A Guide to
Cotntnon
Echinodertns
of Andaman and
Nicobar Islands
A Guide to
Common Echinoderms
of Andaman and Nicobar Islands
C. RAGHUNATHAN, KOUSHIK SADHUKHAN,
TAMAL MONDAL, C. SIVAPERUMAN AND
K. VENKATARAMAN*
Zoological Survey of India, Andaman and Nicobar Regional Centre,
Raddo, Port Blair - 744102, Andaman and Nicobar Islands
*Zoological Survey of India, Prani Vigyan Bhawan, M-Block,
New Alipore, Kolkata - 700053
ZOOLOGICAL SURVEY OF INDIA
KOLKATA
Citation
Raghunathan, C., Sadhukhan, K., Mondal, T., Sivaperuman, C.,
Venkataraman, K., 2013. A Guide to Common Echinoderms of
Andaman and Nicobar Islands: 1-210, (Published by the Director,
Zool. Surv. India, Kolkata)
Published: July, 2013
ISBN 978-81-8171-338-4
© Govt. of India, 2013
All Rights Reserved
•
No part of this publication may be reproduced, stored in a
retrieval system or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording or otherwise
without the prior permission of the publisher.
•
This book is sold subject to the condition that it shall not, by
way of trade, be lent, re-sold hired out or otherwise disposed of
without the publisher's consent, in any form of binding or cover
other than that in which it is published.
•
The correct price of this publication is the price printed on
this page. Any revised price indicated by a rubber stamp or
by a sticker or by any other means is incorrect and should be
unacceptable.
Price
India Rs. 1550.00
Foreign $ 85.00; £ 60.00
Published at the Publication Division by the Director, Zoological
Survey of India, M-Block, New Alipore, Kolkata-700 053 and
printed at Calcutta Repro Graphics, Kolkata-700 006.
CONTENTS
I.
GENERAL INTRODUCTION
1
2.
LITERATURE REVIEW
11
3.
CHARACTERISTICS OF EXTANT CLASSES
24
4.
STUDY AREAS
32
5.
MATERIAL AND METHODS
38
6.
KEY CHARACTERS
39
7.
DESCRIPTIONS
59
Class CRINOIDEA
I. Comanthus parvicirrus
60
2. Comanthina nobilis
61
3. Comaster schlegeli
62
4. Comaster multibrachiata
63
5. Oxycomanthus bennetti
64
6. Himerometra robustipinna
65
7. Heterometra philiberti
66
8. Heterometra crenulata
67
9. Amphimetra molleri
68
10. Lamprometra palmata
69
II. Cenometra bella
70
12. Cenometra emendatrix
71
13. Oligometra serripinna
72
14. Ponti metra andersoni
73
Class ASTEROIDEA
15. Asterina sarasini
75
16. Stellaster equestris
76
17. Culcita novaguineae
77
18. Culcita schmideliana
78
19. Choriaster granulatus
79
20. Anthenea tuberculosa
80
21. Protoreaster lincki
81
22. Protoreaster nodosus
82
23. Pentaceraster regulus
83
24. Asteropsis carinifera
84
25. Acanthaster planci
85
26. Fromia monilis
86
27. Fromia indica
87
28. Linckia guildingi
88
29. Linckia laevigata
89
30. Linckia multifora
90
31. Echinaster luzonicus
91
32. Astropecten indicus
92
33. Astropecten monacanthus
93
34. Luidia maculata
94
Class OPHIUROIDEA
(iv)
35. Ophiocoma erinaceus
96
36. Ophiocoma dentata
97
37. Ophiocoma scolopendrina
98
38. Ophiarthrum pictum
99
39. Ophiomastix annulosa
100
40. Macrophiothrix propinqua
101
41. Macrophiothrix longipeda
102
42. Ophiarachna incrassata
103
Class ECHINOIDEA
43. Prionocidaris verticillata
105
44. Arbacia puntulata
106
45. Diadema setosum
107
46. Diadema savignyi
108
47. Echinothrix calamaris
109
48. Echinothrix diadema
110
49. Echinometra oblonga
111
50. Echinometra mathaei
112
51. Heterocentrotus trigonarius
113
52. Echinodiscus auritus
114
53. Temnopleurus alexendri
115
54. Mespilia globulus
116
55. Microcyphus ceylanicus
117
56. Salmacis belli
118
57. Tripneustes gratilla
119
58. Stomopneustus variolar is
120
59. Clypeaster humilis
121
Class HOLOTHUROIDEA
60. Actinopyga mauritiana
123
61. Actinopyga miliaris
124
62. Actinopyga lacanora
125
63. Actinopyga echinities
126
64. Holothuria cinerascens
127
65. Holothuria (Thymiosycia) hilla
129
66. Holothuria (Thymioscycia) impatiens
131
67. Holothuria atra
133
68. Holothuria (Acanthotrapeza) pyxis
134
69. Holothuria leucospilota
136
70. Holothuria (Mertensiothuria) fuscocinerea
138
(v)
(vi)
71. Holothuria (Metriatyla) scabra
139
72. Holothuria (Halodeima) edulis
141
73. Holothuria (Microthele) nobilis
143
74. Holothuria coluber
144
75. Holothuria (Mertensiothuria) pervicax
145
76. Holothuria arenicola
147
77. Holothuria marmorata
148
78. Bohadschia argus
149
79. Bohadschia graeffei
150
80. Labidodemas semperianum
152
81. Stolus buccalis
153
82. Thelenota annas
154
83. Stichopus horrens
155
84. Stichopus chloronotus
156
85. Stichopus hermanni
157
86. Stichopus vastus
158
87. Synapta maculata
159
88. Euapta godreffroyi
160
89. Acaudina malpadioides
161
CHECKLIST OF ECHINODERMS OF INDIA
162
ACKNOWLEDGEMENTS
192
REFERENCES
192
1. GENERAL INTRODUCTION
Preamble
The Ehinodermata IS one of the best characterized
and most distinct phyla of animal kingdom (Bather,
1900). The echinoderms being common and conspicuous
marine animals have been known since ancient times.
