

glucosamine, and chondroitin—of this species have shown antithrombotic and antitumour
activities besides regulating angiogenesis (Bahrami, 2015; Mondol et al., 2017).
Holothuria (Mertensiothuria) hilla (Lesson, 1830) (= Holothuria (Thymiosycia)

decorata)
Common name(s): Tiger tail sea cucumber
Global distribution: Tropical Indo-West Pacific: Kenya to India, Indonesia, the Philippines,
China, and Japan; to Hawaii, Mexico, Central America, and the Galapagos Islands
Ecology: It is found under rocks on the bottom of coarse sand, generally between 0 and
20 m but also to over 200 m.
Biology
Description: It has a body that is long and cylindrical with blunt ends. The mouth is ven-
tral in position and is surrounded by 20 small yellowish tentacles; the anus is terminal.
Ventral podia are arranged in 3–4 rows in each ambulacra area. Dorsal papillaeare are
large and conical and are scattered on the bivium. The spicules of the papillae are buttons
with three or five pairs of holes (60–100 mm) and tables (80 mm across). The spicules of
the body wall are tables (80 mm across) and buttons (100 mm). The spicules of tentacles
are 130–200 mm long, and the body wall has tables and buttons as well as distinct papillae.
The background colour of this species is reddish brown with numerous white spots mark-
ing the base of creamy white buds. It reaches a length of 200 mm. The body wall, which is
soft, is light brown in colour with scattered white conical papillae.
Food and feeding: It feeds largely on sediments.

Cytotoxic activity: Mondol et al. (2017) reported on the isolation of hillasides B, C from
this species. Among these compounds, hillaside C has shown cytotoxic activity against
human tumour cell lines with IC50 values of 0.15–3.20 µg/mL.
Holothuria (Thymiosycia) impatiens (Forsskål, 1775)
Common name(s): Impatient sea cucumber

Global distribution: It is widespread throughout the tropical Indo-Pacific and subtropical regions
(including in the Red Sea and Persian Gulf) and is also known from the Mediterranean Sea.
Ecology: It lives in coral reefs and among rocks in shallow waters (0–2 m) but can be
observed up to 30 m. When touched or disturbed, the animal very quickly contracts in size,
an adaptive response to escape from predators and living in confined spaces.
Biology
Description: In this species the body is bottle (cylindrical) shaped with a long neck and
rough surface, sandy to the touch. It is covered with conical warts from which filamentous
appendages emerge. The ventral mouth is surrounded by 20 tentacles that are in turn
surrounded by a ring of very small conical papillae. The anus is terminal with five conical
papillae. Tentacles have curved sticks and are rough to the ends. The spicules of the dorsal
and ventral body wall are buttons and tables. Podia have tables (identical to that of the
seed coat) and buttons, which are perforated elongated sticks. Thick Cuvierian tubules
are present but are not readily ejected. The tube feet are short and sparsely arranged on
ambulacral and interambulacral areas. The species is light brown in colour with 4–5 dark
brown transverse bands on the upper side near the anterior end. Background colour is beige
with brown dorsal spots that are more or less dark. The trivium (side or face of the animal
with which the animal stands) has light brown with some lighter areas corresponding to
the base of the tube feet. The bivium (dorsal face of the animal, against the standing base)
is somewhat darker with dark brown areas. The species has a size range of 150–400 mm.
Food and feeding: At night it extends its anterior end from its hole to feed on nearby sediments.

Cytotoxic activity: The alcohol extracts containing lanostane-type triterpene glycoside,
impatienside A and bivittoside D, have shown in vitro cytotoxicities similar to or better
than those of the potent anticancer drug etoposide (V-16) against seven different human
tumour cells, with IC50 values of 0.37–2.75 µg/mL (Sun et al., 2007).
Holothuria (Metriatyla) lessoni (Massin, Uthicke, Purcell, Rowe & Samyn, 2009)
Common name(s): Golden sandfish, Australian sea cucumber
Global distribution: Western Central Pacific: Asia and Africa and Indian Ocean
Ecology: It occurs on sandy and muddy areas, reef flats, inner slopes lagoons, and seagrass
beds between 0 and 25 m.
Biology
Description: The body of this species is slightly flattened, with distinct ventral sole and
rounded extremities. Mouth is ventral, surrounded by 20 large tentacles. Papillae around
mouth are prominent in live specimens. The anus is terminal, without anal teeth but with
five groups of anal papillae. Dorsal surface of the body is brown beige, with or without
black patches, which are sometimes surrounded by a white margin; some specimens are
completely black or mottled with black and beige brown. Tube feet are as brown or black
dots covering bivium. Ventral surface is white to beige or grey in the black colour variants,
with deep longitudinal, median groove, darker than the rest of the surface. Tube feet are
brown or black, spread evenly over trivium. Body wall is smooth to the touch and is with
10–15 mm thickness (in contracted specimens). It is a large holothurian, and living adults
reach a size of 25–48 cm long.
Uses: It has the highest commercial value due to its taste.

Saponins and triterpene glycosides, viz. Holothurin B3, Holothurin, Holothurin B, Nobiliside
B, Holothurin B2, Leucospilotaside B, Leucospilotaside A, DS-pervicoside B, Bivitoside
B, Holothurinosides C/C1, Holothurinosides X/Y/Z, Desholothurin A (Nobiliside 2a),
Desholothurin A1 (Arguside E), Holothurinoside T, Holothurinoside J, Arguside A, Arguside D,
Intercedenside A, Fuscocinerosides B/C, Scabraside A or 24–dehydroechinoside A, Holothurin
A2, Echinoside A, Pervicoside B, Holothurin A, Scabraside B, 17-Hydroxy fuscocineroside B,
25-Hydroxy fuscocinerosiden B, Holothurin A1, Holothurin A4, Scabraside D, Holothurin A3
Holothurin D, Cousteside G, Impatienside B, Cousteside H, Cousteside J, Fuscocineroside A,
Holothurinoside E, Holothurinoside E1, Holothurinoside O, Holothurinoside, 17-dehydroxy
holothurinoside A, Cousteside E, Cousteside F, 22-acetoxy-echinoside A, Griseaside A
Cousteside I, Holothurinoside M, Holothurinoside, Holothurinoside A1, Holothurinoside Q,
Holothurinoside S, Holothurinoside R, Holothurinoside R1, Cousteside C, Holothurinoside
L, Arguside F, Impatienside A, Marmoratoside A, Bivittoside D, Holothurinosides H/H1,
Holothurin C, Cousteside A, 17α-hydroxy impatienside A, Marmoratoside B, Argusides B/C,
Lessoniosides A/B/C/D/E, Holothurinosides I/I1, Holothurinoside K1, 25-acetoxy bivittoside
D, and several unidentified compounds isolated from the ethanolic extraction of the viscera of
this species hold promise for biotechnological applications (Bahrami et al., 2014a, 2014b).
Holothuria (Mertensiothuria) leucospilota (Brandt, 1835)
Common name(s): Black sea cucumber, black tarzan
Global distribution: Indo-Pacific; east coast of Africa and northeast coast of Australia
Ecology: This shallow-water species is found on reefs and rocky coasts; the organisms are
often partly concealed under boulders.
Biology
Description: It is roughly cylindrical, tapering towards the posterior end. At the anterior
end, there are 20 oral tentacles with branched tips. These surround the mouth, which is
on the underside of the body. The animal is soft and pliable and is covered with fleshy
papillae. Its usual colour is charcoal grey or reddish black with pale grey tube feet on the
underside, but it may be bright or dark brown with white patches that are larger towards the
posterior end. If threatened, this species can emit a mass of fine sticky Cuvierian tubules
from its anus, which ensnare the potential predator and allow the sea cucumber to escape.
It is capable of regenerating these tubules in 15–18 days. It is a medium-sized sea cucum-
ber, reaching a length of up to 40 cm when relaxed but can stretch to about a metre when
extended.
Food and feeding: It is a scavenger. When feeding, it usually has its posterior end anchored
underneath a rock or in a crevice so that it can contract back out of sight if disturbed. It
feeds by using its tentacles to shovel organic debris lying on the seabed into its mouth. In
the process it swallows a significant quantity of sand, which passes through the gut.
Reproduction: It spawns more than once in a year i.e., during the post-monsoon (October
to January) and in monsoon periods (June to September). The spawning of this species
appears to be influenced by low temperature and salinity.
Associated species: The worm pearlfish (Encheliophis vermicularis) is a parasite of this

species and each parasitized H. leucospilota will host a male and female pair of the fish,
which live inside its body.

Antimicrobial and cytotoxic activities: The methanol extract of Cuvierian organ of this spe-
cies has shown antifungal activity against A. niger (minimum inhibitory concentration (MIC/
disc) <1 mg); the water-methanol extract of coelomic fluid was against C. albicans (MIC/
disc = 1 mg). Further, significant inhibitory effect was observed in cytotoxic assays, which
was highest in body wall methanol extract (LC50 = 0.4 μg mL−1) (Mokhlesi et al., 2012).
Cytotoxicity: The crude saponin extracted from this species showed marked dose-
dependent effect on viability of B16F10 cancer cells with an IC50 value of 10 µg/mL
(Baharara et al., 2016).
Gomes et al. (2014) reported on the isolation of compounds, viz. leucospilotaside
B, holothurin B2, and echinoside B from this species. Among these compounds,
leucospilotaside B exhibited moderate cytotoxicity against human tumour cell lines (HL-
60, MOLT-4, A-549, and BEL-7402).
Haemolytic and cytotoxic properties: The crude saponin extracted from this species was
cytotoxic to A549 cells at a concentration of 1 μg/mL after 48 hours of treatment. Further,
the 80% ethanol fraction of saponin isolated from it exhibited haemolytic activity and
offers promise as an anticancer candidate (Soltani et al., 2014).
Antioxidant and antiproliferative effects: The dichloromethane fraction of this species

exhibited antioxidant activities to scavenge DPPH. The reducing capacity enhanced with
increasing concentration of the extract and the activity was comparable with the butylated
hydroxyanisole (BHA) as a standard. Further, the dichloromethane fraction exhibited
activity against MCF-7 and A549 cancer cell lines (Soltani and Baharara, 2014).
Holothuria (Holothuria) mammata (Grube, 1840)
Global distribution: Subtropical Atlantic and the Mediterranean Sea including Spain,
Greece, Turkey, Tunisia, Croatia and Italy; Gulf Mexico, southern Portugal and the Azores
and Canary Islands; Algeria and Malta
Ecology: It occurs below 30 m; this benthic species inhabits sand and silt and rocky bot-
tom areas; the depth range is 1–77 m. It stays in crevices and holes during the day and
leaves them in the night for feeding. It can also be found in unusual habitats such as sea-
grass, with diurnal and nocturnal feeding.
Biology
Description: It is more purple and darker than the others. There are thick podia at the outer
surface of the animal. The Cuvierian tubules of this species are few in number, small, and
never expelled. It attains a maximum length of 15 cm.
Reproduction: The spring period, from March to April, corresponds to its maturation and
subsequent spawning. It showed five typical larval stages of most aspidochirote holothuri-
ans. Juvenile stage was reached after 21 days post-fertilization.
Aquaculture value: This species shows high potential for its aquaculture.
Other commercial uses: It is an edible species, and its fishery in Turkish waters started in
1996. Harvested individuals are being exported to Asian countries as frozen, dried, and
salted products.

As functional foods and nutraceuticals: Its total fat content was 1.00% and highest values
of fatty acids were obtained for stearic acid (C18:0) (8.91%), arachidonic acid (C20:4 ω6)
(19.97%), and eicosapentaenoic acid (C20:5 ω3) (10.85%). These findings suggest that this
species has potential uses as functional foods and nutraceuticals (Santos et al., 2017).
Holothuria (Microthele) nobilis (Selenka, 1867) (= Holothuria nobilis)
Common name(s): Black teatfish
Global distribution: Africa and Indian Ocean region: Madagascar, East Africa, Seychelles,
India, Sri Lanka, the Maldives, Comoros, and the Red Sea
Ecology: It is distributed mainly in shallow coral reef areas, on reef flats, slopes, and
shallow seagrass beds; it prefers sandy hard substrate. Depth range is 0–40 m.
Biology
Description: It is a black sea cucumber, often with a fine sand covering. It is a large, stout
species, firm and rigid, with prominent lateral papillae and anal papillae (usually five). It
has translucent Cuvierian tubules. Spicules are stout tables and fenestrated ellipsoids.
Reproduction: In this species, sexual reproduction takes place during the cold season. The
juveniles are very rarely seen and have a different colour.

Antibacterial activity: The methanolic extracts of this species have shown antibacterial
activity against E. coli with MIC (µg/mL) values of 40,116, < 0.01 in 4/8/24 hours of
exposure, respectively. In aqueous extracts, the MIC values were found to be 1431, 21,76 µg/
mL respectively; in choloroform extracts, 179.369, 340 µg/mL, respectively; and in hexane
extracts, 1182, 8004, 3041 µg/m, respectively (Layson et al., 2014).
Antitumour activity: The triterpene glycoside, nobiliside D, isolated from this species has
shown antitumour activity against human leukaemic cell line (K562), human leukaemia
cell line (U937), human lung cancer cell line (A-549), human cervix carcinoma cell line
(HeLa), human breast cancer cell line (MCF-7), and human liver carcinoma cell line
(HepG2). The above compound exhibited the greatest inhibitory effect on K562 and MCF-7
cells with IC50 values of 0.83 and 0.82 µg/mL, respectively. When human tumour cell
lines K562 and MCF-7 were treated by this compound, (0.5 µg/mL) for 24 hours, 45.8%
of K562 cells and 58.7% of MCF-7 cells were apoptotic, whereas only 0.5% of untreated
control cells were apoptotic. These data indicate that nobiliside D should offer potential as
a novel drug for the treatment of a range of cancers (Zhang and Zhu, 2017).

Holothuria (Stauropora) pervicax (Selenka, 1867)
Common name(s): Stubborn sea cucumber
Global distribution: This species is found in tropical Indo-West Pacific, from Madagascar,
Kenya, and Tanzania to India, Australia, and China, and eastward to Hawaii, including the
Maldives, Micronesia, and French Polynesia.
Ecology: This benthic, near-shore species lives in the inner reef flats, most frequently
in seagrass beds and coral rubble to depths of about 20 m. It becomes active at night.
Disturbance causes this species to eject copious amounts of Cuvierian tubules.
Biology
Description: It is a hemicylindrical species and is somewhat flattened below. It has a soft
and pliable body, a smooth tegument, and thick, translucent Cuvierian tubules. Spicules
are rather poorly formed, and their buttons often resemble small beaded rods. It is a grey/
brown sea cucumber, mottled above, with red-brown papillae. It is a medium-sized species
with a maximum length of 35 cm.
Food and feeding: It is a conveyor belt-feeder, meaning it ingests the substrate’s surface
and defecates on it.

Antifungal activity: Desulfated lanostane-type triterpene tetraglycosides, designated
DS-pervicoside A, DS-pervicoside B, and DS-pervicoside C extracted from the body walls
and the Cuvierian tubules of this species displayed antifungal activity as detailed below
(Kitagawa et al., 1989).
MIC (µg/mL)
DS-pervicoside A DS-pervicoside B DS-pervicoside C
Aspergillus niger 12.5 1.6 6.3
Aspergillus oryzae 12.5 3.1 6.3
Penicillium chrysogenum 6.3 1.6 3.1
Penicillium citrinum 6.3 1.6 6.3
Mucor spinescence 12.5 6.3 12.5
Cladosporium harborum 50.0 6.3 12.5
Rhodotorula rubra 100.0 3.1 6.3
Trichophyton mentagarophytes 12.5 1.6 6.3
Trichophyton rubrum 12.5 1.6 6.3
Candida albicans 100.0 6.3 12.5
Candida utilis 100.0 6.3 12.5
Source: Kitagawa, I. et al., Chem. Pharm. Bull., 37, 1230–1234, 1989.

Holothuria (Roweothuria) poli (Delle Chiaje, 1824)
Common name(s): White spot cucumber
Global distribution: Mediterranean including Algeria, Croatia, Italy, Spain (also Canary
Islands), France (also Atlantic coast) and Turkey; and Gulf of Suez in the northern Red Sea
Ecology: It is found in rocky substratum and rocky cavities, and seagrass meadows; depth
range is 5–22 m.
Biology
Description: The body of this species is cylindrical with a ventral sole that is nearly flat.
The anterior end is narrower than the posterior. It has, at one end, a mouth surrounded
by 20 short retractile tentacles, ending in a star, and a cloacal orifice at the other end. Its
skin is thick and very rough. Its dorsal surface is speckled with black, against a variegated
background of brown, grey, or light. The dorsal surface is covered with conical, irregular,
slightly prominent tubercles surmounted by small white papillae. The distribution of these
tubers is irregular. The feet secrete sticky mucus, which allows the animal to cover with
a small film of sand or various organic debris (fragments of dead leaves of Posidonia).
The ventral side, greyish brown, has many nonaligned ambulacral feet. The difference in
colour between the upper and lower faces is weak. It measures about 25 cm long and 5 cm
in diameter.
Food and feeding: This species is limnivorous and swallows the soft substrate and sorts
the organic particles.
Reproduction: It is sexual. Fertilization is done once a year from November to January,
by free emission in water. Each one recovers in “L” to emit ovules or spermatozoa in a
white cloud. The larva lives a long planktonic phase in open water. The first larval stage
is the auricularia stage and successive metamorphoses will give larvae doliolaria (barrel
shaped), then pentactula (appearance of tentacles), and finally adults.
Parasite: The gastropod Entoconcha mirabilis is one of its parasites.

Triterpene glycosides—monosulfated biosides, viz. holothurins B2 (1), B3 (2), and B4 (3)—
along with the previously known holothurins A (4) and B (5) have been isolated from this
species. Though these compounds serve as chemotaxonomic markers, their pharmaceuti-
cal uses/activities are yet to be known (Silchenko et al., 2005; Mondol et al., 2017).
Holothuria (Metriatyla) scabra (Jaeger, 1833)
Common name(s): Sandfish, garlic bread sea cucumber
Global distribution: Throughout the Indo-Pacific region
Ecology: It inhabits soft sediments, reefs and coastal areas; it may be exposed or partially
buried in sand or seagrass flats.
Biology
Description: This loaf-shaped sea cucumber has a rather flat, square-ended, loaf-shaped
body with a distinct upper and underside. The upper side is darker and often has little folds
and sometimes black bars (hence the name garlic bread sea cucumber). The underside is
flat and pale or white. There are little tube feet regularly distributed all over the body. It
grows up to 40 cm long.
Food and feeding: It feeds on and gathers detritus with the 20 or so short feeding tentacles
that surround its mouth, which usually faces downwards towards the ground.
Associated species: Pea crabs (Pinnotheres sp.) are sometimes found living in their rear ends.
Parasite: The worm pearlfish (Encheliophis vermicularis) is a parasite of this species and each
parasitized H. scabra will host a male and female pair of the fish, which live inside its body.
Edible values: This species is harvested and processed into “beche-de-mer” and eaten in
China and other Pacific coastal communities.