They are found at every ocean depth, from the intertidal
zone to the abyssal zone. The first definitive members
of the phylum appeared near the start of the Cambrian
period. The echinoderms are important both biologically
and geologically: biologically because few other groupings
are so abundant in the biotic desert of the deep sea,
as well as the shallower oceans, and geologically as
their ossified skeletons are major contributors to many
limestone formations, and can provide valuable clues as
to the geological environment. Further, it is held by some
that the radiation of echinoderms was responsible for the
Mesozoic revolution of marine life.
The word "echinoderm" is derived from the Greek
word ゥxセカPXエーjャ。キ@
(Echinodermata), plural of ゥxセカVFpjャ。@
(Echinoderma), "spiny skin" from ゥxセカV@
(echinos),
"hedgehog," and 8tpJla (derma), "skin". The name
Echinodermata appears to have originated with Jacob
Klein (1734), who however applied it only to echinoids.
Linnaeus in the 10th edition (1758) of his Systema naturae
relegated all invertebrates except insects to one class,
Vermes. The Echinodermata, (from the Greek meaning
spiny skin), is a phylum containing some 13,000 extinct
and 7,000 extant species. Living representatives are only
found in marine environment, making the echinodermata
the largest phylum lacking in terrestrial and freshwater
forms. Echinoderms evolved from bilaterally symmetric
animals exhibiting five fold radial symmetry in portions of
their body at some stage oflife. Echinoderms can reversibly
1
vary the rigidity of their dermis and general connective
tissue. They are pentamerous symmetrical animals where
the body can be divided into five similar parts arranged
around the central axis. The radial symmetry, however,
evolves within the echinoderms and does not indicate a
close evolutionary relationship with others.
Definition
The Echinodermata or echinoderms are enterocoelous
coelomates, having a pentaradiate construction derived
from an original bilaterality, without definite head or
brain, with a calcareous endoskeleton of separate plates
or pieces, often bearing external spines or protuberances,
and with a water-vascular system of coelomic nature that
sends numerous small projections (podia) to the exterior
and communicates with the external medium by a pore or
cluster of pores, at least in juvenile stages (Hymen,1955).
Body wall
The body wall of echinoderms consists of three
layers. The outer layer, called the epidermis, is only
a single layer of cells which covers the entire animal
including its various spines. The third layer is also a
single layer of cells the main difference being that these
cells are ciliated. This layer encloses the animal's coelom
separating the animal's guts from its skin. It is called the
'coelomic lining'. The middle layer is much thicker and
is called the dermis. It is composed of connective tissue
and contains the endoskeleton. This endoskeleton takes
three different forms: a set of closely joint plates with
little individual movement that exist as a test or shell
(Sea urchins), a set of separately articulating (more freely
moving) small plates called ossicles (Seastar, Brittle Stars
and the arms of Crinoids), and a collection of widely
2
separated microscopic ossicles lying in the lethery dermis
(Sea cucumbers). Whatever form they take these plates
or ossicles are always made from calcite, the ingredients
for which are found in sea water. The endoskeleton of
echinoderms grows continuously throughout the animals
life, thus older animals are always larger than younger
ones. The endoskeleton supports the spines, warts and
tubercles that are often found on the echinoderm surface.
These various protuberances are also generally made
from calcite.
Endoskeleton
Echinoderms have a mesodermal skeleton composed
of calcareous plates or ossicles. Despite the robustness
of the individual skeletal modules, complete echinoderm
skeletons are rare in the fossil record. This is because
they quickly disarticulate once the encompassing skin rots
away, and in the absence of tissue there is nothing to hold
the plates together. The modular construction is a result of
the growth system employed by echinoderms, which adds
new segments at the centre of the radial limbs, pushing
the existing plates outwards in the fashion of a conveyor
belt. The spines of sea urchins are most readily lost, as
each spine can be moved individually and is only loosely
attached in life. A walk above a rocky shore will often
reveal a large number of spineless but otherwise complete
sea urchin skeletons. Echinoderm skeletons are made
up of interlocking calcium carbonate plates and spines.
This skeleton is enclosed by the epidermis and is thus an
endoskeleton. In some, such as sea urchins, the plates fit
together tightly. In others, such as starfish, the plates are
more loosely bound, and in sea cucumbers the plates are
usually microscopic. But whatever their shape, the plates
of echinoderms have a very typical microstructure: electron
3
microscopy reveals them to be, not solid blocks, but fine
networks of calcium carbonate forming a structure known
as stereom. Each skeletal element of an echinoderm is
actually a single crystal of calcium carbonate, very finely
branched and structured. Between the skeletal plates, a
number of special structures protrude, with which the
echinoderm breathes, moves, and defends itself. Typically,
these are tube feet, pedicellaria, and gills.
Water-vascular System
Echinoderms possess a unique water vascular or
"ambulacral" system. The water vascular system of the
echinoderms is unique in the living world and easily
distinguishes them from all other phyla. This is a network
of fluid-filled canals that function in gas exchange, feeding,
and secondarily in locomotion. This system is derived from
both the hydrocoel and axocoel. This system may have
allowed echinoderms to function without the gills found
in other deuterostomes. The system comprises a central
ring, the hydrocoel, and radial ambulacra stretching
along the body or arms. There are extensions (tube feet,
papulae etc.) of the water vascular system which project
out through holes in the skeleton and can be extended or
contracted by the redistribution of fluid between the foot
and internal sac.