Cytotoxic/anticancer activity: The triterpene glycosides, namely holothurins A3 and
A4, of this species have been found to be strongly cytotoxic to cancer cell lines—human
epidermoid carcinoma (KB) and human hepatocellular carcinoma (Hep-G2), with IC50
values of 0.87 and 0.32 µg/mL (for holothurin A3) and of 1.12 and 0.57 µg/mL (for
holothurin A4), respectively (Li et al., 2013).
Li et al. (2013) reported that the triterpene glycosides, arguside A–E, of this species exhibited
significant cytotoxicity against different human tumour cell lines. Arguside A showed the
highest activity towards human colorectal carcinoma (HCT-116) cells (IC50 = 0.14 µM);
argusides B and C showed potent cytotoxicity against adenocarcinomic human alveolar basal
epithelial cells (A549), HCT-116, HepG2, and human breast adenocarcinoma (MCF-7) cell
lines. The IC50 values of these compounds on A549 were 0.48 and 0.43 µg/mL, respectively,
and for HCT-116, the values were 0.46 and 0.38 µg/mL, respectively. Besides, argusides D
and E have also been tested for their anticancer activities in above human cancer cell lines
and revealed a significant activity with IC50 values in the range of 3.36–7.77 µg/mL.
Li et al. (2013) reported that the glycosides, impatienside A and bivittoside D, isolated from
this species have shown in vitro cytotoxicity against A549 (IC50 = 0.35 µg/mL and 0.52 µg/
mL, respectively); HCT-116 (-IC50 = 0.45 µg/mL and 0.37 µg/mL, respectively); DU-145
(IC50 = 1.14 µg/mL and 0.937 µg/mL, respectively); and KB tumour cells (IC50 = 1.6 µg/
mL and 1.42 µg/mL, respectively).
The compounds, 17-Dehydroxyholothurinoside A and griseaside A, of this species have

been identified as promising anticancer agents due to their significantly higher cytotoxicity
against four human tumour cell lines, A549 (IC50 = 0.886 µM and 1.07 µM); HL-60
(IC50 = 0.245 µM and 0.427 µM); BEL-7402 (IC50 = 0.97 µM and 1.114 µM); and
human acute lymphoblastic leukaemia cell line (Molt-4) (IC50 = 0.34 µM and 0.521 µM)
(Li et al., 2013).
The triterpenoid glycoside, hillaside C, has also been tested for the anticancer potential
of this species against eight human tumour cell lines (A-549, MCF-7, human lung
carcinoma cells-IA9, human clear cell carcinoma cells—CAKI-1, human prostate cancer
cells—PC-3, KB, KB-VIN, and human colorectal adenocarcinoma cells—HCT8) and
has exhibited significant cytotoxicity with IC50 values in the range of 0.15–3.20 µg/mL
(Li et al., 2013).
The ethyl acetate extract of the body wall of this species has shown anticancer activity
against two human glioblastoma (brain tumour) cell lines A172 and U87MG with IC50
values of 4.23 and 4.46 mg/mL, respectively. The above extract has also been reported
to induce both early and late stages of apoptosis, loss of mitochondrial membrane
potentials, and nuclear condensation and fragmentation (Sangpairoj et al., 2016).
Antioxidant activity: The crude methanol extracts and three pure compounds isolated from
this species—viz. Friedelin, 3-Hydroxybenzaldehyde, and 4-Hydroxybenzaldehyde—
have shown DPPH radical scavenging activity, and the effective concentration (EC50)
values were found to be 33.77, 14.63, 14.62, and 14.78 mg/mL for the whole body extract,
Friedelin, 3-Hydroxybenzaldehyde and 4-Hydroxybenzaldehyde, respectively (Nobsathian
et al., 2017).
Antifungal activity: Three triterpene glycosides—viz. scabraside A (1), echinoside A (2)

and holothurin A1 (3) of this species—showed antifungal activities (Han et al., 2009).
The MIC80 values of these compounds on different species of fungi (Candida

albicans, Cryptococcus neoformans, Candida pseudotropicalis, Trichophyton rubrum,
Fonsecaea compacta, Aspergillus fumigatus, and Microsporum gypseum) are given
below:
Antifungal Activities of Triterpene Glycosides (MIC80: µg·mL−1)

Cd C. albicans C. pseudotropicalis C. neoformans T. rubrum M. gypseum F. compacta A. fumigates
1 8 4 8 8 4 8 2
2 4 2 8 4 4 8 1
3 1 6 8 8 8 8 1
Source: Han, H. et al., Acta Pharm. Sin., 44, 620–624, 2009.

activities against Aeromonas hydrophila (inhibition zone, distance from the border
of paper disc to the edge of the clear zone, 6.0 mm) and Enterococcus sp., Klebsiella
pneumoniae, Pseudomonas aeruginosa, Salmonella typhi, Vibrio harveyi, and Aspergillus
sp. (1.25 mm) (Abraham et al., 2002).
Anti-Parkinson potential: The organic extracts of this species (whole body-ethyl acetate
(WBEA), body wall-ethyl acetate (BWEA), viscera-ethyl acetate (VIEA), whole body-
butanol (WBBU), body wall-butanol (BWBU), and viscera-butanol (VIBU) have shown anti-
Parkinson potential at 500 µg/mL by significantly attenuating the DA neuron-degeneration
induced by selective cathecholamine neurotoxin 6-hydroxydopamine (6-OHDA) in the

Caenorhabditis elegans BZ555 strain. Moreover, the extracts also reduced α-synuclein
aggregation and restored lipid content in NL5901, as well as improved food-sensing
behaviour and prolonged life span in the 6-OHDA-treated wild-type strain (Chalorak et al.,
2018).
Others: Thanh et al. (2006) reported on the isolation of a new triterpene glycoside,
holothurinogenin B (1) along with three known compounds, holothurin B (2), holothurin
A (3), and holothurin A2 (4), from the methanol extract of this species. The bioactivities of
these compounds are to be known (Thanh et al., 2006).
Holothuria (Holothuria) tubulosa (Gmelin, 1791)
Common name(s): Cotton-spinner, tubular sea cucumber
Global distribution: Temperate regions of the eastern Atlantic Ocean as far north as the
Bay of Biscay, in the Mediterranean Sea.
Ecology: It is found on sandy seabeds, among seagrass (Posidonia spp.), and on muddy
rocks to a depth of about 100 m.
Biology
Description: It grows to a length of 20–45 cm and a diameter of 6 cm. The body
of this species is roughly cylindrical with a flattened base on which there are three
longitudinal rows of tube feet. It has a tough, leathery skin. The general colour is
a shade of brown and the surface is covered with numerous dark-coloured, conical,
thorn-like projections known as papillae. It often appears greyish, as it secretes a
protective film of mucus to which pieces of seaweed and sediment adhere. It has a
fringe of short, flattened tentacles around the mouth at the anterior end, and the anus
is located at the posterior end.
Food and feeding: It feeds on detritus, algae, and plankton.
Reproduction: It is dioecious, with each individual being either male or female but with
no difference between them in external appearance. First, males emit a whitish fluid that
is gradually dispersed in the surrounding water. Next, females emit a more viscous fluid,
in response to the release of the male gametes. Each release lasted about 30 minutes.
The necessary conditions for spawning seemed to be a full moon and a sea temperature
of about 25°C. The fertilized eggs hatch in about 24 hours, and the developing larvae
are planktonic. They feed on microscopic algae and pass through several stages over the
course of a few weeks before settling on the seabed and undergoing metamorphosis into
juveniles.

Antimicrobial activity: The associated bacterial strains of this species showed antifungal
activity (León-Palmero et al., 2018).
Others: Silchenko et al. (2005) reported on the occurrence of holothurins A and B from
this species.
Holothuria sp.1

Antiviral activity: The cold-water extract of this unidentified species exhibited antiviral
activity not only against the virus adsorption to the cells but also on virus intracellular
replication. The IC50 values for the inhibition of the virus adsorption to the cells and virus
intracellular replication were 120.2 and 189.9 µg/mL, respectively (Farshadpour et al., 2014).
Pearsonothuria graeffei (Semper, 1868)
Common name(s): Blackspotted sea cucumber, flowerfish, Graeffe’s sea cucumber
Global distribution: Tropical parts of the Indo-Pacific Ocean: from the east coast of
Africa to the Philippines, Indonesia, and the South Pacific
Ecology: It is found on the seabed and on coral reefs, and reef slopes close to the coast at
depths down to about 25 m. When threatened or disturbed, it ejects its Cuvierian tubules,
thin white sticky strands of viscera, from its cloaca.
Biology
Description: The body of this species is subcylindrical, arched dorsally (bivium), and slightly
flattened ventrally (trivium). White conical papillae are sparsely distributed on bivium. Podia
on trivium are long and large. Its mouth, at one end, is surrounded by a ring of up to 24 leaf-
like, paddle-shaped tentacles with black stalks that are black on the upper side and white
beneath. The anus is located at the other end of the body, and there are several rows of tube
feet along the underside. Spicules on the dorsal and ventral tegument are with pseudo-tables
and rosettes. The colour of the adults is pale brown and white, with black speckles and small
thorn-like protuberances. It has a maximum length of 45 cm and weight of 1.3 kg.
Food and feeding: It is a scavenger and roams around on the seabed sifting through the sedi-
ment with its feeding tentacles. Any organic matter it finds is passed to its mouth by the tentacles.

Cytotoxic activity: The glycosylated triterpenes compounds, echinoside A and
ds-echinoside A, isolated from this species showed cytotoxic activity against the P388,
A549, MKN-28, HCT-116, and MCF-7 cancer cell lines. Both compounds displayed
potent activity in blocking cell cycle progression and induced apoptosis through the
intrinsic mitochondrial pathway, with ds-echinoside being the most potent one (Ruiz-
Torres et al., 2017).
Anticancer activity: Ds-echinoside A of this species inhibited the proliferation of human
hepatocellular liver carcinoma cells Hep G2, with a half-maximal inhibitory concentration
(IC50) of 2.65 μmol/L, and suppressed Hep G2 cell adhesion, migration, and invasion in a
dose-dependent manner. Ds-echinoside A also reduced tube formation of human endothelial
cells ECV-304 on matrigel in vitro and attenuated neovascularization in the chick embryo
chorioallantoic membrane assay in vivo. Immunocytochemical analysis revealed that
Ds-echinoside A significantly decreased the expression of matrix metalloproteinase-9,
which plays an important role in the degradation of basement membrane in tumour
metastasis and angiogenesis (Zhao et al., 2011).

Antifungal, antiprotozoal, and anticancer activities: Two lanostane-type sulfated
triterpene monoglycosides, holothurin A and echinoside A, isolated from this species
showed activities against Candida albicans, Leishmania major, and colon cell line
(Khattab et al., 2018).
Others: The fucoidans 4-O- and 2, 4-di-O-sulfated polysaccharides have been isolated
from this species (Hu et al., 2015).
5.8 FAMILY: CAUDINIDAE (ORDER: MOLPADIDA)

Acaudina molpadioides (Bordbar et al., 2011)
Common name(s): Sea potato
Global distribution: Tropical Indo-West Pacific: India, Thailand, Indonesia, Philippines,

and Australia
Ecology: This benthic species, which lives in slightly less saline waters, is found in reefs
and in estuarine and muddy areas in depths of 0–330 m.
Biology
Description: The body of a sea potato is stout, spindle shaped, and tapers at the posterior
end, making it a distinct species among other sea cucumbers. An additional feature of
this species is the presence of 15 tentacles on its body. The body wall of the sea potato
is smooth and opaque while juveniles have transparent body walls. Tube feet are absent.
It measures about 15–20 cm in length. Its body is consistently flesh coloured, and the
colouration starts to fade if the animal stays out of water for a long period.
Food and feeding: It feeds on detritus and other decomposing matter, leaving a bead-like
faecal pellet trail behind.

Antitumour activity: The cerebrosides of this species displayed antitumour activity with
an inhibitory effect on cell proliferation through induction of apoptosis in S180 cells.
Administration of these cerebrosides (50 mg/kg BW) on S180 tumour-bearing mice
reduced the tumour weight by 45.2%. In S180 ascites tumour model, cerebroside (50 mg/
kg BW) treatment exhibited a significant ascites fluid growth inhibition of 31.2% (Du
et al., 2012a).
Anti-adipogenic activity: The fucoidan of this species showed anti-adipogenic activity by

modulating the Wnt/β-catenin pathway and downregulating the expression of SREBP-1c
(Xu et al., 2014).
Inhibition of pancreatic islet cell apoptosis and inflammation: The fucoidans of this
species inhibited pancreatic islet cell apoptosis and inflammation in type 2 diabetic mice.
These fucoidans have also been reported to repair pancreatic islet cells, decreased serum
C-reactive protein (CRP), macrophage inflammatory protein 1 (MIP-1), interleukin 1β (IL-
1β), IL-6, and tumour necrosis factor-α (TNF-α) levels, and increased the IL-10 level. Further,
these compounds also reduced TNF-α, CRP, MIP-1, IL-1β, and IL-6 mRNA expressions,
and increased IL-10 mRNA expression in epididymal adipose tissues (Wang et al., 2016).
Paracaudina chilensis (Müller, 1850)
Common name(s): Sand sea cucumber, rattail sea cucumber
Global distribution: Subtropical Eastern and Northwest Pacific: China
Ecology: It is a coastal species, buried 10–20 cm below sandy mud. This benthic species
has a depth range of 0–1000 m.
Biology
Description: The body of this species is firm, thin, leathery to parchment-like, and finely
wrinkled. Mid-body ossicles are predominantly octagonal plates. Spicules are in the form
of perforated plates. It is a white sea cucumber with a purple cast near the posterior end.
This small to medium-sized species has a maximum length of 20 cm and diameter of 2 cm.
Its posterior body (caudal) tapers as a discrete tail with a length of 8 mm.
Functional and bioactive properties: It is reported that the collagen and gelatin-derived peptides
of this species have exhibited numerous bioactivities, namely antioxidant and antihypertensive/
ACE inhibitory activity, antimicrobial activity, lipid-lowering effect, immunomodulatory
activity, and beneficial effects on skin, bone, or joint health (Gómez-Guillén et al., https://pdfs.
semanticscholar.org/f004/71261cc9f93aeda9d5049ec3c7351cd32754.pdf)
Pseudostichopus mollis (Théel, 1886) (= Pseudostichopus trachus)
Common name(s): Sandy sea cucumber
Global distribution: Indo-Pacific and the Antarctic: Kerguelen and Antarctica
Ecology: This polar, benthic species is a widespread deep-sea species and is found in mud
and sand strata; depth range is 100–5203 m.
Biology
Description: It grows up to 15 cm in length. It is shaped like a flattened sausage. The skin is
covered with particles of sand or fine debris. It is beige in colour and smooth. There are fine
tube feet, which are not apparent to the naked eye. At the posterior end, the anus is in a vertical
groove. The mouth is on the underside of the anterior end. There are 20 peltate tentacles that are
fairly small and usually retracted inside the mouth. It roughly measures 15 cm long on average.

The triterpene disulfated tetraosides (holostane oligoglycosides), viz. pseudostichoposides
A and B, have been isolated from this species. Activities of these compounds are to be
known (Kalinin et al., 1989; Silchenko et al., 2004).
5.9 FAMILY: STICHOPODIDAE (ORDER: SYNALLICTIDA)

Apostichopus japonicus (Selenka, 1867) (= Stichopus japonicus)
Common name(s): Japanese spiky sea cucumber, Japanese sea cucumber
Global distribution: Temperate Southeast Asia; Russia, China, Japan and Korea
Ecology: It is a shallow-water species. While the red morphs of this species are found on
gravel beds offshore at 40 m. or deeper depths, the other two colours are found intermingled
on muddy and sandy bottoms at shallower depths.
Biology
Description: It has a cylindrical leathery body with blunt, thorny protuberances. At the
anterior or front end there is a mouth surrounded by a ring of short feeding tentacles, and
at the posterior end is the anus. In this species, there are three different colour morphs, viz.
red, green, and black.
Food and feeding: It sifts through the sediment on the seabed with its tentacles and
feeds on detritus and other organic matter including plant and animal remains, bacteria,
protozoa, diatoms, and faeces.
Reproduction: In this species, the sexes are separate. Males and females release a mass of
gametes into the sea where fertilization takes place. The planktonic larvae develop through
several stages before settling on hard surfaces on the seabed, undergoing metamorphosis
and becoming juveniles.
Edible/commercial values: It is used for food. It is also cultivated on a commercial scale
in shallow ponds and by sea ranching in northern China.

Antifungal activity: Mondol et al. (2017) reported on the isolation of holotoxins A, A1,
B, B1, D, D1, and E–I from this species. Among them, holotoxin A1 exhibited antifungal
activity against five pathogenic fungi with MIC values ranging from 0.5 to 1.0 µg/mL).
The nortriterpene glycoside and triterpene glycosides, viz. 26-nor-25-oxo-holotoxin

A1, holotoxin D, holotoxin E-G, holotoxin A1 and B, and cladoloside B isolated from
the ethanolic extracts demonstrated antifungal properties against six fungal strains such
as Candida albicans, C. tropicalis, Cryptococcus neoformans, Trichophyton rubrum,
Microsporum gypseum, and Aspergillus fumigates (Oh et al., 2017).
Anticancer activity: The EtOAc fractions of this species showed higher anticancer
activity and cell growth inhibition in HL-60 and HT29 cells. Further, these fractions
inhibited the HepG2 cell proliferation and induced apoptosis by downregulation of
Bcl-2 and upregulation of nm23-H1 gene expression in a time and dose-dependent manner.
Furthermore, the extracts of this species inhibited the SK-Mel-2 cell proliferation and
metastasis through reduction in the expression of skin tumour-promoting genes such as
Bcl-2, STAT3, and MMP-9. These reports suggest that S. japonicus could be a natural
source of the potential anticancer drug (Oh et al., 2017).
The hot water extract of this species has shown antiproliferative and cytotoxic activities
against human colon cancer CaCo-2 cells (Janakiram et al., 2015).
Antitumour activity: The mucopolysaccharide obtained from the hot water extract of
this species showed antitumour activity in human colon adenocarcinoma Caco-2 cells
(Bordbar et al., 2011).
Anti-inflammatory and anti-allergic activities: The water fraction of this species possesses
anti-inflammatory and anti-allergic effects by inhibiting NO release without cytotoxicity in LPS-
stimulated Raw 264.7 cells. The levels of IL-6 and TNF-α mRNA reduced following treatment
with this water fraction. In addition, this fraction inhibited antigen-induced degranulation (e.g.,
IC50 value of 658 μg/mL) as well as IL-4 mRNA expression (Song et al., 2013).
Anti-inflammatory activity: The ethyl acetate fraction of this species had a higher
inflammatory activity in a dose-dependent manner. In addition, this fraction inhibited the
ERK and p38 MAPK phosphorylation in LPS-induced macrophage. This fraction could
therefore be used as an effective anti-inflammation agent (Oh et al., 2017).
Anticoagulation activity: Oh et al. (2017) isolated a depolymerized fucosylated chondroi-

tin sulphate isolated from this species, which showed slightly higher anticoagulant activity
than heparin, suggesting that this species could be used a potential anticoagulant drug.
Antithrombotic activity: The depolymerized fragment of glycosaminoglycan of this

species demonstrated antithrombotic and anticoagulant activities. At levels greater than
0.3 mg/kg i.v., it significantly prevented the death of mice treated with thrombin (800 U/kg
i.v.). Under the same conditions, FGAG, UFH, and LMWH prevented the death of mice at
dosage higher than 0.3, 0.3, and 0.6 mg/kg i.v., respectively (Bordbar et al., 2011).
Antihyperlipidemic activity: When rats were fed with the polysaccharides (glucosamine,
galactosamine, glucuronic acid, mannose, glucose, galactose, and fucose) at 400 mg/kg,
the TC, TG, and LDL-C concentrations decreased significantly by 17.23%, 20.78%, and
31.18%, respectively, and HDL-C increased by 27.27%. This species could therefore be a
potential therapeutic target for hyperlipidemia (Oh et al., 2017).
Anti-melanogenesis activity: The ethanolic extracts of this species showed anti-melanogenesis