The system takes slightly different forms in the
different classes. In the Crinoidea, which are believed to
be the most ancient of the echinoderms, the tube feet are
branched and secrete mucous. In the Ophiuroidea the tube
feet are simple and slender. In Asteroidea, Echinoidea
and Holothuroidea they are thicker and end in suckers.
The water vascular system starts with an opening to the
external environment called a madreporite. From this a
short straight canal called the 'stone canal' leads to the
4
'ring canal'. The ring canal is a ring as might be expected
and it has five longitudinal canals branching off from it
into each of the arms, or their morphological equivalents
in Echinoidea and Holothuroidea. In species with more
than 5 arms these canals branch out into each arm. On
each side of each canal there arises a series of short lateral
canals that lead, via a valve, into the descending tube feet
and the ampulla that operate them. The tube feet pass
through small holes in the animal's endoskeleton and
muscles around the ampulla (which remains inside the
endoskeleton) squeeze water into them causing them to
extend or relax.
Reproduction
Sexual reproduction: Echinoderms become sexually
mature after approximately two to three years, depending
on the species and the environmental conditions. The
eggs and sperm cells are released into open water, where
fertilization takes place. The release of sperm and eggs is
coordinated temporally at same location and spatially at
different regions. Internal fertilization has currently been
observed in three species of sea star, three brittle stars
and a deep water sea cucumber. In some species of feather
star, the embryos develop in special breeding bags, where
the eggs are held until sperm released by a male happen
to find them and fertilize the contents. This can also
be found among sea urchins and sea cucumbers, where
exhibit care for their young can occur, for instance in a
few species of sand dollars who carry their young between
the pricks of their oral side, and heart urchins possess
breeding chambers. With brittle stars, special chambers
can be developed near the stomach bags, in which the
development of the young takes place. Species of sea
cucumbers with specialized care for their offspring may
5
also nurse the young in body cavities or on their surfaces.
In rare cases, direct development without passing through
a bilateral larval stage can occur in some sea stars and
brittle stars. Another strategy that has evolved in some
sea stars and brittle stars is the ability to reproduce
asexually by dividing in two halves while they are small
juveniles, while turning to sexual reproduction when they
have reached sexual maturity.
Asexual reproduction: Many echinoderms have
remarkable powers of regeneration. Some sea stars are
capable of regenerating lost arms. In some cases, lost
arms have been observed to regenerate a second complete
sea star. Sea cucumbers often discharge parts of their
internal organs if they perceive danger. The discharged
organs and tissues are quickly regenerated. Seaurchins
are constantly losing their spines through damage all parts
are replaceable. Some seastar populations can reproduce
entirely asexually purely by the shedding of arms for long
periods of time.
Larval development: The development of an
echinoderm begins with a bilaterally symmetrical embryo,
with a coeloblastula developing first. Gastrulation marks
the opening of the "second mouth" that places them
within the deuterostomes, and the mesoderm, which will
host the skeleton, migrates inwards. The secondary body
cavity, the coelom, forms by the partitioning of three body
cavities. Upon metamorphosis, each taxon produces a
distinct larva, the left hand side of which develops into
the adult organism, the right hand side eventually being
absorbed; the left hand side typically becomes the oral
plate. The different larval types such as Bipinnaria &
Brachiolaria, Auricularia, Ophiopluteus, Echinopluteus
and Vitellaria for Asteriods, Holothuroids, Ophiuroids,
Echinoids and Crinoids respectivley are classified.
6
Feeding
Echinoderms are filter feeders, su.bstrate eaters or
carnivores. The gut is V-shaped in the Crinoidea with
the mouth and anus being on the same surface. In the
other groups such as Echinoidea and Holothuroidea,
it is straight-through gut with the mouth and anus on
approximately opposite sides of the body. Echinoderms
also have a spacious coelom (an open, fluid-filled body
cavity lined with tissue), large gonads, and (usually) a
complete gut. Many seastar have the peculiar ability to
feed by turning the stomach inside out through the mouth;
sea urchins scrape algae from rocks with five large teeth
arranged in a structure known as "Aristotle's lantern."
The modes of feeding vary greatly between the
constituent taxa. Crinoids and some brittle stars
tend to be passive filter-feeders, absorbing suspended
particles from passing water; sea urchins are grazers,
sea cucumbers deposit feeders, and most seastars are
carnivores. Crinoids employ a large net-like structure to
sieve water as it is swept by currents, and to absorb any
particles of matter sinking from the ocean overhead. Once
a particle touches the arms of the creature, the tube feet
act to swish it to the central mouth of the crinoid, where
it is ingested, nutrients removed, and the remains egested
through its anus to the water column. Many sea urchins
graze on the surfaces of rocks, scraping off the thin layer
of algae covering the surfaces. Sand dollars may perform
suspension or sediment feeding. Sea cucumbers may be
suspension feeders, sucking vast quantities of sea water
through their guts and absorbing any useful matter. Others
use their feeding apparatus to actively capture food from
the sea floor. Yet others deploy their feeding apparatus
as a net, in which smaller organisms become ensnared.
While some sea stars are detrivores, extracting the organic
material from mud, and others mimic the crinoids' filter
7
feeding, attacking other sea stars or shellfish. The latter
are seized and held by the tube feet; sea stars then stiffen
their legs, expanding the shell. The sea stars can use
connective tissue to lock their arms in place and maintain
a force on the prey whilst exerting minimal effort; the
unfortunate victim must expend energy resisting the force
with its adductor muscle. When the adductor tires, the
sea star can insert its stomach through the opening and
release gastric juices, digesting the prey alive.