activity by significantly reducing the melanin content and the expression of melanogenesis-
related proteins (tyrosinase, TRP-1, TRP-2, and MITF) in B16 cells (Oh et al., 2017).
As tyrosinase inhibitors: The adenosine and ethyl-α-d-glucopyranoside of the ethanol

extracts of this species exhibited higher tyrosinase inhibitory activity with IC50
value of 0.191 and 0.126 mg/mL, respectively. Further, its ethanol extracts caused the
reduction expression level of tyrosinase, TRP-1, TRP-2, and MITF by phosphorylation
of ERK. Therefore, this species may be a good source for natural tyrosinase inhibitors
(Oh et al., 2017).
Anti-osteoclastogenesis activity: The types A and B fucan sulphate from the

chloroform/methanol extract of this species inhibited osteoclast-like cell formation by
99.8% and 96.3%, respectively. These results suggest that fucan sulfates A and B of this
species have great potential to be used as a source for natural anti-osteoclastogenesis
reagent (Oh et al., 2017).
Antioxidant and tyrosinase inhibitory activities: A gelatin hydrolysate of this species

showed antioxidant activity by scavenging the superoxide and hydroxyl radicals in a
concentration-dependent manner. Further, it also showed very good inhibitory effect
against melanin synthesis and tyrosinase activity in B16 cells (Bordbar et al., 2011).
Anti-oxidative activity: Two tetrapeptides, viz. Val-Thr-Pro-Tyr and ValLeu-Leu-Tyr; and

a hexapeptide Val-Gly-ThrVal-Glu-Met obtained from the autolysis hydrolysates of the
gut of this species exhibited antioxidant activities by the protection of hydroxyl radical-
induced DNA damage. These oligopeptides exhibited protective barrier against hydroxyl
radical-induced DNA damage at a concentration of 2 mg/mL (Oh et al., 2017).
Immunomodulatory effect: The acid mucopolysaccharide isolated from this species

showed immunomodulatory effect. The treatment groups had significantly reduced nodule
volume and nodule number, serum α-fetoprotein level, proliferating cell nuclear antigen
expression, IL-2 and TNF-α, and increased levels of p21 expression. Further, the protein-
sulfated fucan complex from the body wall of this species significantly induced the mRNA
expression of iNOS and cytokines generation including IL-6, IL-10, TNF-α, and COX-2
(Oh et al., 2017).
Antiprotozoal activity: It has been documented that holotoxins from this species exhibited
activity against Trichomonas vaginalis, and its holothurin showed activity towards Amoeba
proteus (Bahrami, 2015).
Neural-related activities: The neuritogenic active ganglioside isolated from the n-hexane
fraction of chloroform/methanol extracts of this species showed neural-related activities.
The sulfated polysaccharide from this species has been reported to promote the neurosphere
formation by activating nuclear factor NF-κB in NSCs (Oh et al., 2017).
Anti-fatigue effect: The acidic mucopolysaccharides, collagen, bioactive amino

acids, and lipids of the body wall of this species have been reported to possess anti-
fatigue effects. In comparison with the control group, administration of this species for
30 consecutive days prolonged the duration of exhaustive swimming in mice, promoted
the synthesis of liver glycogen and haemoglobin, and kept the level of haemoglobin
(90 minutes post-swimming) similar to that of before swimming. It also significantly
decreased the generation of blood lactic acid and accelerated the elimination of blood
lactic acid and blood urea nitrogen in mice after swimming thus improving the exercise
endurance in mice (Bordbar et al., 2011).
Apostichopus parvimensis (Clark, 1913) (= Stichopus parvimensis)
Common name(s): Warty sea cucumber
Global distribution: Eastern Pacific Ocean; Baja California, Mexico to Monterey Bay,
and California
Ecology: It lives on subtidal habitats, viz. rocky and soft habitats, and kelp forest at depths
between 1 and 60 m.
Biology
Description: The warty sea cucumber, which has a highly contractile and cylindrical
body, is soft as a feather pillow. It can reach a maximum length of 30–40 cm. The
common name “warty” comes from the numerous black-tipped projections covering
the sea cucumber’s brownish skin. There are many small black-tipped papillae on the
dorsal surface. An intestinal tract connects the mouth and anus on opposite ends of
the body. A respiratory tree (a collection of thin-walled tubes) stems from this tract.
The sea cucumber draws water through its anus into the tree where respiration takes
place. It then forcefully expels the water. These warty sea cucumbers and their related
species are sometimes called the “earthworms of the sea,” as they cultivate the seafloor
as earthworms cultivate the soil.
Food and feeding: It is a deposit feeder that feeds on soft sediments, digesting the organic
detritus and small organisms contained within. This animal will lie half buried in the soft
substratum, passing through the intestinal tract quantities of sand and mud from which its
food is extracted. It has mop-like tentacles, which when pressed onto the substratum pick
up particles and transfer them to the mouth. The digestive system processes the organic
matter, and the bits of shell and sand particles pass through the gut.
Reproduction: This species expresses a definite reproductive cycle with spawning in May
and June, possibly in response to increasing water temperatures. The gonad is completely
reabsorbed during September and October, and the gonadal growing phase starts around
January. It uses an external broadcast spawning method, so fertilization is largely a matter
of chance. Environmental cues, such as consecutive sunny days, a plankton bloom, or
a certain temperature, can cause a large number of individuals to spawn simultaneously,
thereby increasing the chances for successful fertilization. After fertilization takes place, a
larva is formed that metamorphoses into a sea cucumber, after only a few weeks.
Predators: Predators of this species include sea stars such as Pycnopodia helianthoides,
fish, gastropods, and crustaceans as well as humans.

Mondol et al. (2017) reported on the isolation of parvimosides A, B from this species.
Astichopus multifidus (Sluiter, 1910) (= Stichopus multifidus)
Common name(s): Furry sea cucumber, fissured sea cucumber
Global distribution: This species is found in the tropical Western Central Atlantic, from
the southeastern US and the Gulf of Mexico, south to Colombia and Venezuela, and east to
Jamaica, Dominican Republic, Puerto Rico, Cuba, Panama, and the Bahamas.
Ecology: This benthic species prefers soft bottoms with muddy or sandy patches, in and
around Thalassia seagrass beds. It is also found in areas of calcareous algae and deeper,
calmer reef environments; the depth range is 1–37 m.
Biology
Description: It is a large sea cucumber, and its back is covered with numerous small, 1 cm
long, pointed podia. Colour of the body is a mottled chocolate brown to grey. Sometimes
white blotches and spots are seen on the body. It attains a maximum length of 50 cm.

Cytotoxic activity: The triterpene oligoglycoside, astichoposide D isolated from this species
showed antiproliferative/cytotoxic activity against 2 HTCL, Vero cells (Blunt et al., 2017).
Australostichopus mollis (Hutton 1872)
Common name(s): Brown sea cucumber, Australasian sea cucumber, soft sea cucumber,
New Zealand sea cucumber
Global distribution: From Cook Strait to Stewart Island and from New South Wales to
south Western Australia
Ecology: It inhabits the littoral at low tide level on sand, as well as in mud in protected
areas. Its vertical range goes from shallows waters on the coast to around 200 m depth.
Biology
Description: It is a medium-sized species of sea cucumber, reaching up to 300 g in wet
weight and up to 25 cm in length. This species has the typical cylindrical shape of sea
cucumbers, with the mouth located anteriorly facing the seafloor and the anus positioned
at the posterior end. The dorsal surface of this species is characterized by a series of
irregularly positioned papillae of different sizes and shapes. The ventral surface, on the
other hand, is smoother, with the presence of ambulacral podia or tube feet, which are used
for locomotion. The colouration of this species varies significantly. Individuals may be
with a single colour (from dark brown to white) or have a combination of different brown,
yellow, and cream-white tones. Like other holothurians, it has the ability to regenerate its
internal organs after undergoing a phenomenon called auto-evisceration. This process,
which is thought to be induced by external stimuli, is characterized by violent and rapid
expulsion of organs, followed by regeneration.
Food and feeding: It is a detritivore feeding on particles of organic matter from the substrata.
Reproduction: In this species, the sexes are generally separate; however, some
hermaphrodites have been found. Reproductive season is from October to February, and
periodicity of spawning is regulated by lunar cycles, specifically following a full moon.
Aquaculture uses: This species is cultivated with the green-lipped mussel, Perna
canaliculus, and the native abalone, Haliotis iris. Further, it is capable of reducing wastes
produced by aquaculture.

Mondol et al. (2017) reported on the isolation of compounds, viz. mollisoside A, B2 and
neothyonidoside.
Isostichopus badionotus (Selenka, 1867)

Common name(s): Chocolate chip cucumber, cookie dough sea cucumber, three-rowed
sea cucumber
Global distribution: Warm parts of the Atlantic Ocean; from North Carolina (US),
through the Caribbean, to north Brazil, at Ascension Island and in the Gulf of Guinea
Ecology: This benthic species lives on the sandy bottoms around reefs, sandy rubble areas,
and in grass beds (Thalassia beds); depth range is 0–65 m.
Biology
Description: This is a large species that can grow to a maximum length of 45 cm. It has
distinctive dark-coloured “warts” in three coarse rows on its dorsal surface. The rest of
the body may vary from white through to shades of orange to brown, with sometimes
large brownish stains. The mouth is located ventrally and surrounded by about 20 large
tentacles. On the upper side are small knob-like podia that are often in contrasting
colour.

Anti-inflammatory activity: Lyophilized extracts of this species exerted a strong
anti-inflammatory activity in vivo. They attenuated histological disruption caused by
inflammatory agents, repressed the expression of pro-inflammatory genes including TNFα
iNOS, COX2, NFκB or IL-6, and slightly enhanced the expression of anti-inflammatory or
survival genes (Olivera-Castillo et al., 2018).
against Plasmodium falciparum FcB1 strain and the recorded IC50 value was 86.7 µg/mL
Anticancer activity: The clarified extracts of this species showed anticancer activity against
3LL and PC3 cancer cell lines with IC50 values of 57.1 and 83.1 µg/mL, respectively
Stichopus chloronotus (Brandt, 1835)
Common name(s): Okinawan sea cucumber, greenfish sea cucumber, spiky sea cucumber
and black knobby sea cucumber
Global distribution: This species is found in the Indo-Pacific region, from the Red Sea
and the East Coast of Africa, through Madagascar, Seychelles, Comores, and Réunion to
Australia, Indonesia, China, Japan, Guam, Fiji, Tonga, and Samoa.
Ecology: It lives on reefs but can also be found on rubble located on the outer reef flats at
depths down to 12 m.
Biology
Description: It is a fairly large species, growing to 25 cm. It has a firm but pliable body and
a squarish cross section. The skin is smooth but there are numerous conical fleshy papillae
in longitudinal rows, and these are larger on the lower lateral angles. This species is with
deep blackish green in colour, and it has yellow or red tips to the papillae.
Food and feeding: It is a detritivore and sifts through the sediment on the seabed with
its tentacles and feeds on detritus and other organic matter including plant and animal
remains, bacteria, protozoa, diatoms, and faeces. In the feeding process, it swallows a lot
of sand and plays an important part in churning up and aerating the seabed.
Reproduction: It can reproduce asexually by undergoing transverse fission, forming two
new individuals that each regenerate the missing parts. It can also reproduce sexually.

(Layson et al., 2014).
Antifungal activity: The lanostane-type triterpene-oligoglycosides, viz. stichlorosides A1,
B1, C1, A2, B2, and C2 of this species showed antifungal activity (Kitagawa et al., 1981).
Antiproliferative activity: The aqueous extract of this species exhibited antiproliferative

activity against C33A cells (IC50 = 10.0 µg/mL) and A549 cell line (Bordbar et al., 2011).
Cytotoxicity: The organic extract from this species showed more cytotoxicity against C33A
cells (IC50 = 6.0 µg/mL), but little action was found against A549 cells (IC50 = 21.0 µg/
mL) (Bordbar et al., 2011).
Antioxidant activity: The aqueous extract (50 mg/mL) of this species scavenged DPPH
free radical (IC50 = 2.1 mg/mL) more effectively. In this regard, the antioxidant activity
was to the extent of 80.6% (Bordbar et al., 2011).
As functional foods: Fatty acids-profile in terms of myristic (C14:0), palmitic (C16:0),
stearic (C18:0), linoleic (C18:2), arachidic (C20:0), eicosapentaenoic (C20:5, EPA), and
docosahexaenoic acid (C22:6, DHA) suggest that this species could be of great use as
functional foods (Bordbar et al., 2011).
Stichopus hermanni (Semper, 1868)
Common name(s): Herman’s sea cucumber, curryfish herrmanni
Global distribution: Tropical Indo-West Pacific
Ecology: This bottom-dwelling sea cucumber is usually found on sandy areas, coral reefs,
and seagrass beds between 0 and 30 m.
Biology
Description: The body of this species is hard, wrinkled, but without large bumps. Colour
of the body is uniform sandy brown or grey-green-brown. Specimens with light mustard-
yellow to orangey brown or brown and olive green have also been collected. With scattered
papillae, these individuals are orange brown. Short tube feet are present on the flat
underside. The mouth is on the underside with 8–16 stout, green feeding tentacles. It has
a maximum length of 55 cm. Unlike other echinoderms, the oral surface of holothurians is
extended aborally along the elongate polar axis, i.e., that they lie on their side. This species
respires using a respiratory tree, which opens out of the cloaca.
Food and feeding: It grazes on organic materials.
Reproduction: In the Great Barrier Reef (Australia), this species has a single reproductive
event in November, and in New Caledonia it occurs in February. In the Pacific, it reproduces
during the warm season, and in the Islamic Republic of Iran it also reproduces during
summer. This species reaches first maturity at 310 mm.

Antifungal and cytotoxic activities: The organic extracts of this species showed signifi-
cant antifungal activity only against Aspergillus niger with MICs ranging from 3 to 15 µg/
mL. The highest activity was found in body wall (MeOH) extract with an inhibition zone
of 38 mm at a concentration of 18 µg/mL. Body wall and Cuvierian organ extracts of this
species also exhibited cytotoxic activities (Sarhadizadeh et al., 2014).
Antibacterial activity: The saponins of the organic extract of this species showed antibac-
terial activity (Salari et al., 2018).
Antileishmanial activity (antiprotozoal activity): The dichloromethane/methanol extracts of this
species showed antileishmanial activity against Leishmania amazonensis (Oliviera et al., 2016).
Stichopus horrens (Selenka, 1867) (= Stichopus variegatus)
Common name(s): Dragonfish, peanutfish, Selenka’s sea cucumber, Durian sea cucumber,
Vietnamese sea cucumber
Global distribution: This species is found in the tropical Pacific Ocean, from Madagascar
to the Red Sea and Egypt, India to Australia, Indonesia, the Philippines and Japan; its
range extends eastward to the Marianas, Hawaii, Samoa and Tonga, French Polynesia, and
to the Galapagos Islands, Coco Island, and Malpelo Island in the Eastern Pacific.
Ecology: This reef-associated, shallow-water species is found in lagoons and seagrass beds
over sand and rubble; the depth range is 0–15 m. This nocturnal species is often found
hidden in reef flats.
Biology
Description: The body of this species is firm, rigid, and flattened ventrally (trivium). Its
body wall easily disintegrates outside seawater. Bivium is covered with irregular warts,
arranged in 10 longitudinal rows. These warts are larger near the mouth. Trivium is delimited
by a characteristic double row of large papillae. Its stout podia are arranged in four rows,
on three radial bands. Mouth is ventral and is surrounded by a half row of papillae and
18 brown, short tentacles. The anus is terminal. Calcareous ring is with deeply indented
radial pieces and triangular interradials. Cuvierian tubules are absent in this species. Bivium
is whitish to grey, with brown irregular dots, and the trivium is lighter. Spicules are with
very characteristic rosettes (X-shaped or elongate), which are numerous in the tegument.
Numerous C-shaped spicules of three sizes are also present. Ventral tegument is with tables
of two sizes. The species has a maximum length of 50 cm and weight of 500 g.

Mondol et al. (2017) and Khotimchenko (2018) reported on the isolation of Stichoposide
A, B, Variegatuside A–F, Stichorrenoside A–D, 3β-O-[β-D-xylopyranosyl-(1→2)-β-D-
xylopyranosyl]-23S-acetoxyholost-7-ene, and 3β-O-[β-D-xylopyranosyl-(1→2)-β-D-
xylopyranosyl]-23S-hydroxyholost-7-ene from this species.
Cytotoxic activity: The triterpene diglycosides, viz. stichorrenosides A–D, and stichoposides
A, B isolated from the methanol extract of this species showed in vitro cytotoxic activity against
five human cancer cell lines, viz. Hep-G2 (hepatoma cancer), KB (epidermoid carcinoma),
LNCaP (prostate cancer), MCF7 (breast cancer), and SK-Mel2 (melanoma) (Cuong et al., 2017).
Antimicrobial activity: The aqueous methanol extract (8 mg/mL concentration) of the body
wall of this species exerted antimicrobial activity against E. coli with an inhibition zone diameter
of 12.3 mm. On the other hand, the methanol extract (2 mg/mL concentration) showed poor
activity against C. albicans with an inhibition zone diameter of 1.2 mm (Shakouri et al., 2017).
Angiotensin-converting enzyme (ACE) inhibitory capacity: The alcalase hydrolysate
extracted from this species exhibited the highest ACE inhibitory activity (IC50 value of 0.4 mg/
mL) followed by flavourzyme hydrolysate (IC50 value of 2.2 mg/mL), trypsin hydrolysate
(IC50 value of 2.3 mg/mL), papain hydrolysate (IC50 value of 2.5 mg/mL), bromelain
hydrolysate (IC50 value of 4.2 mg/mL), and protamex hydrolysate (IC50 value of 6.4 mg/mL).
These findings suggest that the hydrolysates of this species could be used as a potential source
of functional ACE inhibitory peptides for physiological benefits (Forghani et al., 2012).
Antioxidant and cytotoxic properties: Aqueous and organic extracts of this species
showed antioxidant activity by inhibiting 79.6% and 46.7% of β-carotene oxidation by
linoleate free radical. On the other hand, the organic extract of this species exhibited the
highest cytotoxic effects against A549 and TE1 cancer cells with IC50 values of 15.5 and
4.0 μg/mL, respectively (Althunibat et al., 2013).
Anticancer activity: Triterpene diglycosides: stichorrenosides A, B, C, D, stichoposides
A, B, 3β-O-[β-D-xylopyranosyl-(1→2)-β-D-xylopyranosyl]-23S-acetoxyholost-7-ene,
3β-O-[β-D-xylopyranosyl-(1→2)-β-D-xylopyranosyl]-23S-hydroxyholost-7-en; enzyme
hydrolysates, and cerebrosides isolated from this species showed activity against human
colon cancer cells Caco-2, DLD-1 and WiDr with reduction of cell viability and induction
of apoptosis (Khotimchenko, 2018).
Cytotoxic activity: Three triterpene tetraglycosides, namely stichorrenoside E (1), thelenotoside
B (2), and deacetyl thelenotoside B (3), have been isolated from the MeOH extract of this
species. Among these compounds, thelenotoside B (2) showed strong cytotoxicities against
five human cancer cell lines, viz. HepG2, KB, LNCaP, MCF7, and SK-Mel2 with the IC50
values from 0.95 to 1.90 μM, whereas stichorrenoside E (1) and deacetyl thelenotoside B (3)
exhibited significant activities with IC50 values from 6.87 to 11.62 μM (Vien et al., 2018b).
Thelenota ananas (Jaeger, 1833)
Common name(s): Pineapple sea cucumber, tripang, prickly skin cucumber, pointed teat
sea cucumber, armoured sea cucumber, giant sea cucumber, sand fish, prickly redfish
Global distribution: This species is found on tropical Indo-Pacific waters, from the Red
Sea and East Africa to Hawaii and Polynesia.
Ecology: It is a common reef species mostly found on hard bottoms, large rubble, and coral
patches; it is also found on reef slopes and near passes. Depth range is 0–30 m.
Biology
Description: The body of this species is firm, rigid, and flattened ventrally (trivium).
Bivium is entirely covered with characteristic, large, leaf-shaped, lobate papillae.
Trivium is with brown to pink podia, which are more numerous on the radii. The mouth
is ventral and surrounded by a circle of conical papillae, which are larger on the dorsal
side, and 20 large, brown tentacles. The anus is terminal. Calcareous ring is with large
radial pieces and narrow interradials. Cuvierian tubules are absent. Colour of the body
is variable on bivium where it is reddish orange to brown. Trivium is generally red.
Dorsal tegument has larger branched picules. Ventral podia and tentacles are also with
branched spicules. The individuals of this species attain a maximum length of 80 cm
and weight of 7.0 kg.
Reproduction: Sexual reproduction takes place during the warm season. It is a species with a
low potential fecundity and late sexual maturity. Spawning occurs from January to March.
Parasite: The worm pearlfish (Encheliophis vermicularis) is a parasite of this species and each
parasitized individual will host a male and female pair of the fish, which live inside its body.
Edible values: This species is edible and considered a delicacy in the Far East countries.