Despite their low nutrition value and the abundance
of indigestible calcite, many organisms, such as crabs,
sharks, sea birds and larger starfish, make a living by
feeding on echinoderms. Defensive strategies employed
include the presence of spines, toxins, which can be
inherent or delivered through the tube feet, and the
discharge of sticky entangling threads by sea cucumbers.
Being stabbed by a sea urchin may result in painful injury.
Distribution and habitat
Echinoderms are globally distributed in almost all
depths, latitudes and environments in the ocean. They
reach highest diversity in reef environments but are also
widespread on shallow shores, around the poles refugia
where crinoids are at their most abundant, and throughout
the deep ocean, where bottom-dwelling and burrowing
sea cucumbers are common, sometimes accounting for up
to 90% of organisms. Whilst almost all echinoderms are
benthic, that is, they live on the sea floor and some sealilies can swim at great velocity for brief periods of time,
and a few deep-sea sea cucumbers are fully floating. Some
crinoids are pseudo-planktonic, attaching themselves
to floating logs and debris, although this behaviour
was exercised most extensively in the Paleozoic, before
competition from such organisms as barnacles restricted
the extent of the behaviour. Some sea cucumbers employ
8
a similar strategy, hitching lifts by attaching to the sides
of fish. The larvre of many echinoderms, especially starfish
and sea urchins, are pelagic, and with the aid of ocean
currents can swim great distances, reinforcing the global
distribution of the phylum.
Ecology
Echinoderms provide a key ecological role in
ecosystems. The grazing of sea urchins reduces the rate
of colonization of bare rock; the burrowing of sand dollars
and sea cucumbers depleted the sea floor of nutrients and
encouraged deeper penetration of the sea floor, increasing
the depth to which oxygenation occurs and allowing a
more complex ecological tiering to develop. Seastar and
brittle stars prevent the growth of algal mats on coral
reefs, which would obstruct the filter-feeding constituent
organisms. Some sea urchins can bore into solid rock; this
bioerosion can destabilise rock faces and release nutrients
into the ocean. The echinoderms are also the staple diet
of many organisms, most notably the otter; conversely,
many sea cucumbers provide a habitat for associates,
including crabs, worms and snails. The extinction of
large quantities of echinoderms appears to have caused a
subsequent overrunning of ecosystems by seaweed, or the
destruction of an entire reef.
Evolution
The first universally accepted echinoderms appear
the Lower Cambrian period (Paul and Smith 1984).
Echinoderms left behind an extensive fossil record.
Despite this, there are numerous conflicting hypotheses
on their phylogeny. Based on their bilateral larvae, many
zoologists argue that echinoderm ancestors were bilateral
and that their coelom had three pairs of spaces (trimeric).
III
9
Some have proposed that radial symmetry arose in a freemoving echinoderm ancestor and that sessile groups were
derived several times independently from free-moving
ancestors. Unfortunately, this view does not address the
significance of radial symmetry as an adaptation for a
sessile existence.
The more traditional view is that the first echinoderms
were sessile, became radial as an adaptation to that
existence, and then gave rise to free-moving groups. This
view perceives the evolution of endoskeletal plates with
stereom structure and of external ciliary grooves for
feeding as early echinoderm developments.
The extinct members of Class Homalozoa, commonly
referred to as carpoids, had stereom ossicles but were
not radially symmetrical, and the status of their watervascular system is not known. Further, extinct members
of the Class Helicoplacoidea possessed three, true
ambulacral grooves, and their mouth was on the side of
their body.
Attachment to a substratum would have selected for
radial symmetry and may have marked the origin of the
Class Crinoidea. Members of Crinoidea, along with the
extinct members of Class Cystoidea, were primitively
attached to a substratum by an aboral stalk. An ancestor
that became free-moving might have given rise to
Asteroidea, Ophiuroidia, Holothuroidea, and Echinoidea.
Economic importance
Echinoderms are also elements of many CUIsmes.
Around 50,000 tons of sea urchins are captured each
year, the gonads of which are consumed particularly in
Japan, Peru and in France. The taste is described as soft
and melting, like a mix of seafood and fruit. The quality
10
depends on the colour, which can range from light yellow
to bright orange.
Economically, sea cucumbers are important in two
main ways. First, some species produce toxins that are
of interest to pharmaceutical firms seeking to learn their
medical value. Some compounds isolated to date exhibit
antimicrobial activity or act as anti-inflammatory agents
and anticoagulants. Second, as a gourmet food item in the
orient, they form the basis of a multimillion-dollar industry
that processes the body wall for sale as beche-de-mer or
trepang. However, the high value of some species, the
ease with which such shallow-water forms can be collected
and their top-heavy age structures all contribute to overexploitation and collapse of the fisheries in some regions.
Sea cucumbers are also considered a delicacy in some
countries of southeast Asia; particularly popular are the
pineapple roller Thelenota ananas and the red Halodeima
edulis. They are well known as heche de mer or Trepang
in China and Indonesia. The sea cucumbers are dried, and
the potentially poisonous entrails removed. The strong
poisons of the sea cucumbers are often psychoactive, but
their effects are not well studied. It does appear that some
sea cucumber toxins restrain the growth rate of tumour
cells, which has sparked interest from cancer researchers.