Antineoplastic activity: The compound, ananaside C, isolated from this species showed
in vitro antitumour activity against A-549 lung cancer and IA-9 ovarian cancer and eight
other kinds of tumour cell strains. This compound can provide a lead compound for the
development of new antitumour drugs, which has important values for the development
and the utilization of the marine medicinal and biological resources of China (Anon,
https://patents.google.com/patent/CN101245092A/en).
Antiviral activity: The saponin, bivittoside D, isolated from this species showed antiviral
activity against HIV-1 by inhibiting chemokine receptor subtype 5 (Bahrami, 2015).
As functional food and nutraceuticals: The compound, desulfated holothurin A (desHA),

isolated from this species showed highest activity in decreasing low-density lipoprotein-
cholesterol (LDL-C). This suggests that this species could be a valuable resource to exploit
new functional food and nutraceuticals (Han et al., 2018).
Others: Wu et al. (2005) reported on the isolation of a new triterpene glycoside, named as
ananaside D. Mondol et al. (2017) reported on the presence of two new compounds, viz.
thelenotosides A, B.
Thelenota anax (Clark, 1921)
Common name(s): Amberfish, royal sea cucumber
Global distribution: Tropical Indo-West Pacific: from eastern Africa and Madagascar and
the Comoros Islands, east to Australia, Indonesia, the Philippines and Spratly Islands, as
far north as Japan, and southeast to French Polynesia and the Cook Islands
Ecology: This reef-associated species generally occurs on hard ground, large rub-
ble, coral sand patches, on reef slopes, outer lagoons, and near passes; depth range is
10–30 m.
Biology
Description: The body of this species is firm, rigid, and flattened ventrally (trivium).
Bivium is entirely covered with numerous characteristic, conical papillae and minute
podia; it is demarcated from the trivium by a row of large papillae. Podia are numerous
on trivium. The mouth is ventral and surrounded by a circle of 18 large brown tentacles.
The anus is terminal. Calcareous ring is with large radial pieces and narrow interradials.
Cuvierian tubules are absent. Cloaca is large. While the bivium is cream coloured,
with large beige dots, the trivium is generally beige. Branched spicules are seen on the
tegument. Spicules in the form of a “rose window” are mostly abundant in the tentacles,
which are also with straight, curved, or X-shaped rods. Ventral podia are with short,
smooth rods; and dorsal papillae are with sparse, very long, spiny rods. This species has
a maximum length of 1 m and weight of 6 kg.

Mondol et al. (2017) reported on the isolation of glycosides, stichoposides C and D, from
this species.
Antitumour activity: The compound, stichoposide C (STC), isolated from this species
markedly reduced tumour growth of HL-60 xenograft and CT-26 subcutaneous tumours
and increased ceramide generation in vivo (Yun et al., 2012).
Antileukaemic activity: Stichoposide D of this species (STD) induced apoptosis in human

leukaemic cells in a dose- and time-dependent manner (Trincone et al., 2007).
5.10 FAMILY: SYNALLACTIDAE (ORDER: SYNALLICTIDA)

Synallactes nozawai (Mitsukuri, 1912)
Global distribution: It is distributed in the Northwest Pacific from northern Japan to the
Bering Sea.
Ecology: It inhabits the deep-sea areas at 540 m depth.

Biology
Description: The distinct morphological characteristics of this species are as follows:
body flexible, with thin gelatinous body wall; presence of numerous tubercles along dorsal
ambulacra table of body wall consisted of a three- or four-armed disc and a spire-form
pillar.

Silchenko et al. (2002) reported on the occurrence of non-sulfated triterpene glycosides,
synallactosides A1, A2, B1, B2, and C, from this species.
5.11 MISCELLANEOUS
Hemoiedema spectabilis
Common name(s): Patagonian sea cucumber, Malaysian sea cucumber
No other information is available

Antifungal activities: Sulfated triterpene glycosides, hemoiedemosides A and B, isolated
from this species exhibited considerable antifungal activity against Cladosporium cuc
umerinum (Chludil et al., 2002b; Mondol et al., 2017).
Pharmaceutical Sea Lilies or
Feather Stars (Class: Crinoidea)
6.1 FAMILY: ANTEDONIDAE (CLASS CRINOIDEA; ORDER: COMATULIDA)

Heliometra glacialis (Owen, 1833 ex Leach MS) (= Heliometra glacialis maxima)
Common name(s): It has no common names.
Global distribution: Northeast Atlantic and the Arctic: Svalbard and the Canadian Arctic
Archipelago
Ecology: This is a polar, demersal species. It is a free-moving species of sheltered habitats

after the pentacrinoid has detached itself from its stalk; depth range is 18–1358 m.
Biology
Description: The body of this species is centrodorsal hemispherical, with cirrus sockets
irregularly arranged. Cirri, which are rather stout and numerous (about 100 or more), cover
nearly the whole of the centrodorsal. They have mostly ca. 40–60 joints. Brachials are usu-
ally smooth. Proximal segments are somewhat longer than wide, and the distal segment is
short, with more or less prominent dorsal spine. Oral pinnules are greatly elongated and
flagellate, and they are composed of very numerous short and wide joints. The distal part
of these pinnules is more or less serrate. Pinnule 2 is of about the same length as pinnule 1.
The genital pinnules are with fairly large calcareous plates in the side walls. Colour in life
is yellowish. It is a large and stout species, reaching an arm length of more than 200 mm.
373

Shubina et al. (1998) reported on the isolation of seven polyhydroxylated steroids, viz. 24-nor-
cholesta-7,22E-diene-3b,5a,6b-triol, (24j)-methyl-27-nor-cholesta-7,22E-diene-3b,5a,6btriol,
cholesta-7,22E-diene-3b,5a,6b-triol, (24S)-methyl-cholesta-7,22E-diene-3b,5a,6b-triol, (24S)-
ethyl-cholesta-7,22E-diene-3b,5a,6b-triol, (24R)-ethyl-cholesta-7,22E-diene-3b,5a,6b-triol,
and (24R)-ethyl-cholest-7-ene-3b,5a,6b-triol from this species.
Biology and Ecology of Pharmaceutical Sea Lilies or Feather Stars (Class: Crinoidea) 375
6.2 FAMILY: COMATULIDAE (ORDER: COMATULIDA)

Anneissia bennetti (Müller, 1841) (= Oxycomanthus bennetti; Comanthus bennetti)
Common name(s): Bennett’s feather star, Bennett’s rainbow crinoid
Global distribution: This species is found in the Indo-West Pacific, from the Bay of
Bengal and Maldives to Marshall Islands, and from China to Australia, Bali, and Indonesia
Ecology: It inhabits exposed coral heads with strong currents, at a depth of 0–50 m.
Biology
Description: This species grows up to 300 mm. It has 31–120 feathery arms, which are
usually held up into the water in order to trap the food. The mouth is on the upper side of
the small disc-like body, among the arms. The arms have numerous finger-like append-
ages known as pinnules. The colour of this species is quite variable, ranging from yellow
to brown and purple. At the beginning of their life, these feather stars stay attached to the
seabed by a stalk, while mature specimen break the stalk and become free-living.
Food and feeding: It feeds on detritus, phytoplankton, and zooplankton.

Anti-inflammatory and analgesic effects: The crude extract of this species significantly
inhibited the expression of pro-inflammatory proteins in lipopolysaccharide (LPS)-
stimulated murine RAW 264.7 macrophage cells. Further, its compound, comaparvin, sig-
nificantly decreased the expression of inducible nitric oxide synthase (iNOS) protein and
mRNA in LPS-stimulated macrophage cells. Moreover, the post-treatment with comaparvin
significantly inhibited mechanical allodynia, thermal hyperalgesia, and weight-bearing
deficits in rats with carrageenan-induced inflammation. Comaparvin also attenuated leuko-
cyte infiltration and iNOS protein expression in carrageenan-induced inflamed paws. These
findings suggest that comaparvin of this species is a potential anti-inflammatory therapeu-
tic agent against inflammatory pain (Chen et al., 2014).
Comatula (Validia) rotalaria (Lamarck, 1816)
No information is available on the biology and ecology of this species.

Khokhar et al. (2016) reported on the isolation of anthraquinones, rhodocomatu-
lin 5,7-dimethyl ether, rhodocomatulin 7-methyl ether, 12-desethylrhodocomatulin
5,7-dimethyl ether, and 12-desethylrhodocomatulin 7-methyl ether from this species.
Francesconi (1980) reported on the presence of rhodocomatulin 6-methyl ether from

this species.
6.3 FAMILY: HIMEROMETRIDAE (ORDER: COMATULIDA)

Himerometra robustipinna (Carpenter, 1881) (= Himerometra magnipinna)
Global distribution: This species is found in the Western Pacific and in the Indian Ocean,
from the Bay of Bengal up to the China Sea, Great Barrier Reef, Indonesia, Philippines,
and South Japan.
Ecology: It lives in the coastal waters with a coral reef ecosystem, relatively strong cur-
rents, clear water, and plankton availability, at a depth of 0–57 m. It can cling onto corals
with short appendages called cirri, but it also can freely swim.
Biology
Description: This species can reach a diameter of 35–40 cm. It has a cup-shaped body
with 33–62 arms (usually 45). These arms, which are of about 20 cm in length, extend out
from the central disc. The mouth and anus are both on the upper side of the body. Usually
this species has reddish or maroon arms, but it may have yellow or pale brown brachials
with maroon pinnules.
Food and feeding: It feeds on detritus, phytoplankton, and zooplankton caught by means
of a sticky substance on the arms.
Reproduction: The larvae of this feather star swim freely with plankton for a few weeks
before they settle down and grow into a stalked form. Mature specimens, however, break
the stalk and become free-living.
Compounds and Activities

Antibacterial activity: A new tetrabromospirocyclohexadienylisoxazole, (+)-12-hydroxy-
homoaerothionin, together with the known compounds (+)-aerothionin and crinemodin-
rhodoptilometrin bianthrone isolated from this species showed i nhibitory activity with the
hyphae formation inhibition assay in Streptomyces 85E (Shao et al., 2007).
6.4 FAMILY: MARIAMETRIDAE (ORDER: COMATULIDA)

Lamprometra palmata (Müller, 1841) (= Lamprometra klunzingeri)
Common name(s): Feather star, feather Zebra
Global distribution: India
Ecology: Shallow, coastal waters
Biology
Description: It is a brilliantly coloured species occurring in numerous colour varieties.
Individuals are black, brown, or yellow coloured with intercalating white bands forming
a concentric banding pattern. Arms are pinnate, soft, and 26–40 in number. The largest
proximal pinnule of this species is stout and stiffened basally. The enlarged proximal pin-
nules are tapering to a flexible point. The distal pinnules are also flexible. Brachials are
usually wider than long on the proximal half of the arm. Calyx is with flexible tegmen
bearing a relatively short anal tube. Centrodorsal plate is discoidal. The number of cirri is
25–32 with less than 40 segments. Distal cirral segment is with a median crust.

Antibacterial activity: The organic extracts of this species exhibited trace antibacterial
activity against Bacillus subtilis, Staphylococcus aureus, and E. coli. (Anon., http://shod-
hganga.inflibnet.ac.in/bitstream/10603/136970/8/08_chapter%203.pdf).
Antiangiogenic and cytotoxic activities: The chloroform fraction (CC) isolated from the
methanol extract of this species significantly and dose-dependently inhibits proliferation,
migration of endothelial cells and exhibits antiangiogenic effect with ID50 10 ng/10 mL. It
also showed moderate cytotoxicity with IC50 192 mg/mL (Pandit et al., 2009).
Cytotoxic activity: The carotenoids diadinochrome, alloxanthin (pectenoxanthin), cyn-
thiaxanthin, and asterinic acid isolated from the calyx and arms of this species displayed
cytotoxicity (Dembitsky, 2006).
6.5 FAMILY: HEMICRINIDAE (ORDER: CYRTOCNIDA)

Neogymnocrinus richeri (Bourseau et al., 1987) (= Gymnocrinus richeri)
Common name(s): No common name has yet been provided.
Global distribution: Tropical Western Central Pacific: New Caledonia
Ecology: This benthic species appears to be quite numerous on seamounts at depths

between 300 and 500 m.
Biology
Description: It is a weird stalked crinoid. This conic columnal species is with expanded
attachment base. Its calyx is elongated and subcylindrical, with completely fused radials.

Antiviral activity: The compounds gymnochromes A–D, isogymnochrome D, cholest-4-
en-3-one and cholesta-1,4-dien-3-one have been isolated from this species. Of these com-
pounds, gymnochrome, B and D and isogymnochrome D showed antiviral activity against
dengue virus (Riccardis et al., 1991; Motuhi et al., 2016).
6.6 FAMILY: HOLOPODIDAE (ORDER: CYRTOCNIDA)

Holopus rangii (Orbigny, 1837)
Global distribution: It occurs on steep Caribbean slopes, but significant populations have
recently become accessible via submersible off Roatan Island, Honduras.
Ecology: It is cemented to rock walls or even to roofs of submarine caves at depths of 300–400 m.
Biology
Description: This living fossil is one of only eight living species of Cyrtocrinida, an aber-
rant, enigmatic group that first diversified during the Jurassic period and became perhaps
the most diverse crinoid group during the remaining Mesozoic period. It has a conical
to tubular dorsal cup and 10 tapered arms that are able to enroll swiftly into a “fist-like”
configuration when disturbed. Its dorsal cup resembles that of acorn barnacles and other
tubular encrusters. During arm enrollment, the adjacent arms abut and form an impervi-
ous seal. The presence of an anus and absence of the chambered organ and glandular axial
organ (peculiar to crinoids) are also the characteristic features of this species. It is also sug-
gested that the species of this genus may have evolved from stalked cyrtocrinid ancestors
by saltatory loss of major body parts.
Food and feeding: It may feed raptorially by rapidly closing its arms over demersal
zooplankton.
Reproduction: The species is capable of regenerating its damaged arms.
Antiproliferative and antibacterial activities: Compounds such as gymnochrome E,

gymnochrome F, Isogymnochrome B, emodic acid, and a bromo derivative have been iso-
lated from this species. Among them, gymnochrome E (1) of this species inhibited the pro-
liferation of the NCI/ADR-Res (multi-drug-resistant ovarian cancer cell line) with an IC50
value of 3.5 µM and did not show significant inhibitory activity at a concentration of 6.4 µM
against the PANC-1 pancreatic carcinoma or DLD-1 human colorectal adenocarcinoma
cell lines. Gymnochrome E also inhibited histone deacetylase-1 (HDAC-1) with an IC50 of
10.9 µM. Gymnochrome F (2) did not show significant i nhibitory activity at a concentration

of 5.2 µM against the PANC-1, NCI/ADR-Res or DLD-1 tumour cell lines but was a mod-
erate inhibitor of MCL-1 binding to Bak with an IC50 of 3.3 µM. Further, gymnochrome
E (1) also showed antibacterial activity and exhibited minimum inhibition concentrations
(MICs) of 25 µg/mL against Staphylococcus aureus while gymnochrome F (2) exhibited
MICs of 12.5 µg/mL against S. aureus (Wangun et al., 2010).
6.7 FAMILY: PROISOCRINIDAE (ORDER: ISOCRINIDA)

Proisocrinus ruberrimus (A. H. Clark, 1910)
Common name(s): Sea lily or stone lily
Global distribution: Tropical Western Central Pacific: New Caledonia, Japan
Ecology: This benthic species lives at depths of >1000 m.
Biology
Description: The defining feature of all sea lilies is their stalks, which extend their head
(calyx) and arms above the seabed. Their tall stature allows them to filter-feed in the higher
currents found just above the seabed. The species has a strikingly scarlet-coloured stalked
crinoid.