The calcareous tests or shells of echinoderms are used
as a source of lime by farmers in areas where limestone
is unavailable; indeed 4,000 tons of the animals are used
annually for this purpose. This trade is often carried out in
conjunction with shellfish farmers, for whom the starfish
pose a major irritation by eating their stocks
2. LITERATURE REVIEW
The
Phylum
Echinodermata,
comprIsmg
approximately 7,000 living species, and 13,000 fossil
11
species, is epitomized by the familiar sea star, a universal
symbol of the marine realm. The history of the phylum
is fraught with misconceptions. Linnaeus (1758) did
not recognize the echinoderms as a separate group, and
placed the echinoderms that were known to him in his
"Mollusca", a subdivision of "Vermes". Bruguiere (1791)
revived Klein's (1734) name Echinodermata-a short-lived
independence for the group, for Lamarck (1801) referred
the echinoderms to his "Radiata", where they stayed for
several decades until finally Leuckart (1854) successfully
established the Echinodermata as a distinct phylum. Over
the past 160 years, progress on the higher classification of
the extant and fossil echinoderms has been fairly steady,
with such authors as Ludwig (1889-1907), Bather (1900),
Cuenot (1948), and Hyman (1955) providing authoritative
summaries of classification history. The numerous
authors contributed to the Echinodermata volumes of the
Treatise on Invertebrate Paleontology (Moore, 1966-1978)
to revise the groups for which they were responsible, so
that the 1960's became an era of great change in our
knowledge of the phylum. Publication of the Treatise
volumes stimulated much research work, particularly on
fossil groups. New and exciting approaches to taxonomy
have resulted in a reassessment of most major groups
of fossil and living echinoderms. Cladistic analyses of
extant and fossil groups helped to reshape some major
classifications, and Mooi and David's (2000 et. al.)
extraxial-axial theory (EAT) provided a new framework
for study of interrelationships. Within the past couple of
decades, molecular analyses are offering powerful tools,
especially in combination with morphology, with which
to address long-standing problems. Thus, the past 40
years have witnessed upheavals in classification at the
family, order, and even class level. In the past decade
numerous important volumes have been published on
the echinoderms, living and fossil. Some of the most
12
comprehensive include: Candia Carnevali & Bonasoro
(1999), Mooi & Telford (1998), Barker (2001), Jangoux &
Lawrence (2001), Feral & David (2001), Kasyanov (2001),
Heinzeller & Nebelsick (2004), Matranga (2005). Despite
the increasing attention received by to this phylum,
there remain many major uncertainties and unresolved
problems.
The extant Deuterostomia ("second mouth") are
usually defined as animals in which the mouth develops
from a second opening in the embryo, opposite to the
initial opening, the blastopore, of the rudimentary gut. In
addition the coelom develops by enterocoely, or pouching
from the primitive gut. Smith (2004b) noted that there
was fossil evidence to show that the major deuterostome
groups were established by about 520 million years ago.
Composition of the extant Deuterostomia has changed
in recent years. At present, it is usually regarded as
comprising the phyla Chordata, Hemichordata, and
Echinodermata (Cameron et aZ., 2000), as well as the
recently-defined phylum Xenoturbella (Bourlat et aZ.,
2006). In echinoderms with planktotrophic larval stages,
the deuterostome affinities of the group are evident. Fell
(1948) and others have pointed out that in a significant
percentage of echinoderms the coelom develops from a
splitting in mesoderm and not from pouching, and the
larval mouth becomes the adult mouth. Pawson and Kerr
(2001) reported presence of chitin in one species of sea
cucumber. Chitin is usually unknown in deuterostomes,
but it has been reported from a blenniid fish (Wagner et
aZ., 1993). These various exceptions to the deuterostome
"norm" are believed to be relatively minor, and the
echinoderms are regarded as fully qualified members of
the Deuterostomia.
Five extant classes of echinoderms are universally
recognized:
Asteroidea,
Ophiuroidea,
Echinoidea,
13
Holothuroidea,
and
Crinoidea.
A
sixth
class,
Concentricycloidea, was described 21 years ago (Baker et
aZ., 1986). Smith (1988a), in a phylogenetic analysis offossil
evidence, suggested times of divergence of the five (then)
modern classes at 450-590 million years ago. Smith noted
that the long subsequent history for each class allowed
for introduction of a great deal of "noise" via mutations.
The surge in molecular taxonomy occurred at around this
time. Wada & Satoh (1994) analyzed one species from
each class, and concluded that phylogenetic relationships
among extant classes matched those deduced from the
fossil record. Littlewood (1995) discussed molecular and
morphological data sets, noting that poor phylogenetic
resolution arose from inconsistent morphological data
sets, and inadequate molecular data, based upon a small
representation of taxa. Soon thereafter, Littlewood et
aZ. (1997) combined more comprehensive morphological,
molecular, and stratigraphic data, and arrived at three
competing phylogenetic solutions, of which two appeared
to be the most plausible. Their study demonstrated that
frustrating pitfalls can appear even when data sets are
fairly detailed . Smith (1997) reaffirmed w ha t many previous
authors had noted, that larval morphology of echinoderms
has evolved independently of adult morphology, and that
larval morphology, when taken alone, cannot inform the
phylogeny of adult echinoderms. Janies (2001), providing
additional sequence data, especially for the refractory
ophiuroids and asteroids, concluded that echinoids and
holothuroids "are related", and crinoids are the sister
taxon to the remaining extant echinoderm classes. The
relationship between asteroids and ophiuroids was more
difficult to address, but Janies found strong support for
monophyly of each of these two classes. McEdward &
Miner (2001) comprehensively summarize what is known
about larval development in echinoderms, and construct
phylogenies for the extant classes based upon patterns of
14
development. As more molecular data become available,
and more comprehensive and refined morphological
databases are produced. The various supra-class names
that have been applied to groupings of echinoderms Echinozoa, Asterozoa, Crinozoa, Pelmatozoa, Eleutherozoa
- are occasionally used in the formal sense, but for the
most part they are used as informal and convenient
adjectives in describing life habits, body form, feeding
propensities, and the like.