Wolkenstein et al. (2009) reported on the isolation of brominated anthraquinone pigments,
proisocrinins A−F, from this species. The bioactivities of these compounds are however to
be known.
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Compounds
(1S,3S)1methyl 1,2,3,4tetrahydroβcarboline3carboxylic acid
(20R)-24-methyl-5α-cholest-24(28)-ene-3α,21-diyl disulfate
(20R)-24-methylcholest-5,24(28)-diene-3α,21-diyl disulfate
(20R)-3α,4β-dihydroxycholest-5-ene-21-yl sulfate
(20R)-3α-hydroxycholest-5-ene-2β,21-diyl disulfate
(20R)-4β-hydroxycholest-5-ene-3α,21-diyl disulfate
(20R)-5acholestane-3a,21-diol disulfate
(20R)-5α-cholestan-3β,6α,8,15α,24,26-hexaol
(20R)-5α-chlostane-3α,21-didyl disulfate
(20R)-5α-cholest-24-ene-2β,3α,21-triol 3,21-disulfate
(20R)-5α-cholestane-2β,3α,21-triol 3,21-disulfate
(20R)-5α-cholestane-3α,21-diol 3,21-disulfate
(20R)-5α-cholestane-3α,21-diyl disulfate,
(20R)-5α-cholestane-3α,4β,21-triol 3,21-disulfate
(20R)-cholest-5-ene3a,4b,21-triol 3-sulfate
(20R)-cholest-5-ene-3α,21-diol 3,21-disulfate
(20R)-cholest-5-ene-3α,21-diyl disulfate
(20R)-cholest-5-ene-3α,4β,21-triol 3, 21-disulfate
(20R)-cholesta-5,24-diene-2α,3α,4β,21-tetrol 3,21-disulfate
(20R)-cholesta-5,24-diene-2β,3α,21-triol 2,21-disulfate
(20R,25S)-5α-cholestan-3β,6α,15β,16β,26-pentaol
(20R,25S)-5α-cholestan-3β6α,8,15β,16β,26-hexaol
(22E,24R,25R)-24-methyl-5α-cholest-22-en-3β,5,6β,15α,25,26-hexol 26-O-sulfate
(22E,24ξ)-26,27-bisnor-24-methyl-5α-cholest-22-en-3β,5,6β,15α,25-pentol 25-O-sulfate
(24E)-5α-cholest-24-ene-26-yde-3β,6α,8,14,15α-pentaol 15-sulfate
(24j)-methyl-27-nor-cholesta-7,22E-diene-3b,5a,6btriol‑
(24R)-ethyl-cholest-7-ene-3b,5a,6b-triol
(24R)-ethyl-cholesta-7,22E-diene-3b,5a,6b-triol
(24S)-5α-cholestane-3β,4β,5,6α,7β,8,14,15α,24-nonaol 6-sulfate
(24S)-5α-cholestane-3β,6α,8,15β,24-pentaol 24-sulfate
(24S)-ethyl-cholesta-7,22E-diene-3b,5a,6b-triol
(24S)-methyl-cholesta-7,22E-diene-3b,5a,6b-triol
(24S)-O-(β-d-giucopyranosyll-5α-cholestane-3β,6α,8,15β,24-pentaol 6′-sulfate
24(S)-5α-cholestane-3β,6α,8,15β,24-pentol
(25S)-5α-cholestane-3β,4β,6α,7α,8,15α,16β,26-octaol
(25S)-5α-Cholestane-3β,4β,6α,8,15β,16β,26-heptol
(25S)-5α-cholestane-3β,4β,6β, 7α,8,15α,16β,26-octol
(25S)-5α-cholestane-3β,5,6β,15α,16β,26-hexaol (4), and Δ7-sitosterol
(25S)-5α-cholestane-3β,6α,7α,8,15α,16β,26-heptaol
(25S)-5α-Cholestane-3β,6α,8,15β,16β,26-hexol
(28R)-24-ethyl-5αcholesta-3β,5,6β,8,15α,28,29-heptaol-24-sulfate
(2S,3R,4E,8E,10E)-1-(β-D-glucopyranosyloxy)-3 -hydroxy-2-[(R)-2-hydroxyheptadecanoyl)amino]-9-methyl-4,8,10-o
ctadecatriene
(2S,3R,4E,8E,10E)-1-(β-D-glucopyranosyloxy)-3 -hydroxy-2-[(R)-2-hydroxyoctadecanoyl)amino]-9-methyl-4,8,10-oc
tadecatriene
(E) 25-O-β-d-xylopyranosyl-26, 27-dinor-24(S)-methyl-22-ene-15α-O-sulfated-5α-cholest-3β,6α-ol
1-(β-d-ribofuranosyl)-1,2,4- triazole
10-acetoxy-18-hydroxy-2,7-dolabelladiene
12-desethylrhodocomatulin 5,7-dimethyl ether
12-desethylrhodocomatulin 7-methyl ether
12-hydroxyhomoaerothioni
14–methyl–methyl ester
15-O-sulfate of echinasteroside C
16-keto-holothurinogenin
17α-hydroxy impatienside A
17-dehydroxy holothurinoside A
17-Hydroxy fuscocineroside B
17α-hydroxy impatienside A
405
406 Compounds
1-methyl-1,2,3,4-tetrahydro-β-carboline-3-carboxylic acid
1-methyl-6,7-dihydroxy-1,2,3,4-tetrahydroisoquinoline
1-methyl-6,7-dihydroxy-1,2,3,4-tetrahydroisoquinolinium salts of 3-O-sulfoasterone
1′-O-palmitoyl-3′-O-(6-sulfo-α-d-quinovopyranosyl) glycerol
2, 3-Dihydroxy propyl elaidate
2,10-dichloro-3-chlorochamigran-7-en-9-ol
22 ξ-acetoxy-echinoside A
22 ξ-hydroxy-24-dehydro-echinoside A
22,23-dihydro, laeviuscoloside I
22(23)-dihydroechinasteroside A desulfated
22-acetoxy-echinoside A
24(S)-hydroxy-25-dehydro-echinoside A
24–dehydroechinoside A
24-nor-cholesta-7,22E-diene-3b,5a,6b-triol
24-O-β-d-xylopyranoside 4ʺ-sulfate
25-acetoxy bivittoside D
25-hydroxy fuscocinerosiden B
25-hydroxy-dehydroechinoside A
26-nor echinasteroside A desulfated
26-nor-25-oxo-holotoxin A1
26-norechi-nasteroside A
2E,4Z-tanzawaic acid D
2-methyl-8-hydroxy-2H-pyrano [3,2-g] naphthazarin
2-Sulfated β-Galactan
2-sulfated, 3-linked α-l-galactan
3-{O-β-d-fucopyranosyl-(1→3)-β-d-fucopyranosyl-(1→4)-[β-d-quinovopyranosyl-(1→2)]-β-d-quinovopyranosyl}-2-
acetyl-pyrrole
3β-O-sulfated-cholest-5-ene-7α-ol
3-Hydroxybenzaldehyde
3-O-sulfo-24,25-dihydromarthasterone
3-O-sulfoasterone
3-O-sulfomarthasterone
3-O-sulfoisoasterone
3-O-sulfothornasterol A
3′-sulphonoquinovosyl-1′,2′-diacylglyceride
3′-sulphonoquinovosyl-1′-monoacylglyceride
3β,6α-dihydroxy-5α-cholest-9(11)-en-23-one 3-sulfate, were isolated
3β,6α-dihydroxy-5α-cholesta-9(11),24-dien-23-one 3-sulfate
4ʺ-O-methylhalityloside A 6-O-sulfate
4-Hydroxybenzaldehyde
4-keto-cynthiaxanthin
4-O- and 2, 4-di-O-sulfated polysaccharides
5 α-cholest-7-en-3β-yl-sulfate
5,8-Epidioxycholest-6-en-3-ol
5-deoxyisonodososide
5α,8α-epidioxycholest-6-en-3β-ol
5α-cholest-3β,6α,8,14,15α,24,25,26-octaol 15-sulfate
5α-cholest-7,9(11)-diene-3β-ol
5α-cholest-7-ene-3β,6α-diol (5)
5α-cholest-8(14)-ene-3β,7α-diol
5α-cholestane-3β,5,6β,15α,16β,26-hexaol 3-sulfate
5α-cholestane-3β,5,6β,15α,26-pentaol 15-sulfate
5α-cholestane-3β,6α,8,15β,16β,26-hexaol
5α-cholestane-3β,6α,8,15β,24-pentaol 24-sulfate
6,8,9-Trihydroxy-2-methyl-2H-naphtho[2,3-b]pyran-5,10-dione
6,9-dihydroxy-2-methyl-2H-naphtho[2,3-b]pyran-5,10-dione
6,9-dihydroxy-2-methyl-7,8-dichloro-2H-naphtho[2,3-b]pyran-5,10-dione
9-Octadecenoic acid
A
acanthacerebroside A
acanthaglycoside B–D; F
achlioniceosides A1–A3
Compounds 407
adenosine
aerothionin
alkanesulfonic acid
alloxanthin (pectenoxanthin)
α-tocopherol
amurensoside A
amuresterol
ananaside C, D
anasterosides A, B
anthenosides A–X; A1, A2; S1–S6
anthraquinone
aphelaketotriol
aphelasteroside C, D, F
arachidonic acid
archasterosides A, B
argusides A–F
astaxanthin
asterinic acid (7,8-Didehydroastaxanthin)
asterogenins
asterolloside
asterosaponin P1, P2
asterosterol
astichoposide D
astrocerebroside A–C
astropectenols A–D
astrosteriosides A–D
attenuatoside B–1
aurantiamide acetate
axilogoside A
B
bivittosides A–D
borealoside C
brasiliensoside
bromo derivative
butenolide
C
calcigerosides B, C1, C2, D2, E
cariniferosides A–F
carolisterols A–C
celeromycalin
cercodemasoides A–E
cerebrosides, CE‐1‐1, CE‐1‐2, CE‐1‐3, CE‐3‐1, CE‐3‐2
certonardosides A–N; B2,B3; H2–H4; I2,I3,J2,J3,O1,P1,P2
certonardosterol A; A2–A4; B2–B4; C2, D2–D5; E2
cheliferoside L1
cholest-4-en-3-one
cholest-5-ene-2α,3α,4β, 21-tetraol 3,21-disulphate
cholest-5-ene-2β,3α,21-triol 2,21-disulphate
cholest-5-ene-2β,3α,21-triol tri(sodium sulphate)
cholest-5-ene-3α,4β,21-triol 3,21-disulfate
cholesta-1,4-dien-3-one
cholesta-7,22E-diene-3b,5a,6b-triol
cholesterol
cholesterol sulfate
chondroitin
chondroitin sulfates EF1, EF2
cis 11-eicosenoic acid
cis 11,14 eicosadienoic acids
cladoloside B
cladolosides Al–A6; B1, B2, C; C1–C4; D, D1, D2, E1, E2, F1, F2, G, H1, H2, M, M1, M2, N, Q
408 Compounds
colochirosides A, A1, A2, A3, B2, D, E

comaparvin
contractin A
coscinasterosides A–F
cosmasterosides A–D
coustesides A–J
crambescidin 800
crinemodin-rhodoptilometrin bianthrone
cucumarioside A1–A15; A0-1–A0-3; A1-2; A2-2–A2-4; A2-5, A3-2, A3-3, A4-2, A7-1, A7-3; B1,B2,C1,C2,F1,F2,G;
G1–G4; H; H2–H8; I1–I3; I, II
cucumechinosides A–F
culcinosides A–D
culcitoside C2, C6
curacin E
cynthiaxanthin
D
deacetyl thelenotoside B
desholothurin A (nobiliside 2a); A1 (arguside E)
desulfated holothurin A
d-galactose-binding lectins
diadinochrome
diatoxanthin
dictyol E
dictyolactone
dihydroxycrenulide
dimethyl sulfoxide
diplasteriosides A, B
distolasterosides D1–D7
disulfated polyhydroxysteroids
disulfated sterols
docosahexaenoic acid
downeyosides A–L
ds-patagonicoside A
ds-echinoside A
DS-penaustrosides A, B
DS-pervicosides A–C
E
echinamines A, B
echinasterosides A–C
echinochrome A
echinosides A, B
EeCentrocin 1
EeStrongylocin 2
eicosapentaenoic acid
emodic acid
epidioxysterol
ergosta-7,22-dien-3-ol
ethyl-α-D-glucopyranoside
evasteriosides A–E
eximisoside A
F
fallaxosides B1, C1,C2; D1–D7
fellutanine A
fisherioside A
forbesides A–D; E; E1–E3; F–L
forbesin
friedelin
Compounds 409
fromiamycalin
frondosides A, A1; A2-1–A2-4; A2-7, A2-8; A7-1, A7-2; B
fucan sulphate A, B
fucose
fucosylated chondroitin sulphate
fuscocineroside A–C
G
galactosamine
galactose
gangliosides LG-1, LG-2
gelatin hydrolysate
gliotoxin
glucosamine
glucose
glucuronic acid
glycerol 1,3-dioleate-2-stearate
glycerol 1-palmitate
glycine
glycoproteins
granulatoside A
griseaside A
gymnochromes A–F
H
halicerebroside A
halistanol trisulfate C
halityloside A 6-O-sulfate
halityloside H,6-O-sulfate
halitylosides A, B; D–F; H, I
hedathiosulfonic acids A, B
hemoiedemosides A, B
hesperusides A–C
hillasides B, C
hippasteriosides A–D
holothurins A, B, A1–A4; B, B2–B4; C, D
holothurinosides A–C; A1, B, C1, D, E1, H, H1, I, I1, J, J1, K1, L, M, O, Q, R, R1, S, T; X–Z
holotoxins A, A1, B, B1, D, D1; E–G
homarine
HSO3-8NeuGc α 2-6Glc β 1-1Cer (T1 ganglioside)
HSO3-8NeuGc(α)2-6Glc(β)1-1ceramide
hylodoside A
hypoxanthine
imbricatine
impatiensides A, B
indicosides A–C
intercedensides A–I
isogymnochrome B, D
isokoreoside A
isonodososide
J
johnstonol
K
kolgaosides A, B
kurilensosides A, B; D–J
kurilosides A, C
410 Compounds
L
labidiasteroside A
laeviusculosides C; G–I
laurinterol
lectin CEL-III
lectin-III
lefevreiosides A1, A2; B–D
leptaochotensosides A–C
leptasteriosides A–F
lessoniosides A–E
lethasteriosides A, B
leucospilotasides A, B
leviusculoside G (forbeside J); J
linckosides A, B; F–K; LI, L3
liouvillosides A, B; A1–A5; B1, B2
longicaudosides A, B
lucunterperacetate
luidiaquinoside
luridoside A
lutein
luzonicosides A–F
lysaketodiol
lysaketotriol
lysastroside A
M
maculalactones A, E
magnumosides A1–A4; B1–B4; C1–C4
mannose
marmoratosides A, B
marthasterone sulfate
marthasterosides A1, A2, B, C
metallothionein
minutosides A, B
mirabiquinone A
mithrotriol
mollisoside A, B2
monosulfated sterol
myxodermoside A
N
N-(2-(1H-indol-3-yl)ethyl)-2-phenylethanamine
N-Acyl taurine
neothyonidoside
NeuGc(α)2-6Glc(β)1-1ceramide (M5 ganglioside)
nipoglycosides A–D
nobilisides B, D
novaeguinosides A, Y, I, II
O
okhotosides B1–B3
ophidiacerebrosides B–D; E
ophidianoside F
ophiodilactones A, B
ophiurasaponin
ophydianoside F
ovarian asterosaponin 1
ovothiol A (N1-methyl-4-mercaptohistidine)
Compounds 411
P
pachydictyol A
pacifenol
palmitic acid
parvimosides A, B
patagonicosides A–C
pectenolone
peditoxin
penta decanoic acid
pentactasides I, II and III
pentaregulosides A–G
peroxylucunterine
pervicosides B–D
philinopsides A, B, E, F
phosphatidylcholine
phrygiasterol
phrygioside B
phytosphingosine
pisasterosides A; D–F
plancipyrrosides A, B
plancisides A–D
polyhalogenated monoterpene
polyhydroxysteorids
prepacifenol epoxide
proisocrinins A−F
propanephosphonic acid
protolinckiosides A–D
pseudaboydin A
pseudocnoside A
pseudostichoposides A, B
psilasteroside
psolusosides A, B
ptilomycalin A
pycnopodiosides A–C
pyropheophorbide-a
R
rathbuniosides R1, R2
regularosides A–C; S1, S2
rhodocomatulin 5,7-dimethyl ether
rhodocomatulin 7-methyl ether
ruberosides A−D
S
salsolinol
sanguinosides A–C
santiagoside
saropeptate
scopariosides A–D
sodium 1-(12-hydroxy) octadecanyl sulphate
sodium 24-O-β-d-glucopyranoside 6ʺ-sulfate
spinamine E
spinazarin
spinochromes AE
stichlorosides A1, A2, B1, B2, C1, C2
stichoposides A–D
stichorrenoside A–E
strongylocins 1, 2
strongylostatins 1, 2
412 Compounds
sulfated sterols 1, 2
synallactosides A1, A2, B1, B2, C
synaptosides A, A1
T
temnosides A, B
tenuispinosides A–C
tetradecanoic acid
thelenotosides A, B
thornasreroside A
thornasterol A sulphate
thornasteroside A
tornasterol A sulfate
turquetoside A
typicosides A1, A2, B1, C1, C2
V
Val-Gly-ThrVal-Glu-Met
ValLeu-Leu-Tyr
Val-Thr-Pro-Tyr
variegatuside A–F
versicoside A
violaceusides A, E; G; I–III
X
xanthosine
Z
zeaxanthin
Index
Note: Page numbers in bold refer to tables respectively.
Acanthasteridae (Order: Valvatida): Acanthester planci Arbacia lixula (black sea urchin) 197–8
106–10 Arbaciidae (Order: Arbacioida): Arbacia lixula 197–8
Acanthester planci (crown-of-thorns starfish) 106–10 Arbacioida (Family: Arbaciidae): Arbacia lixula 197–8
Acaudina molpadioides (sea potato) 349–50 Archasteridae (Order: Valvatida):
Achlionice violaecuspidata 293 Archaster typicus 111–13
Actinocucumis typica 244–6 Archaster typicus (common sea star/sand-sifting
Actinopyga agassizi (five-toothed sea cucumber/ star/sand star) 111–13
West Indian sea cucumber/Bahamian sea Aristotle’s lantern 2, 8, 202
cucumber) 294–5 armoured sea cucumber (Thelenota ananas) 366–8
Actinopyga caerulea (blue sea cucumber) 295–6 ashy pink sea cucumber (Holothuria fuscocinerea) 320–2
Actinopyga crassa 296–7 Aslia lefevrei (brown sea cucumber) 247–8
Actinopyga echinites (brown sea cucumber/spiny sea Asterias amurensis (Northern Pacific sea star/Japanese
cucumber/brownfish/deepwater redfish) 297–8 common starfish) 28–30
Actinopyga flammea 298–9 Asterias forbesi (forbes sea star/common sea star) 30–4
Actinopyga lecanora (stonefish/white-rumped sea Asterias nipon 43–7
cucumber/white-bottomed sea cucumber) 300–2 Asterias rathbuni 34–5
Actinopyga mauritiana (surf redfish) 302–4 Asterias rollestoni 35–7
Actinopyga miliaris (blackfish/harry blackfish) 304–5 Asterias rubens (common starfish/common sea star) 38–9
African red knob sea (Protoreaster lincki) 164–6 Asteriidae (Class: Asteroidea; Order: Forcipulatida):
African sea star (Protoreaster lincki) 164–6 Anasterias antarctica 23–5; Aphelasterias
Aleutian star (Leptasterias hylodes) 51–2 japonica 26–7; Asterias amurensis 28–30;
Alexanders sea urchin (Temnopleurus alexandri) 216–17 Asterias forbesi 30–4; Asterias rathbuni
amberfish (Thelenota anax) 369–70 34–5; Asterias rollestoni 35–7; Asterias
ambulacral areas 7 rubens 38–9; Coscinasterias tenuispina 40–2;
Amphilepidida (Class: Ophiuroidea; Family: Diplasterias brucei 42–3; Distolasterias
Ophiolepididae): Ophiolepis superba 171–2 nipon 43–7; Evasterias echinosoma 47–8;
Amphilepidida (Family: Ophiopholidae): Ophiopholis Evasterias retifera 48–9; Leptasterias fisheri
aculeata 172–3; Ophiopholis mirabilis 173–4 50; Leptasterias hylodes 51–2; Leptasterias
Amphilepidida (Family: Ophiotrichidae): Ophiocnemis ochotensis 52–3; Lethasterias fusca 54–6;
marmorata 175; Ophiothrix fragilis 176–7 Lethasterias nanimensis chelifera 56–8;
Anasterias antarctica (cinderella starfish/subantarctic Lysastrosoma anthosticta 58–9; Marthasterias
sea star) 23–5 glacialis 60–2; Pisaster giganteus 62–5
Anasterias minuta 23–5 Asterina pectinifera 113–15
anatomy: Asteroidea 5; Crinoidea 10; echinoderms 2; Asterinidae (Order: Valvatida): Patiria pectinifera 113–15;
Echinoidea 7–8; Holothuroidea 9; Tremaster mirabilis 168–9
Ophiuroidea 6–7 Asteroidea (Order: Forcipulatida) 5–6; see also sea stars
Anneissia bennetti (Bennett’s feather star/Bennett’s (Class: Asteroidea)
rainbow crinoid) 375 Asteropseidae (Order: Valvatida): Asteropsis carinifera
Antarctic basket star (Gorgonocephalus chilensis) 178–9 116–17; Dermasterias imbricata 117–19
Antarctic brittle star (Astrotoma agassizii) 177–8 Asteropsis carinifera (Sheriff-badge sea star) 116–17
Antarctic sea cucumber (Staurocucumis liouvillei) 273–5 Astichopus multifidus (furry sea cucumber/fissured
Antarctic starfish (Neosmilaster georgianus) 69–70 sea cucumber) 357–8
Antarctic sun starfish (Labidiaster annulatus) 66–7 Astropecten indicus (plain sand star) 72–3
Antedonidae (Class: Crinoidea; Order: Comatulida): Astropecten irregularis (sand sea star) 73–4
Heliometra glacialis 373–4 Astropecten latespinosus (sea star) 74–6
Anthenea aspera (cake sea star) 150–3 Astropecten monacanthus (Vietnamese starfish) 76–7
Anthenea chinensis 153–5 Astropecten polyacanthus (sand sifting starfish/comb sea
Anthenea pentagonula 153–5 star/brown spotted combstar) 77–9
Anthenea sibogae 155–6 Astropecten scoparius 80–1
Anthocidaris crassispina 204–5 Astropectinidae (Order: Paxillosida): Astropecten indicus
Aphelasterias japonica 26–7 72–3; Astropecten irregularis 73–4; Astropecten
Apodida (Family: Synaptidae): Synapta maculata 243–4 latespinosus 74–6; Astropecten monacanthus
Apostichopus japonicus (Japanese spiky sea cucumber/ 76–7; Astropecten polyacanthus 77–9;
Japanese sea cucumber) 352–5 Astropecten scoparius 80–1; Craspidaster
Apostichopus parvimensis (warty sea cucumber) 356–7 hesperus 81–3; Psilaster cassiope 83–4
413
414 Index
Astrophyton chilense 178–9 188–9; Ophiocomidae 179–85;