Since the first concentricycloid was described (Baker
et aZ., 1986) numerous authors have investigated and
discussed the status of these small (usually <1 cm
diameter), discoidal echinoderms with the mouth frame in
the form of a ring, and a ring of tube feet. The only known
habitat is deep-sea wood, either naturally occurring
waterlogged wood, or wood that has been placed at depth
by submersibles for extended periods of time. Baker et
aZ. (1986) referred these extraordinary animals to a new
class Concentricycloidea. Three species are now known:
XyZopZax medusiformis Baker, Rowe and Clark, 1986; X.
turnerae Rowe, Baker and Clark, 1988; X. janetae Mah,
2006. In the 21 years since description of the first species,
numerous authors have voiced their opinions about the
status of these strange echinoderms. Smith (1988a),
Belyaev (1990), Janies & Mooi (1999), Janies (2001), argued
for placement of the concentricycloids within the Class
Asteroidea, perhaps near the caymanostellids. Rowe et aZ.
(1988), Pearse & Pearse (1994), Mooi et aZ. (1998), Baker
(2003) and others have been a little more cautious in their
assignment of the group. In a detailed recent summary,
Mah (2006), having examined the morphological, cladistic,
and molecular evidence, placed the concentricycloids
within the Asteroidea (Figure 2), where they comprise the
mono typic Infraclass Concentricycloidea of the Subclass
Ambuloasteroidea (sensu Blake & Hagdorn, 2003).
15
Crinoidea
The 650 species of extant crinoids (Class Crinoidea,
Extant crinoids), represented by 100 stalked crinoids and
550 feather stars, have received considerable attention
from specialists, partly because of their intrinsic interest,
and partly because these extant animals can throw some
light on the living habits of the fossil crinoids. The living
crinoids are all referred to the Subclass Articulata Zittel,
1879. Most authors regard the feather stars as comprising
a single order, Comatulida A.H. Clark, 1908. Messing
(1997) notes that Simms (1988) reduced the comatulids
to an infraorder, Comatulidia. Rasmussen & SievertsDoreck (1978) recognized seven superfamilies in the order
Comatulida. Further details are provided by Messing
(1997). The extant stalked crinoids are assigned to four
orders: Millericrinida Sieverts-Doreck, 1952; Cyrtocrinida
Sieverts-Doreck, 1952; Bourgueticrinida Sieverts- Doreck,
1952; Isocrinida Sieverts-Doreck, 1952. In a recent review
of the comatulids, Messing (1997) covers classification,
ecology, and taphonomy, among other topics. For the
stalked crinoids Roux (2002) provide keys and checklists
to the genera. Comprehensive and detailed revisions,
such as David (2006) of the stalked crinoid genus
Endoxocrinus are regrettably rare, as are regional studies
such as Messing (2007). The emphasis on morphology is
gradually being complemented by molecular approaches,
some producing astonishing results. Cohen et aZ. (2004)
found that a clade containing the genera CaZedonecrinus,
Gymnocrinus, Ho Zop us, and Proisocrinus, and also
possibly Cyathidium raises many intriguing questions
about the taxonomic placement of these entities. More
than 6,000 species of fossil crinoids are known as extict,
and revisionary work, usually below the order level, is
proceeding apace. The current classification of the class
16
Crinoidea, in the opinion of Ausich (1998) recognizes six
subclasses: Aethocrinea Ausich 1998; Cladida Moore &
Laudon, 1943; Camerata Wachsmuth & Springer, 1885;
Flexibilia Zittel, 1895; Articulata Zittel, 1879; Disparida
Moore & Laudon, 1943. Hess (2003) describe assemblages
of crinoids from the Ordovician to the Tertiary, and
they study form and function, evolutionary history,
classification, among other topics. Ausich (1999) discussed
the origin of crinoids in light of the re-interpretation of
the Cambrian Echmatocrinus as an octocoral. Ausich
suggests an early Ordovician origin for the crinoids from
primitive rhombiferans via paedomorphosis. Ausich &
Kammer (2001), emphasizing mostly fossils, comment on
the status of research on crinoids, and identify areas for
future research.
A useful and comprehensive classification of fossil
echinoderm groups is provided by Simms (1993). Mooi
(2001) discussed and critically assessed publications on
fossil echinoderms for the period 1980-2000. Loven's law
and ray homologies are described in echinoids, ophiuroids,
edrioasteroids, and an ophiocistioid by Hotchkiss (1995). In
a related study with broad implications, Sumrall & Wray
(2007) discuss pentamerous symmetry and its origin in the
30 Cambrian-Ordovician clades of echinoderms. Shu et aZ.
(2004) describe, and Smith (2004b) comments on, what are
believed to be ancestral echinoderms ("velulocystids") from
the Lower Cambrian of China. The vetulocystids have a
globosetheca and a tail. Smith (2004a) studied phylogeny
of deuterostomes, and anatomy of carpoids, and concluded
that early deuterostomes with a stereom skeleton and gill
slits may have existed, but it is not likely that sterom and
a notochord co-occurred. These conclusions support in part
(gill slits), and disagree in part (notochord) with the ideas
of Jefferies (1996). David (1999), applying the extraxiall
axial theory, regard the four non-pentamerous classes
17
comprising the homalozoans as early echinoderms, but not
indicative of the plesiomorphic morphology of the phylum.
Further, the Homalozoa is not a monophyletic assemblage.
Lefebvre (2007) studied in detail the palaeobiogeography
and palaeoecology of cornutes and mitrates. Parsley (1999),
using a cladistic approach, determined that the Cincta
(Homostelea) are blastozoans. Ophiocistioids have been
reviewed by Haude (2004), and Reich & Haude (2004).
Dominguez-Alonso (1999) presented new data on the
structure of ctenocystoids and proposed a new approach to
the early evolution of echinoderms. Molecular analyses are
transforming our view of the echinoderms at all taxonomic
levels. Wilson (2007) found that the Antarctic comatulid
previously known as the single widely distributed species
Promachocrinus kerguelensis may in fact embrace at least
five species-level clades.
Plancus and Gaultire (1743) first reported the
presence of echinoderms from Goa, coastal area of India
(James, 1987). Bell (1887) compiled the list of echinoderms
from Bay of Bengal. Nagabhushanam and Rao reported
the echinoderms of Orissa and Lakshadweep in 1969 and
1972. Soota (1983) gave distribution of some holothurians
in Andaman and Nicobar Islands without mentioning
the species wise distribution. Venkataraman (2002) also
compiled a report on echinoderms. Sastry (2005, 2007)
reported an annotated checklist on echinodermata of
Andaman and Nicobar Islands with 424 species and in
India with 649 species. The present study dealt with
pictorial description of 89 species of most commonly found
echinoderms belonging to 53 genera, 31 families and 15
orders recorded from Andaman and Nicobar Islands.
Asterioidea
Today's oceans are graced by 2100 species of sea
stars. In their magisterial monograph of Atlantic sea stars
18
(Class Asteroidea, Extant asteroids), Clark and Downey
(1992) noted that the status of the entire group-whether
it should be referred to a Class or a Subclass-was
'controversial" and they declined to discuss the topic. At
the ordinal level, they elected to follow the classification
proposed by Blake (1987, 1989), and recognized seven
orders: Paxillosida Perrier, 1884; Notomyotida Ludwig,
1910; Valvatida Perrier, 1884; Velatida Perrier, 1893;
Spinulosida Perrier, 1893; Forcipulatida Perrier, 1893;
Brisingida Fisher, 1928. Clark and Downey chose to
ignore, or did not accept, the three Superorders proposed
by Blake (1987). Gale (1987) proposed a new Subclass,
Neoasteroidea for the post-Paleozoic crow group asteroids,
and he differed from Blake in naming just four orders:
Paxillosida, Notomyotida, Valva tid a (including the
velatids and spinulosids), and Forcipulatida (including
the brisingids).
Subsequent to the Blake and Gale papers, Lafay et aZ.
(1995) made a combined morphological and molecular study
of nine sea star species, in an attempt to provide a broader
framework for discussion of the differing classifications.
Papers published following a symposium on evolution of
starfishes (Blake, 2000; Blake et aZ., 2000; Mooi & David,
2000; Hotchkiss, 2000; Vickery & McClintock, 2000;
Hrincevich et aZ., 2000; Mah, 2000; Knott & Wray, 2000)
offered possibilities for resolution of some differences in
morphology-based phylogenies by the addition of more
comprehensive molecular data. The consensus of opinion
seemed to be that molecular/morphological studies are
very necessary and desirable, especially for resolving
problems at the family-level and below (Hrincevich et
aZ., 2000; Knott & Wray, 2000; O'Loughlin & Waters,
2004; Mah, 2007). Matsubara et aZ. (2005) found that
comparisons of genome structure were "uninformative for
the purposes of asteroid phylogeny, but that phylogenetic
19
analyses based upon nucleotide and amino acid sequences
indicated that paxillosidan characters are secondarily
derived, and perhaps the paxillosids are specialized
rather than primitive asteroids". In a comprehensive
paper describing the Ordovician to recent history of the
Asteroidea, Blake & Hagdorn (2003) diagnosed a new
Subclass Ambuloasteroidea, to reflect their findings and
those of Mooi & David (2000) on their extraxial/axial
theory; in regard to the axial skeleton of Paleozoic and
post- Paleozoic asteroids. Blake & Elliott (2003) found that
axial skeletal characters could be compared across stem
and crown-groups of asteroids. The Ambuloasteroidea
embrace two infraclasses, one "Unknown" which includes
the late Paleozoic Families Calliasterellidae Schondorf
1910, and Compsasteridae Schuchert 1914, the other
Gale's (1987) Infraclass (formerly Subclass) Neoasteroidea.
The Neoasteroidea embraces the remammg postPaleozoic asteroids. Shackleton (2005) comprehensively
reviewed Ordovician stem-group asterozoans (asteroids
plus ophiuroids) and proposed a revised classification.
A new Plesion (Order) Eopentaroida was proposed to
accommodate three distinctive Silurian genera. It was
noted that the asteroid and ophiuroid body plans, already
established in the Ordovician, indicated that these groups
had a considerable, but unknown, pre-Ordovician history.
In this context, Herringshaw et aZ. (2007) established
that the Silurian Lepidaster grayi was the earliest-known
multiradiate sea star, so that the multiradiate body plan
was also established early. Blake & Hotchkiss (2004)
characterized the post-Paleozoic asteroid crown group,
identifying apomorphies of crown-group diversification.