Astrotoma agassizii (predatory snake star/Antarctic Ophiodermatidae 186–8; Ophiolepididae 171–2;
brittle star) 177–8 Ophiomyxidae 186; Ophiopholidae 172–4;
Athyonidium chilensis (burrowing shaggy sea cucumber) Ophiopyrgidae 195–6; Ophiotrichidae 175–7;
248–9 Ophiuridae 190–5
auricularia 9 brownfish (Actinopyga echinites) 297–8
Australasian sea cucumber (Australostichopus mollis) 358–9 brown sandfish: Bohadschia marmorata 311–13;
Australian sea cucumber (Holothuria (Metriatyla) lessoni) Bohadschia vitiensis 315–16
328–31 brown sea cucumber: Actinopyga echinites 297–8; Aslia
Australostichopus mollis (brown sea cucumber/ lefevrei 247–8; Australostichopus mollis 358–9
Australasian sea cucumber/soft sea cucumber/ brown spotted combstar (Astropecten polyacanthus)
New Zealand sea cucumber) 358–9 77–9
auto-evisceration phenomenon 358 brownspotted sandfish (Bohadschia vitiensis) 315–16
brown-tipped sea star (Thromidia catalai) 130–1
Bahamian sea cucumber (Actinopyga agassizi) 294–5 burrowing sea cucumber (Massinium magnum) 276–8
banded brittle star (Ophiolepis superba) 171–2 burrowing shaggy sea cucumber (Athyonidium chilensis)
banded diadema (Diadema savignyi) 229–31 248–9
basket rock sea cucumber (Psolus paradubiosus) 285
bêche-de-mer/gamat 16 cake sea star (Anthenea aspera) 150–3
Bennett’s feather star/Bennett’s rainbow crinoid Camarodonta (Family: Echinidae): Echinus esculentus
(Anneissia bennetti) 375 198–9
blackfish (Actinopyga miliaris) 304–5 Camarodonta (Family: Echinometridae): Echinometra
black knobby sea cucumber (Stichopus chloronotus) 360–1 lucunter 199–201; Echinometra mathaei 201–3;
black longspine urchin (Diadema savignyi) 229–31 Heliocidaris crassispina 204–5; Heliocidaris
black sea cucumber: Holothuria (Halodeima) atra 316–17; erythrogramma 205–6; Heliocidaris
Holothuria (Panningothuria) forskali 319–20; tuberculata 206–7
Holothuria (Mertensiothuria) leucospilota 332–4 Camarodonta (Family: Parechinidae): Paracentrotus
black sea urchin: Arbacia lixula 197–8; Stomopneustes lividus 208–10
variolaris 239–41; Temnopleurus toreumaticus Camarodonta (Family: Strongylocentrotidae):
217–18 Mesocentrotus nudus 210–11;
black-spined star (Lethasterias nanimensis chelifera) 56–8 Strongylocentrotus droebachiensis 211–12;
blackspotted sea cucumber (Pearsonothuria graeffei) 347–9 Strongylocentrotus intermedius 212–13;
black tarzan (Holothuria (Mertensiothuria) leucospilota) Strongylocentrotus pallidus 213–14;
332–4 Strongylocentrotus purpuratus 214–15
black teatfish: Holothuria (Microthele) fuscogilva 322–3; Camarodonta (Family: Temnopleuridae): Temnopleurus
Holothuria (Microthele) nobilis 335–6 alexandri 216–17; Temnopleurus toreumaticus
blood star: Echinaster (Echinaster) sepositus 96–7; 217–18
Henricia leviuscila 101–3 Camarodonta (Family: Toxopneustidae): Lytechinus
blood starfish (Henricia leviuscila) 101–3 variegatus 218–19; Sphaerechinus granularis
blue bat star (Patiria pectinifera) 113–15 219–20; Toxopneustes pileolus 220–2;
blue-black urchin (Echinothrix diadema) 233–4 Tripneustes gratilla 222–4; Tripneustes
blue Linckia (Linckia laevigata) 16, 144–5 ventricosus 225–6
blue phataria starfish (Phataria unifascialis) 149 catala’s sea star (Thromidia catalai) 130–1
blue sea cucumber (Actinopyga caerulea) 295–6 Caudinidae (Order: Molpadida): Acaudina molpadioides
blue sea star (Phataria unifascialis) 149 349–50; Paracaudina chilensis 350;
blue spiny starfish (Coscinasterias tenuispina) 40–2 Pseudostichopus mollis 351
blue star (Linckia laevigata) 16, 144–5 Celerina heffernani 119–20
Bohadschia argus (leopard sea cucumber) 305–6 Cercodemas anceps (red box sea cucumber/pink warty
Bohadschia bivittata (two-ribboned sea cucumber) 307 sea cucumber/spiny sea cucumber/pink sea
Bohadschia cousteaui 308–11 cucumber/green sea cucumber) 249–51
Bohadschia marmorata (brown sandfish/chalky Certonardoa semiregularis 132–42
cucumber/pale marbled sea cucumber) 311–13 chalky cucumber (Bohadschia marmorata) 311–13
Bohadschia subrubra (leopardfish) 313–15 Chilean basket star (Gorgonocephalus chilensis) 178–9
Bohadschia tenuissima 315–16 chocolate chip cucumber (Isostichopus badionotus)
Bohadschia vitiensis (brownspotted sandfish/brown 359–60
sandfish) 315–16 chocolate chip starfish (Protoreaster nodosus) 166–8
bordered sea star (Craspidaster hesperus) 81–3 cinderella starfish (Anasterias antarctica) 23–5
Bradley’s sea star: Mithrodia bradleyi 127–8; Pharia circulatory system 3
pyramidata 148 Cladolabes schmeltzii (Vietnamese sea cucumber/
Breviturma dentata 179–80 Indo-West Pacific sea cucumber) 285–8
Brisaster latifrons (Northern heart urchin/wide heart sea Clypeasteroida (Family: Scutellidae): Scaphechinus
urchin) 236 mirabilis 226–9
brittle stars and basket stars (Class: Ophiuroidea) 4; collector urchin (Tripneustes gratilla) 222–4
Gorgonocephalidae 177–9; Hemieuryalidae Colochirus anceps 249–51
Index 415
Colochirus quadrangularis (thorny sea cucumber) 251–3 269–70; Staurocucumis liouvillei 273–5;
Colochirus robustus (robust sea cucumber/yellow sea Staurocucumis turqueti 275; Thyonidium
cucumber) 253–5 kurilensis 276
Comanthus bennetti 375 Culcita novaeguineae (cushion star) 157–9
Comatula (Validia) rotalaria 376–7 curryfish herrmanni (Stichopus hermanni) 362
Comatulida (Class: Crinoidea; Family: Antedonidae): cushion star (Culcita novaeguineae) 157–9
Heliometra glacialis 373–4 Cyrtocnida (Family: Hemicrinidae): Neogymnocrinus
Comatulida (Family: Comatulidae): Anneissia bennetti richeri 380–1
375; Comatula (Validia) rotalaria 376–7 Cyrtocnida (Family: Holopodidae): Holopus rangii 382–3
Comatulida (Family: Himerometridae): Himerometra
robustipinna 377–8 daisy brittle star (Ophiopholis aculeata) 172–3
Comatulida (Family: Mariametridae): Lamprometra dark sea star (Fromia heffernani) 119–20
palmata 379–80 deepwater redfish (Actinopyga echinites) 297–8
Comatulidae (Order: Comatulida): Anneissia bennetti 375; Dendrochirotida (Family: Cucumariidae):
Comatula (Validia) rotalaria 376–7 Actinocucumis typica 244–6; Aslia lefevrei
comb sea star (Astropecten polyacanthus) 77–9 247–8; Athyonidium chilensis 248–9;
common brittle star (Ophiothrix fragilis) 176–7 Cercodemas anceps 249–51; Colochirus
common fjord starfish (Cosmasterias lurida) 67–9 quadrangularis 251–3; Colochirus robustus
common sea star: Archaster typicus 111–13; Asterias 253–5; Cucumaria conicospermium 255–6;
forbesi 30–4; Asterias rubens 38–9 Cucumaria fallax 257–8; Cucumaria
common sea urchin (Echinus esculentus) 198–9 frondosa 258–61; Cucumaria frondosa
common starfish (Asterias rubens) 38–9 japonica 261–2; Cucumaria koreaensis
cookie-cutter sea star (Ctenodiscus crispatus) 84–5 263; Cucumaria okhotensis 263–4;
cookie dough sea cucumber (Isostichopus badionotus) Hemioedema spectabilis 264–5; Mensamaria
359–60 intercedens 265–7; Pentactella leonina
Coscinasterias tenuispina (blue spiny starfish/white 267; Plesiocolochirus australis 268;
starfish) 40–2 Pseudocnus californicus 271; Pseudocnus
Cosmasterias lurida (common fjord starfish) 67–9 echinatus 271–3; Pseudocolochirus violaceus
cotton-spinner: Holothuria (Panningothuria) forskali 269–70; Staurocucumis liouvillei 273–5;
319–20; Holothuria (Holothuria) tubulosa 346 Staurocucumis turqueti 275; Thyonidium
Craspidaster hesperus (bordered sea star) 81–3 kurilensis 276
crevice brittle star (Ophiopholis aculeata) 172–3 Dendrochirotida (Family: Phyllophoridae): Massinium
Crinoidea 9–11; see also sea lilies/feather stars (Class: magnum 276–8; Pentamera calcigera
Crinoidea) 278–80
crown-of-thorns starfish (Acanthester planci) 106–10 Dendrochirotida (Family: Psolidae): Psolus eximius 281;
Ctenodiscidae (Order: Paxillosida): Ctenodiscus crispatus Psolus fabricii 281–2; Psolus paradubiosus
84–5 285; Psolus patagonicus 283–4
Ctenodiscus crispatus (mud star/cookie-cutter sea star) Dendrochirotida (Family: Sclerodactylidae): Cladolabes
84–5 schmeltzii 285–8; Eupentacta fraudatrix
Cucumaria conicospermium 255–6 288–91; Sclerodactyla briareus 291–2
Cucumaria echinata 271–3 Dermasterias imbricata (leather star) 117–19
Cucumaria fallax (pale sea football) 257–8 Diadema savignyi (long-spined sea urchin/black longspine
Cucumaria frondosa (orange-footed sea cucumber) 258–61 urchin/banded diadema) 229–31
Cucumaria frondosa japonica (Japanese sea cucumber) Diadema setosum (long-spined sea urchin) 231–2
261–2 Diadematidae (Order: Diadematoida): Diadema savignyi
Cucumaria japonica 261–2 229–31; Diadema setosum 231–2; Echinothrix
Cucumaria koreaensis 263 diadema 233–4
Cucumaria lefevrii 247–8 Diadematoida (Family: Diadematidae): Diadema savignyi
Cucumaria okhotensis 263–4 229–31; Diadema setosum 231–2; Echinothrix
Cucumariidae (Order: Dendrochirotida): diadema 233–4
Actinocucumis typica 244–6; Aslia lefevrei diadema urchin (Echinothrix diadema) 233–4
247–8; Athyonidium chilensis 248–9; digestive system 3
Cercodemas anceps 249–51; Colochirus Diplasterias brucei 42–3
quadrangularis 251–3; Colochirus robustus Diplopteraster multipes (pincushion star) 103–4
253–5; Cucumaria conicospermium 255–6; Distolasterias nipon 43–7
Cucumaria fallax 257–8; Cucumaria dragonfish (Stichopus horrens) 363–6
frondosa 258–61; Cucumaria frondosa drugs development 18–19; from brittle stars 20; NP as 19;
japonica 261–2; Cucumaria koreaensis from sea cucumbers 20–1; from sea lilies 21;
263; Cucumaria okhotensis 263–4; from sea stars 19; from sea urchins 20
Hemioedema spectabilis 264–5; Mensamaria Duasmodactyla kurilensis 276
intercedens 265–7; Pentactella leonina Durian sea cucumber (Stichopus horrens) 363–6
267; Plesiocolochirus australis 268;
Pseudocnus californicus 271; Pseudocnus “earthworms of the sea” 356
echinatus 271–3; Pseudocolochirus violaceus Echinaster (Othilia) brasiliensis 93–4
416 Index
Echinaster (Othilia) echinophorus (orange knobby star) forbes sea star (Asterias forbesi) 30–4
97–8 Forcipulatida (Class: Asteroidea; Family: Asteriidae):
Echinasteridae (Order: Spinulosida): Echinaster (Othilia) Anasterias antarctica 23–5; Aphelasterias
brasiliensis 93–4; Echinaster (Othilia) japonica 26–7; Asterias amurensis 28–30;
echinophorus 97–8; Echinaster luzonicus Asterias forbesi 30–4; Asterias rathbuni 34–5;
94–6; Echinaster (Echinaster) sepositus 96–7; Asterias rollestoni 35–7; Asterias rubens 38–9;
Henricia downeyae 98–101; Henricia leviuscila Coscinasterias tenuispina 40–2; Diplasterias
101–3 brucei 42–3; Distolasterias nipon 43–7;
Echinaster luzonicus (Luzon sea star) 94–6 Evasterias echinosoma 47–8; Evasterias retifera
Echinaster (Echinaster) sepositus (Mediterranean red sea 48–9; Leptasterias fisheri 50; Leptasterias
star/purple starfish/blood star) 96–7 hylodes 51–2; Leptasterias ochotensis 52–3;
Echinidae (Order: Camarodonta): Echinus esculentus 198–9 Lethasterias fusca 54–6; Lethasterias
Echinocardium cordatum (sea potato) 234–5 nanimensis chelifera 56–8; Lysastrosoma
echinoderms 1–2; in aquaculture 15; in aquarium anthosticta 58–9; Marthasterias glacialis 60–2;
and souvenir trades 16; in bioindicators of Pisaster giganteus 62–5
environmental quality 17; biology of 2–5; in Forcipulatida (Family: Heliasteridae): Labidiaster
bionics 17; characteristics 5–11; in commercial annulatus 66–7
fisheries 12–15; conservation measures 11; in Forcipulatida (Family: Stichasteridae): Cosmasterias
drugs development 18–21; ecology of 2, 17–18; lurida 67–9; Neosmilaster georgianus 69–70
features 5; as food 15–16; importance of 11–12; Forcipulatida (Family: Zoroasteridae): Myxoderma
as medicine 18; in ornament industry 16; in platyacanthum 70–1
scientific research 16–17 Fromia heffernani (Heffernan’s starfish/Heffernan’s sea
Echinoidea 7–8; see also sea urchins and sand dollars star/dark sea star) 119–20
(Class: Echinoidea) Fromia monilis (necklace starfish/tiled starfish/red tile
Echinometra lucunter (rock boring urchin) 199–201 starfish) 121–2
Echinometra mathaei (rock—burrowing urchin) 201–3 furry sea cucumber (Astichopus multifidus) 357–8
Echinometridae (Order: Camarodonta): Echinometra
lucunter 199–201; Echinometra mathaei 201–3; galloping sea star (Stellaster childreni) 126–7
Heliocidaris crassispina 204–5; Heliocidaris gangliosides 29
erythrogramma 205–6; Heliocidaris garlic bread sea cucumber (Holothuria (Metriatyla)
tuberculata 206–7 scabra) 339–45
Echinothrix diadema (diadema urchin/blue-black giant sea cucumber (Thelenota ananas) 366–8
urchin/horned sea urchin/long-spined giant sea star (Pisaster giganteus) 62–5
urchin) 233–4 giant spined star (Pisaster giganteus) 62–5
Echinus esculentus (European edible sea urchin/common Global Marine Aquarium Database (GMAD) 16
sea urchin) 198–9 Glyptocidaridae (Order: Stomopneustoida): Glyptocidaris
economic importance, Echinoidea 8 crenularis 237–9
eight-armed Luidia sea star (Luidia maculata) 88–9 Glyptocidaris crenularis 237–9
Elasipodida (Family: Elpidiidae): Kolga hyalina 292–3; golden sandfish (Holothuria (Metriatyla) lessoni) 328–31
Rhipidothuria racovitzai 293 Goniasteridae (Order: Valvatida): Fromia heffernani
elephant trunkfish (Holothuria (Halodeima) edulis) 119–20; Fromia monilis 121–2; Hippasteria
323–4 phrygiana 122–6; Rosaster species 126;
Elpidiidae (Order: Elasipodida): Kolga hyalina 292–3; Stellaster childreni 126–7
Rhipidothuria racovitzai 293 Gorgonocephalidae (Order: Euryalida): Astrotoma
Eupentacta fraudatrix (far Eastern sea cucumber) 288–91 agassizii 177–8; Gorgonocephalus chilensis
European edible sea urchin (Echinus esculentus) 198–9 178–9
Euryalida (Family: Gorgonocephalidae): Astrotoma agassizii Gorgonocephalus chilensis (Antarctic basket star/Chilean
177–8; Gorgonocephalus chilensis 178–9 basket star) 178–9
Evasterias echinosoma (spiny net starfish) 47–8 Graeffe’s sea cucumber (Pearsonothuria graeffei) 347–9
Evasterias retifera (Western star) 48–9 gray sea cucumber (Holothuria (Halodeima) grisea) 324–5
evisceration 9 green brittle star (Ophiarachna incrassata) 186
excretory system 3 greenfish sea cucumber (Stichopus chloronotus) 360–1
green sea cucumber (Cercodemas anceps) 249–51
far Eastern sea cucumber (Eupentacta fraudatrix) 288–91 green sea urchin: Lytechinus variegatus 218–19;
far Eastern starfish (Lethasterias fusca) 54–6 Strongylocentrotus droebachiensis 211–12
fat sea star (Thromidia catalai) 130–1 grey sea star (Luidia clathrata) 85–8
feather star/feather zebra (Lamprometra palmata) Gymnocrinus richeri 380–1
379–80
fissured sea cucumber (Astichopus multifidus) 357–8 habitats: Asteroidea 5; Crinoidea 9–10; Echinoidea 7;
five-toothed sea cucumber (Actinopyga agassizi) 294–5 Holothuroidea 9; Ophiuroidea 6
flowerfish (Pearsonothuria graeffei) 347–9 hairy brittle star (Ophiothrix fragilis) 176–7
food and feeding habits: Asteroidea 6; Crinoidea 10; hairy sea cucumber (Sclerodactyla briareus) 291–2
echinoderms 2–3; Echinoidea 8; Holothuroidea Halityle regularis (Mosaic cushion star/regular cushion
9; Ophiuroidea 7 star) 159–61
Index 417
hard-fingered sea cucumber (Sclerodactyla briareus) 291–2 Holothuria (Metriatyla) scabra (sandfish/garlic bread sea
harry blackfish (Actinopyga miliaris) 304–5 cucumber) 339–45
heavy starfish (Thromidia catalai) 130–1 Holothuria species 347
Heffernan’s sea star (Fromia heffernani) 119–20 Holothuria (Holothuria) tubulosa (cotton-spinner/tubular
Heffernan’s starfish (Fromia heffernani) 119–20 sea cucumber) 346
Heliasteridae (Order: Forcipulatida): Labidiaster Holothuriida (Family: Holothuriidae): Actinopyga
annulatus 66–7 agassizi 294–5; Actinopyga caerulea
Heliocidaris crassispina (purple sea urchin) 204–5 295–6; Actinopyga crassa 296–7; Actinopyga
Heliocidaris erythrogramma (purple sea urchin) 205–6 echinites 297–8; Actinopyga flammea 298–9;
Heliocidaris tuberculata (red tipped urchin/tuberculate Actinopyga lecanora 300–2; Actinopyga
urchin) 206–7 mauritiana 302–4; Actinopyga miliaris
Heliometra glacialis 373–4 304–5; Bohadschia argus 305–6; Bohadschia
Heliometra glacialis maxima 373–4 bivittata 307; Bohadschia cousteaui 308–11;
Hemicrinidae (Order: Cyrtocnida): Neogymnocrinus Bohadschia marmorata 311–13; Bohadschia
richeri 380–1 subrubra 313–15; Bohadschia vitiensis 315–16;
Hemieuryalidae (Order: Ophiurida): Ophioplocus Holothuria (Halodeima) atra 316–17; Holothuria
japonicus 188–9 (Halodeima) edulis 323–4; Holothuria
Hemioedema spectabilis (Patagonian sea cucumber) 264–5 (Panningothuria) forskali 319–20; Holothuria
Hemoiedema spectabilis (Patagonian sea cucumber/ fuscocinerea 320–2; Holothuria (Microthele)
Malaysian sea cucumber) 372 fuscogilva 322–3; Holothuria (Microthele)
Henricia downeyae (slender-armed sea star) 98–101 fuscopunctata 317–18; Holothuria (Halodeima)
Henricia leviuscila (Pacific blood star/blood star/blood grisea 324–5; Holothuria (Mertensiothuria)
starfish) 101–3 hilla 325–6; Holothuria (Thymiosycia)
Herman’s sea cucumber (Stichopus hermanni) 362 impatiens 326–8; Holothuria (Metriatyla)
Himerometra magnipinna 377–8 lessoni 328–31; Holothuria (Mertensiothuria)
Himerometra robustipinna 377–8 leucospilota 332–4; Holothuria (Holothuria)
Himerometridae (Order: Comatulida): Himerometra mammata 334; Holothuria (Microthele) nobilis
robustipinna 377–8 335–6; Holothuria (Stauropora) pervicax 336–7;
Hippasteria kurilensis 122–6 Holothuria (Roweothuria) poli 338–9; Holothuria
Hippasteria phrygiana (Kurile spiny star/Pacific starfish/ (Metriatyla) scabra 339–45; Holothuria species
rigid cushion star) 122–6 347; Holothuria (Holothuria) tubulosa 346;
Holopodidae (Order: Cyrtocnida): Holopus rangii 382–3 Pearsonothuria graeffei 347–9
Holopus rangii 382–3 Holothuriidae (Order: Holothuriida): Actinopyga
Holothuria (Halodeima) atra (black sea cucumber/ agassizi 294–5; Actinopyga caerulea
lollyfish) 316–17 295–6; Actinopyga crassa 296–7; Actinopyga
Holothuria axiologa 317–18 echinites 297–8; Actinopyga flammea 298–9;
Holothuria bivittata 315–16 Actinopyga lecanora 300–2; Actinopyga
Holothuria briareus 291–2 mauritiana 302–4; Bohadschia argus
Holothuria (Thymiosycia) decorata 325–6 305–6; Bohadschia bivittata 307; Bohadschia
Holothuria (Halodeima) edulis (elephant trunkfish) 323–4 cousteaui 308–11; Bohadschia marmorata
Holothuria (Halodeima) edulis (pink and black sea 311–13; Bohadschia subrubra 313–15;
cucumber/pink fish sea cucumber) 323–4 Bohadschia vitiensis 315–16; Holothuria
Holothuria (Panningothuria) forskali (black sea (Halodeima) atra 316–17; Holothuria
cucumber/cotton-spinner) 319–20 (Halodeima) edulis 323–4; Holothuria
Holothuria fuscocinerea (ashy pink sea cucumber) 320–2 (Panningothuria) forskali 319–20; Holothuria
Holothuria (Microthele) fuscogilva (black teatfish/white fuscocinerea 320–2; Holothuria (Microthele)
teatfish) 322–3 fuscogilva 322–3; Holothuria (Microthele)
Holothuria (Halodeima) grisea (gray sea cucumber) fuscopunctata 317–18; Holothuria (Halodeima)
324–5 grisea 324–5; Holothuria (Mertensiothuria)
Holothuria (Mertensiothuria) hilla (tiger tail sea hilla 325–6; Holothuria (Thymiosycia)
cucumber) 325–6 impatiens 326–8; Holothuria (Metriatyla)
Holothuria (Thymiosycia) impatiens (impatient sea lessoni 328–31; Holothuria (Mertensiothuria)
cucumber) 326–8 leucospilota 332–4; Holothuria (Holothuria)
Holothuria (Metriatyla) lessoni (golden sandfish/ mammata 334; Holothuria (Microthele)
Australian sea cucumber) 328–31 nobilis 335–6; Holothuria (Stauropora)
Holothuria (Mertensiothuria) leucospilota (black sea pervicax 336–7; Holothuria (Roweothuria)
cucumber/black tarzan) 332–4 poli 338–9; Holothuria (Metriatyla) scabra
Holothuria (Holothuria) mammata 334 339–45; Holothuria species 347; Holothuria
Holothuria nobilis 335–6 (Holothuria) tubulosa 346; Pearsonothuria
Holothuria (Microthele) nobilis (black teatfish) 335–6 graeffei 347–9
Holothuria (Stauropora) pervicax (stubborn sea Holothuroidea 9; see also sea cucumbers (Class:
cucumber) 336–7 Holothuroidea)
Holothuria (Roweothuria) poli (white spot cucumber) horned sea star (Protoreaster nodosus) 166–8
338–9 horned sea urchin (Echinothrix diadema) 233–4
418 Index
impatient sea cucumber (Holothuria (Thymiosycia) madreporite 1, 5–7

impatiens) 326–8 magnum sea cucumber (Massinium magnum) 276–8
Indo-West Pacific sea cucumber: Cladolabes schmeltzii Malaysian sea cucumber (Hemoiedema spectabilis) 372
285–8; Massinium magnum 276–8 marbled brittle star (Ophiocnemis marmorata) 175
Isocrinida (Family: Proisocrinidae): Proisocrinus Mariametridae (Order: Comatulida): Lamprometra
ruberrimus 383–4 palmata 379–80
Isostichopus badionotus (chocolate chip cucumber/ Marthasterias glacialis (spiny starfish) 60–2
cookie dough sea cucumber/three-rowed sea Massinium magnum (magnum sea cucumber/burrowing
cucumber) 359–60 sea cucumber/Indo-West Pacific sea cucumber/
Vietnamese sea cucumber/sea basket sea
Japanese common sea urchin (Strongylocentrotus cucumber) 276–8
intermedius) 212–13 Mediterranean red sea star (Echinaster (Echinaster)
Japanese common starfish (Asterias amurensis) 28–30 sepositus) 96–7
Japanese sea cucumber: Apostichopus japonicus 352–5; Mensamaria intercedens (orange sea cucumber) 265–7
Cucumaria frondosa japonica 261–2 Mesocentrotus nudus (purple sea urchin) 210–11
Japanese smooth brittle star (Ophioplocus japonicus) millipede brittle star (Ophiocoma scolopendrina) 183–4
188–9 Mithrodia bradleyi (Bradley’s sea star) 127–8
Japanese spiky sea cucumber (Apostichopus japonicus) Mithrodia clavigera (nail starfish/nail sea star) 128–9
352–5 Mithrodia enriquetacasoi 127–8
Mithrodiidae (Order: Valvatida): Mithrodia bradleyi
knobbly sea star (Protoreaster nodosus) 166–8 127–8; Mithrodia clavigera 128–9; Thromidia
Kolga hyalina 292–3 catalai 130–1
Kurile sea cucumber (Thyonidium kurilensis) 276 Molpadida (Family: Caudinidae): Acaudina molpadioides
Kurile spiny star (Hippasteria phrygiana) 122–6 349–50; Paracaudina chilensis 350;
Pseudostichopus mollis 351
Labidiaster annulatus (Antarctic sun starfish) 66–7 Mosaic cushion star (Halityle regularis) 159–61
lagoon brittle star (Ophiocoma scolopendrina) 183–4 mud star (Ctenodiscus crispatus) 84–5
Lamprometra klunzingeri 379–80 Myxoderma platyacanthum 70–1
Lamprometra palmata (feather star/feather zebra)
379–80 nail sea star (Mithrodia clavigera) 128–9
leather star (Dermasterias imbricata) 117–19 nail starfish (Mithrodia clavigera) 128–9
Leiaster teres (sunstar starfish/purple linckia starfish/ Narcissia canariensis 146–7
purple starfish/smooth sea star) 143 Narcissia gracilis malpeloensis 147
leopardfish (Bohadschia subrubra) 313–15 natural products (NP) 19
leopard sea cucumber (Bohadschia argus) 305–6 necklace starfish (Fromia monilis) 121–2
Leptasterias fisheri 50 Neogymnocrinus richeri 380–1
Leptasterias hylodes (Aleutian star) 51–2 Neosmilaster georgianus (Antarctic starfish) 69–70
Leptasterias ochotensis 52–3 Neothyonidium magnum 276–8
Lethasterias fusca (far Eastern starfish/warm-water sea nervous system and senses 3
stars) 54–6 New Zealand sea cucumber (Australostichopus mollis)
Lethasterias nanimensis chelifera (Russian starfish/ 358–9
black-spined star) 56–8 Northern heart urchin (Brisaster latifrons) 236
Linckia laevigata (blue Linckia/blue star) 16, 144–5 Northern Pacific sea star (Asterias amurensis) 28–30
lined sea star (Luidia clathrata) 85–8 northern sun star (Solaster endeca) 169–70
lollyfish (Holothuria (Halodeima) atra) 316–17 notched brittle star (Ophiura sarsii) 194–5
long-spined sea urchin: Diadema savignyi 229–31;
Diadema setosum 231–2; Stomopneustes Okinawan sea cucumber (Stichopus chloronotus) 360–1
variolaris 239–41 Ophiacanthida (Family: Ophiocomidae): Breviturma
long-spined urchin (Echinothrix diadema) 233–4 dentata 179–80; Ophiocoma erinaceus 180–2;
Loveniidae (Order: Spatangoida): Echinocardium Ophiocoma schoenleinii 182; Ophiocoma
cordatum 234–5 scolopendrina 183–4; Ophiomastix brocki
Ludwigothuria grisea 324–5 184–5; Ophiomastix mixta 185
Luidia clathrata (grey sea star/lined sea star) 85–8 Ophiacanthida (Family: Ophiodermatidae): Ophioderma
Luidia maculata (eight-armed Luidia sea star) 88–9 longicauda 186–8
Luidia quinaria (spiny sand sea star) 89–90 Ophiacanthida (Family: Ophiomyxidae): Ophiarachna
Luidia sarsii 91–2 incrassata 186
Luidiidae (Order: Paxillosida): Luidia clathrata 85–8; Ophiarachna incrassata (green brittle star) 186
Luidia maculata 88–9; Luidia quinaria 89–90; Ophidiasteridae (Order: Valvatida): Certonardoa
Luidia sarsii 91–2 semiregularis 132–42; Leiaster teres
Luzon sea star (Echinaster luzonicus) 94–6 143; Linckia laevigata 144–5; Narcissia
Lysastrosoma anthosticta (Pacific starfish) 58–9 canariensis 146–7; Narcissia gracilis
Lytechinus variegatus (green sea urchin/variegated sea malpeloensis 147; Pharia pyramidata 148;
urchin) 218–19 Phataria unifascialis 149
Index 419
Ophiocnemis marmorata (marbled brittle star) 175 orange knobby star (Echinaster (Othilia) echinophorus)
Ophiocoma erinaceus (spiny brittle star/Persian Gulf 97–8
brittle star) 180–2 orange sea cucumber (Mensamaria intercedens) 265–7
Ophiocoma schoenleinii (Schoenlein’s brittle star) 182 Oreasteridae (Order: Valvatida): Anthenea aspera
Ophiocoma scolopendrina (lagoon brittle star/millipede 150–3; Anthenea pentagonula 153–5; Anthenea
brittle star) 183–4 sibogae 155–6; Culcita novaeguineae 157–9;
Ophiocomidae (Order: Ophiacanthida): Breviturma Halityle regularis 159–61; Pentaceraster
dentata 179–80; Ophiocoma erinaceus 180–2; regulus 162–4; Protoreaster lincki 164–6;
Ophiocoma schoenleinii 182; Ophiocoma Protoreaster nodosus 166–8
scolopendrina 183–4; Ophiomastix brocki Oreaster nodusus 166–8
184–5; Ophiomastix mixta 185 Oxycomanthus bennetti 375
Ophioderma longicauda (smooth brittle star/snake brittle
star) 186–8 Pacific blood star (Henricia leviuscila) 101–3
Ophiodermatidae (Order: Ophiacanthida): Ophioderma Pacific starfish: Hippasteria phrygiana 122–6;
longicauda 186–8 Lysastrosoma anthosticta 58–9
Ophiolepididae (Class: Ophiuroidea; Order: painted serpent starfish (Ophiolepis superba) 171–2
Amphilepidida): Ophiolepis superba 171–2 pale marbled sea cucumber (Bohadschia marmorata)
Ophiolepis superba (banded brittle star/superb brittle star/ 311–13
painted serpent starfish) 171–2 pale sea football (Cucumaria fallax) 257–8
Ophiomastix brocki 184–5 pale urchin (Strongylocentrotus pallidus) 213–14
Ophiomastix elegans 184–5 Paracaudina chilensis (sand sea cucumber/rattail sea
Ophiomastix mixta 185 cucumber) 350
Ophiomyxidae (Order: Ophiacanthida): Ophiarachna Paracentrotus lividus (purple sea urchin/rock sea urchin/
incrassata 186 stony sea urchin) 208–10
Ophiopholidae (Order: Amphilepidida): Ophiopholis Parechinidae (Order: Camarodonta): Paracentrotus lividus
aculeata 172–3; Ophiopholis mirabilis 208–10
173–4 Patagonian rock sea cucumber (Psolus patagonicus)
Ophiopholis aculeata (crevice brittle star/daisy brittle star) 283–4
172–3 Patagonian sea cucumber: Hemioedema spectabilis 264–5;
Ophiopholis mirabilis 173–4 Hemoiedema spectabilis 372
Ophioplocus japonicus (Japanese smooth brittle star) Patiria pectinifera (blue bat star) 113–15
188–9 paxillae 88
ophiopluteus larvae 173 Paxillosida (Family: Astropectinidae): Astropecten
Ophiopyrgidae (Order: Ophiurida): Ophiosparte gigas indicus 72–3; Astropecten irregularis
195–6 73–4; Astropecten latespinosus 74–6;
Ophiosparte gigas (snake star) 195–6 Astropecten monacanthus 76–7; Astropecten
Ophiothrix fragilis (common brittle star/hairy brittle star) polyacanthus 77–9; Astropecten scoparius
176–7 80–1; Craspidaster hesperus 81–3; Psilaster
Ophiotrichidae (Order: Amphilepidida): Ophiocnemis cassiope 83–4
marmorata 175; Ophiothrix fragilis 176–7 Paxillosida (Family: Ctenodiscidae): Ctenodiscus
Ophiura albida (serpent’s table brittle star) 190–1 crispatus 84–5
Ophiura irrorata 191 Paxillosida (Family: Luidiidae): Luidia clathrata 85–8;
Ophiura (Ophiuroglypha) irrorata irrorata 191 Luidia maculata 88–9; Luidia quinaria 89–90;
Ophiura kinbergi 192–3 Luidia sarsii 91–2
Ophiura leptoctenia 193 Paxillosida (Family: Porcellanasteridae): Styracaster
Ophiura sarsii (notched brittle star) 194–5 caroli 92
Ophiurida (Family: Hemieuryalidae): Ophioplocus peanutfish (Stichopus horrens) 363–6
japonicus 188–9 Pearsonothuria graeffei (blackspotted sea cucumber/
Ophiurida (Family: Ophiopyrgidae): Ophiosparte gigas flowerfish/Graeffe’s sea cucumber) 347–9
195–6 pedicellaria 219
Ophiurida (Family: Ophiuridae): Ophiura albida 190–1; Pentaceraster regulus (spotted sea star) 162–4
Ophiura (Ophiuroglypha) irrorata irrorata 191; Pentacta australis 268
Ophiura kinbergi 192–3; Ophiura leptoctenia Pentacta quadrangularis 251–3
193; Ophiura sarsii 194–5 Pentactella leonina 267
Ophiuridae (Order: Ophiurida): Ophiura albida 190–1; Pentamera calcigera (stony sea cucumber) 278–80
Ophiura (Ophiuroglypha) irrorata irrorata 191; periproct 7–8
Ophiura kinbergi 192–3; Ophiura leptoctenia peristome 7–8, 221
193; Ophiura sarsii 194–5 Persian Gulf brittle star (Ophiocoma erinaceus)
Ophiuroidea 6–7; see also brittle stars and basket stars 180–2
(Class: Ophiuroidea) Pharia pyramidata (yellow-spotted starfish/Bradley’s
orange cushion star (Pteraster pulvillus) 104–5 sea star) 148
orange-footed sea cucumber (Cucumaria frondosa) Phataria unifascialis (blue sea star/tan starfish/blue
258–61 phataria starfish) 149
420 Index
Phyllophoridae (Order: Dendrochirotida): Massinium red tipped urchin (Heliocidaris tuberculata) 206–7
magnum 276–8; Pentamera calcigera 278–80 regeneration ability 4; Asteroidea 6; Ophiuroidea 7
pincushion star (Diplopteraster multipes) 103–4 regular cushion star (Halityle regularis) 159–61
pineapple sea cucumber (Thelenota ananas) 366–8 reproduction system 3; Asteroidea 6; Crinoidea 11;
pink and black sea cucumber (Holothuria (Halodeima) Echinoidea 8; Holothuroidea 9; Ophiuroidea 7
edulis) 323–4 respiratory system 3
pink fish sea cucumber (Holothuria (Halodeima) edulis) Rhipidothuria racovitzai 293
323–4 rigid cushion star (Hippasteria phrygiana) 122–6
pink sea cucumber (Cercodemas anceps) 249–51 robust sea cucumber (Colochirus robustus) 253–5
pink warty sea cucumber (Cercodemas anceps) 249–51 rock boring urchin (Echinometra lucunter) 199–201
Pisaster giganteus (giant sea star/giant spined star) 62–5 rock—burrowing urchin (Echinometra mathaei) 201–3
plain sand star (Astropecten indicus) 72–3 rock sea urchin (Paracentrotus lividus) 208–10
Plesiocolochirus australis 268 Rosaster species 126
pluteus larvae 219 royal sea cucumber (Thelenota anax) 369–70
pointed teat sea cucumber (Thelenota ananas) 366–8 Russian starfish (Lethasterias nanimensis chelifera) 56–8
Porcellanasteridae (Order: Paxillosida): Styracaster
caroli 92 sand dollar (Scaphechinus mirabilis) 226–9
predators 4; of crinoids 10; Echinoidea 8 sandfish (Holothuria (Metriatyla) scabra) 339–45
predatory snake star (Astrotoma agassizii) 177–8 sand fish (Thelenota ananas) 366–8
prickly redfish (Thelenota ananas) 366–8 sand sea cucumber (Paracaudina chilensis) 350
prickly skin cucumber (Thelenota ananas) 366–8 sand sea star (Astropecten irregularis) 73–4
Proisocrinidae (Order: Isocrinida): Proisocrinus sand-sifting star (Archaster typicus) 111–13
ruberrimus 383–4 sand sifting starfish (Astropecten polyacanthus) 77–9
Proisocrinus ruberrimus (sea lily/stone lily) 383–4 sand star (Archaster typicus) 111–13
Protoreaster lincki (red knob sea star/red spine star/ Scaphechinus mirabilis (sand dollar) 226–9
African sea star/African red knob sea) 164–6 scarlet psolus (Psolus fabricii) 281–2
Protoreaster nodosus (horned sea star/chocolate chip Schizasteridae (Order: Spatangoida): Brisaster
starfish/knobbly sea star) 166–8 latifrons 236
Pseudocnus californicus 271 Schoenlein’s brittle star (Ophiocoma schoenleinii) 182
Pseudocnus dubiosus leoninus 267 Sclerodactyla briareus (hairy sea cucumber/hard-fingered
Pseudocnus echinatus 271–3 sea cucumber) 291–2
Pseudocolochirus violaceus (sea apple sea cucumber) Sclerodactylidae (Order: Dendrochirotida): Cladolabes
269–70 schmeltzii 285–8; Eupentacta fraudatrix
Pseudostichopus mollis (sandy sea cucumber) 351 288–91; Sclerodactyla briareus 291–2
Pseudostichopus trachus 351 Scutellidae (Order: Clypeasteroida): Scaphechinus
Psilaster cassiope 83–4 mirabilis 226–9
Psolidae (Order: Dendrochirotida): Psolus eximius 281; sea apple sea cucumber (Pseudocolochirus violaceus)
Psolus fabricii 281–2; Psolus paradubiosus 269–70
285; Psolus patagonicus 283–4 sea basket sea cucumber (Massinium magnum) 276–8
Psolus eximius 281 sea cucumbers (Class: Holothuroidea) 5, 17–18;
Psolus fabricii (scarlet psolus) 281–2 Caudinidae 349–51; Cucumariidae 244–76;
Psolus marcusi 283–4 Elpidiidae 292–3; Holothuriidae 294–349;
Psolus paradubiosus (basket rock sea cucumber) 285 Phyllophoridae 276–80; Psolidae 281–5;
Psolus patagonicus (Patagonian rock sea cucumber) Sclerodactylidae 285–92; species 14–15;
283–4 Stichopodidae 352–70; Synallactidae 370–1;
Pterasteridae (Order: Spinulosida): Diplopteraster Synaptidae 243–4
multipes 103–4; Pteraster pulvillus 104–5 sea lilies/feather stars (Class: Crinoidea) 5;
Pteraster pulvillus (orange cushion star) 104–5 Antedonidae 373–4; Comatulidae 375–7;
purple linckia starfish (Leiaster teres) 143 Hemicrinidae 380–1; Himerometridae 377–8;
purple sea urchin: Heliocidaris crassispina 204–5; Holopodidae 382–3; Mariametridae 379–80;
Heliocidaris erythrogramma 205–6; Proisocrinidae 383–4
Mesocentrotus nudus 210–11; Paracentrotus sea lily (Proisocrinus ruberrimus) 383–4
lividus 208–10; Strongylocentrotus purpuratus sea potato: Acaudina molpadioides 349–50;
214–15 Echinocardium cordatum 234–5
purple spiny urchin (Sphaerechinus granularis) 219–20 sea stars (Class: Asteroidea) 2; Acanthasteridae 106–10;
purple starfish: Echinaster (Echinaster) sepositus 96–7; Archasteridae 111–13; Asteriidae 23–65;
Leiaster teres 143 Asterinidae 113–15, 168–9; Asteropseidae
purple sun star (Solaster endeca) 169–70 116–19; Astropectinidae 72–84; Ctenodiscidae
84–5; Echinasteridae 93–103; Goniasteridae
rattail sea cucumber (Paracaudina chilensis) 350 119–27; Heliasteridae 66–7; Luidiidae 85–92;
red box sea cucumber (Cercodemas anceps) 249–51 Mithrodiidae 127–31; Ophidiasteridae 132–49;
red knob sea star (Protoreaster lincki) 164–6 Oreasteridae 150–68; Porcellanasteridae 92;
red spine star (Protoreaster lincki) 164–6 Pterasteridae 103–5; Solasteridae 169–70;
red tile starfish (Fromia monilis) 121–2 Stichasteridae 67–70; Zoroasteridae 70–1
Index 421
sea urchins and sand dollars (Class: Echinoidea) 2–5, 13; Stichopus chloronotus (Okinawan sea cucumber/greenfish
Arbaciidae 197–8; Diadematidae 229–34; sea cucumber/spiky sea cucumber/black
Echinidae 198–9; Echinometridae 199–207; knobby sea cucumber) 360–1
Glyptocidaridae 237–9; Loveniidae 234–5; Stichopus hermanni (Herman’s sea cucumber/curryfish
Parechinidae 208–10; Schizasteridae herrmanni) 362
236; Scutellidae 226–9; species 13–14; Stichopus horrens (dragonfish/peanutfish/Selenka’s sea
Stomopneustidae 239–41; Strongylocentrotidae cucumber/Durian sea cucumber/Vietnamese
210–15; Temnopleuridae 216–18; sea cucumber) 363–6
Toxopneustidae 218–26 Stichopus japonicus 352–5
Selenka’s sea cucumber (Stichopus horrens) 363–6 Stichopus multifidus 357–8
serpent’s table brittle star (Ophiura albida) 190–1 Stichopus parvimensis 356–7
Sheriff-badge sea star (Asteropsis carinifera) 116–17 Stichopus variegatus 363–6
short-spined sea urchin (Strongylocentrotus intermedius) Stomopneustes variolaris (black sea urchin/long-spined
212–13 sea urchin) 239–41
skeletal system 2 Stomopneustidae (Order: Stomopneustoida):
slender-armed sea star (Henricia downeyae) 98–101 Stomopneustes variolaris 239–41
smooth brittle star (Ophioderma longicauda) 186–8 Stomopneustoida (Family: Glyptocidaridae):
smooth sea star (Leiaster teres) 143 Glyptocidaris crenularis 237–9
smooth sun star (Solaster endeca) 169–70 Stomopneustoida (Family: Stomopneustidae):
snake brittle star (Ophioderma longicauda) 186–8 Stomopneustes variolaris 239–41
snake sea cucumber (Synapta maculata) 243–4 stonefish (Actinopyga lecanora) 300–2
snake star (Ophiosparte gigas) 195–6 stone lily (Proisocrinus ruberrimus) 383–4
soft sea cucumber (Australostichopus mollis) 358–9 stony sea cucumber (Pentamera calcigera) 278–80
Solaster endeca (purple sun star/northern sun star/smooth stony sea urchin (Paracentrotus lividus) 208–10
sun star) 169–70 striped spine sea urchin (Temnopleurus toreumaticus)
Solasteridae (Order: Valvatida): Solaster endeca 169–70 217–18
Spatangoida (Family: Loveniidae): Echinocardium Strongylocentrotidae (Order: Camarodonta):
cordatum 234–5 Mesocentrotus nudus 210–11;
Spatangoida (Family: Schizasteridae): Brisaster latifrons 236 Strongylocentrotus droebachiensis
Sphaerechinus granularis (purple spiny urchin) 219–20 211–12; Strongylocentrotus intermedius
spiky sea cucumber (Stichopus chloronotus) 360–1 212–13; Strongylocentrotus pallidus 213–14;
Spinulosida (Family: Echinasteridae): Echinaster Strongylocentrotus purpuratus 214–15
(Othilia) brasiliensis 93–4; Echinaster Strongylocentrotus droebachiensis (green sea urchin)
(Othilia) echinophorus 97–8; Echinaster 211–12
luzonicus 94–6; Echinaster (Echinaster) Strongylocentrotus intermedius (short-spined sea urchin/
sepositus 96–7; Henricia downeyae 98–101; Japanese common sea urchin) 212–13
Henricia leviuscila 101–3 Strongylocentrotus nudus 210–11
Spinulosida (Family: Pterasteridae): Diplopteraster Strongylocentrotus pallidus (white sea urchin/pale urchin)
multipes 103–4; Pteraster pulvillus 104–5 213–14
spiny brittle star (Ophiocoma erinaceus) 180–2 Strongylocentrotus purpuratus (purple sea urchin) 214–15
spiny net starfish (Evasterias echinosoma) 47–8 stubborn sea cucumber (Holothuria (Stauropora)
spiny sand sea star (Luidia quinaria) 89–90 pervicax) 336–7
spiny sea cucumber: Actinopyga echinites 297–8; Styracaster caroli 92
Cercodemas anceps 249–51 subantarctic sea star (Anasterias antarctica) 23–5
spiny starfish (Marthasterias glacialis) 60–2 sunstar starfish (Leiaster teres) 143
spotted sea star (Pentaceraster regulus) 162–4 superb brittle star (Ophiolepis superba) 171–2
spotted worm sea cucumber (Synapta maculata) 243–4 surf redfish (Actinopyga mauritiana) 302–4
SpStrongylocins 215 Synallactes nozawai 370–1
starfishes 4–5; see also sea stars (Class: Asteroidea) Synallactidae (Order: Synallictida): Synallactes nozawai
Staurocucumis liouvillei (Antarctic sea cucumber) 273–5 370–71
Staurocucumis turqueti 275 Synallictida (Family: Stichopodidae): Apostichopus
Stellaster childreni (galloping sea star) 126–7 japonicus 352–5; Apostichopus parvimensis
Stellaster equestris 126–7 356–7; Astichopus multifidus 357–8;
steroidal components 18 Australostichopus mollis 358–9; Isostichopus
Stichasteridae (Order: Forcipulatida): Cosmasterias lurida badionotus 359–60; Stichopus chloronotus
67–9; Neosmilaster georgianus 69–70 360–1; Stichopus hermanni 362; Stichopus
Stichopodidae (Order: Synallictida): Apostichopus horrens 363–6; Thelenota ananas 366–8;
japonicus 352–5; Apostichopus parvimensis Thelenota anax 369–70
356–7; Astichopus multifidus 357–8; Synallictida (Family: Synallactidae): Synallactes nozawai
Australostichopus mollis 358–9; Isostichopus 370–1
badionotus 359–60; Stichopus chloronotus Synapta maculata (spotted worm sea cucumber/snake sea
360–1; Stichopus hermanni 362; Stichopus cucumber/Vietnamese sea cucumber) 243–4
horrens 363–6; Thelenota ananas 366–8; Synaptidae (Class: Holothuroidea; Order: Apodida):
Thelenota anax 369–70 Synapta maculata 243–4
422 Index
tan starfish (Phataria unifascialis) 149 Valvatida (Family: Goniasteridae): Fromia heffernani
Temnopleuridae (Order: Camarodonta): Temnopleurus 119–20; Fromia monilis 121–2; Hippasteria
alexandri 216–17; Temnopleurus toreumaticus phrygiana 122–6; Rosaster species 126;
217–18 Stellaster childreni 126–7
Temnopleurus alexandri (Alexanders sea urchin) 216–17 Valvatida (Family: Mithrodiidae): Mithrodia bradleyi
Temnopleurus toreumaticus (black sea urchin/striped 127–8; Mithrodia clavigera 128–9; Thromidia
spine sea urchin) 217–18 catalai 130–1
tetrodotoxin (TTX) 80 Valvatida (Family: Ophidiasteridae): Certonardoa
Thelenota ananas (pineapple sea cucumber/tripang/ semiregularis 132–42; Leiaster teres 143; Linckia
prickly skin cucumber/pointed teat sea laevigata 144–5; Narcissia canariensis 146–7;
cucumber/armoured sea cucumber/giant sea Narcissia gracilis malpeloensis 147; Pharia
cucumber/sand fish/prickly redfish) 366–8 pyramidata 148; Phataria unifascialis 149
Thelenota anax (amberfish/royal sea cucumber) 369–70 Valvatida (Family: Oreasteridae): Anthenea aspera
thorny sea cucumber (Colochirus quadrangularis) 150–3; Anthenea pentagonula 153–5; Anthenea
251–3 sibogae 155–6; Culcita novaeguineae 157–9;
three-rowed sea cucumber (Isostichopus badionotus) Halityle regularis 159–61; Pentaceraster
359–60 regulus 162–4; Protoreaster lincki 164–6;
Thromidia catalai (catala’s sea star/brown-tipped sea star/ Protoreaster nodosus 166–8
heavy starfish/fat sea star) 130–1 Valvatida (Family: Solasteridae): Solaster endeca 169–70
Thyone briareus 291–2 variegated sea urchin (Lytechinus variegatus) 218–19
Thyonidium kurilensis (Kurile sea cucumber) 276 venomous/poisonous echinoderms 4–5
tiger tail sea cucumber (Holothuria (Mertensiothuria) Vietnamese sea cucumber: Cladolabes schmeltzii 285–8;
hilla) 325–6 Massinium magnum 276–8; Stichopus horrens
tiled starfish (Fromia monilis) 121–2 363–6; Synapta maculata 243–4
Toxopneustes pileolus (flower urchin) 220–2 Vietnamese starfish (Astropecten monacanthus) 76–7
Toxopneustidae (Order: Camarodonta): Lytechinus
variegatus 218–19; Sphaerechinus granularis warm-water sea stars (Lethasterias fusca) 54–6
219–20; Toxopneustes pileolus 220–2; warty sea cucumber (Apostichopus parvimensis) 356–7
Tripneustes gratilla 222–4; Tripneustes water vascular system 1, 5
ventricosus 225–6 Western star (Evasterias retifera) 48–9
Tremaster mirabilis 168–9 West Indian sea cucumber (Actinopyga agassizi) 294–5
Tremaster novaecaledonia 168–9 West Indian sea egg (Tripneustes ventricosus) 225–6
tripang (Thelenota ananas) 366–8 white-bottomed sea cucumber (Actinopyga lecanora)
Tripneustes gratilla (collector urchin) 222–4 300–2
Tripneustes ventricosus (West Indian sea egg/white sea white-rumped sea cucumber (Actinopyga lecanora) 300–2
urchin) 225–6 white sea urchin: Strongylocentrotus pallidus 213–14;
tuberculate urchin (Heliocidaris tuberculata) 206–7 Tripneustes ventricosus 225–6
tubular sea cucumber (Holothuria (Holothuria) white spot cucumber (Holothuria (Roweothuria) poli)
tubulosa) 346 338–9
two-ribboned sea cucumber (Bohadschia bivittata) 307 white starfish (Coscinasterias tenuispina) 40–2
white teatfish (Holothuria (Microthele) fuscogilva) 322–3
Valvatida (Family: Acanthasteridae): Acanthester planci wide heart sea urchin (Brisaster latifrons) 236
106–10 Wnt signalling pathway 191
Valvatida (Family: Archasteridae): Archaster typicus
111–13 yellow sea cucumber (Colochirus robustus) 253–5
Valvatida (Family: Asterinidae): Patiria pectinifera yellow-spotted starfish (Pharia pyramidata) 148
113–15; Tremaster mirabilis 168–9
Valvatida (Family: Asteropseidae): Asteropsis carinifera Zoroasteridae (Order: Forcipulatida): Myxoderma
116–17; Dermasterias imbricata 117–19 platyacanthum 70–1

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Biology and Ecology of

Pharmaceutical Marine Life:

Biology and Ecology of Edible Marine Bivalve Molluscs

Biology and Ecology of Edible Marine Gastropod Molluscs

Biology and Ecology of Pharmaceutical Marine Cnidarians

Biology and Ecology of Pharmaceutical Marine Life: Echinoderms

Biology and Ecology of Pharmaceutical Marine Mollusks

Biology and Ecology of Pharmaceutical Marine Plants

Biology and Ecology of Pharmaceutical Marine Sponges

Biology and Culture of Portunid Crabs of World Seas

Biology and Ecology of Toxic Pufferfish

Biology and Ecology of Venomous Marine Snails

Biology and Ecology of Venomous Stingrays

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1.6.10 Echinoderms as Sources of Pharmaceutical (Bioactive)

Chapter 2 Biology and Ecology of Pharmaceutical Sea Stars (Class: Asteroidea)...................... 23

Chapter 6 Biology and Ecology of Pharmaceutical Sea Lilies or Feather Stars

1.2 ECOLOGY OF ECHINODERMS

1.3 BIOLOGY OF ECHINODERMS

1.3.2 Food and Feeding Habits

1.3.4 Nervous System and Senses

1.3.5 Circulatory System

Purple sea urchin or stony sea urchin, Paracentrotus lividus

1.4 CHARACTERISTICS OF ECHINODERMS

1.4.2 Characteristics of Different Classes of Echinoderms

1.4.2.1 Class Asteroidea

1.4.2.1.3 Food and Feeding

1.4.2.1.5 Regenerative Ability

1.4.2.2 Class Ophiuroidea

1.4.2.2.3 Food and Feeding

1.4.2.2.5 Regenerative Ability

1.4.2.3 Class Echinoidea

1.4.2.3.3 Food and Feeding

1.4.2.3.6 Economic Importance

1.4.2.3.7 Research Use

1.4.2.4 Class Holothuroidea

1.4.2.4.3 Food and Feeding

1.4.2.5 Class Crinoidea

1.4.2.5.3 Food and Feeding

1.5 CONSERVATION MEASURES OF ECHINODERMS

1.6 IMPORTANCE OF ECHINODERMS

1.6.1 Echinoderms in Commercial Fisheries

Source: Keesing and Hall, 1998.

Species Fishery Distribution

Source: Micael et al., 2009.

About 42 species of sea cucumbers (“bêche-de-mer” or “trepang”) are fished commercially

1.6.1.2 Sea Cucumber Species that Contribute to the Modern Fishery

Species Fishery Distribution

Source: Micael et al., 2009.

1.6.2 Echinoderms in Aquaculture (Farming)

1.6.4 Echinoderms in the Aquarium and Souvenir Trades

1.6.4.1 Potential Echinoderms in the Ornament Industry

Source: Micael et al., 2009.

1.6.5 Echinoderms in Scientific Research (Experimental Model Systems)

1.6.7 Echinoderms in Bioindicators of Environmental Quality

1.6.8 Ecological Role of Echinoderms

1.6.10 Echinoderms as Sources of Pharmaceutical (Bioactive)

anticoagulant, antifungal, antimalarial, antiprotozoal, anti-tuberculosis, anti-inflammatory, antitu-

1.6.10 Echinoderms as Sources of Pharmaceutical (Bioactive)

1.6.10.1 Present Status of New Natural Products (NP) as