Ophiuroidea
The 2,000 species of ophiuroids (Class Ophiuroidea,
Extant ophiuroids) are traditionally assigned to two
20
orders such as Order Euryalida Lamarck, 1816 and Order
Ophiurida Muller & Troschel, 1840. Smith, Paterson
& Lafay (1995) summarized the state of knowledge of
ophiuroid classification after a cladistic analysis based
upon morphological characters. Their molecular data
were equivocal, and did not help to resolve some problems
arising from the morphological analyses. Smith & Paterson
assigned the mysterious Ophiocanops fugiens to Subclass
Oegophiuridea Matsumoto, 1915, and referred all other
ophiuroids to Subclass Ophiuridea Gray, 1840. However,
Pearse et aZ. (1998), with access to new material of
Ophiocanops, demonstrated that this genus belonged in the
Family Ophiomyxidae. These findings, coupled with those
of Hotchkiss (1977) have resulted in the disappearance of
the Oegophiuridea as a subclass of ophiuroids. Cisternas
et aZ. (2004) studied development patterns in 23 species,
suggested some evolutionary pathways, and noted that
good molecular phylogenies would help to refine their
hypotheses. The puzzling Ordovician stelleroid Stenaster
was studied in detail by Dean (1999), who concluded, after
a cladistic analysis of early asteroids, soma steroids, and
ophiuroids, that this asteroid-like animal was in fact an
ophiuroid, converging secondarily with asteroids. Kroh
(2004) reported on the first fossil record of a euryalid
ophiuroid from the Miocene of the Mediterranean. Jagt
(1999) found a rich fauna of ophiuroids in the Cretaceous
of the Maastrichtian Stage. Hotchkiss (1993) has described
several Ordovician and Devonian ophiuroids from North
America, and commented extensively on origins and
relationships of skeletal features.
Echinoidea
The 800 species of echinoids (Class Echinoidea, Extant
and fossiZ echinoids) alive today, coupled with the wonderful
21
fossil record, make the echinoids excellent candidates for
a great variety of studies, from reproductive biology to
evolution. Echinoids are also of commercial importance;
Yokota et aZ. (2002) cover biology and aquaculture of sea
urchins, and Lawrence (2001, 2006) deals with biology,
ecology, and aquaculture of edible sea urchins worldwide.
Extant echinoids are informally divided into "regular"
forms with obvious radial symmetry, spherical bodies and
uniformly long spines, and "irregular" forms - bilaterally
symmetrical urchins, including sand dollars and heart
urchins. Echinoids are typically classified into two
subclasses and 12 orders, as listed by Smith (1984), and
modified slightly by Littlewood & Smith (1995). Subclass
Cidaroidea Claus, 1880 comprises the Order Cidaroida
Claus, 1880. Subclass Euechinoidea Bronn, 1860 includes
all other echinoids: Order Echinothurioida Claus, 1880;
Order Diadematoida Duncan, 1889; Order Pedinoida
Mortensen, 1939; Order Calycina Gregory, 1900; Order
Arbacioida Gregory, 1900; Order Phymosomatoida
Mortensen, 1904; Order Temnopleuroida Mortensen,
1941; Order Echinoida Claus, 1876; Order Cassiduloida
Claus, 1880; Order Clypeasteroida Agassiz, 1872; Order
Spatangoida Claus, 1876. All of the above orders except
the last three are regular echinoids; the last three comprise
the irregular echinoids. Phylogenetic relationships of the
echinoid higher taxa have been studied by several authors,
using morphological data (test and spine morphology,
structure of teeth, pedicellariae, and structure of
larvae) and molecular sequences. Some of the more
comprehensive studies include Smith (1988b), Smith et aZ.
(1992), Littlewood & Smith (1995), Lee (2003), Stockley
et aZ. (2005), Smith et aZ. (2006) and Smith (2007). Most
of the regular echinoids seem to have diverged relatively
recently, between 65 and 35 million years ago. Solovjev
22
& Markov (2004) studied earliest divergence of irregular
echinoids, and selected 15 binary characters for cladistic
analysis. They agreed in general with the conclusions
of Smith (1984) and Rose & Olver (1988) but differed in
some details.
Holoth uroidea
The approximately 1400 species of holothurians
(Class Holothuroidea, Extant holothuroids), in particular
the tropical shallow-water forms, are receiving increasing
attention from specialists, and substantial changes in
classification below the family level are expected to
occur over the next decade. Above the family level, the
classification has been fairly stable for many years. The
orders Aspidochirotida Grube, 1840; Elasipodida Theel,
1882; Molpadiida Haeckel, 1896; and Apodida Brandt,
1835 remain well-characterized. The distinction between
the Dendrochirotida Grube, 1840 and Dactylochirotida
Pawson & Fell, 1965 is less well-defined. At some points
the alleged morphological differences between these two
groups are less well defined, but at the molecular level,
the distinctions are more robust (Kerr & Kim, 2001). The
three subclasses introduced by Pawson & Fell (1965)Apodacea, Aspidochirotacea, and Dendrochirotaceaas a convenient grouping of the orders have essentially
been abandoned over the years, and Kerr & Kim (2001)
convincingly demonstrate their impracticality. The
interrelationships between extant holothurians at the
family level and above were rather neglected until the
1990's, when a few authors used morphological and
molecular approaches to investigate relationships. The
publications by Kerr & Kim (1999, 2001) and Kerr (2001)
were pioneering efforts, and they surely stimulated the
surge in popularity of holothurians as research subjects
23
at all levels of classification. Lacey (2005) found cause to
disagree in some areas with Kerr & Kim's conclusions,
but all of these authors agreed with several earlier
authors in suggesting a basal position for the Apodida.
Smirnov (1998) revised the classification of the Order
Apodida, using morphological characters. Phylogenetic
studies of family-level and lower groups are burgeoning
(Kerr, 2005). The fossil record of extinct holothurians
has long been regarded as poor, an opinion disputed by
Kerr & Kim (1999). Since the attempts by Pawson (1966,
1980) to integrate the higher-level classifications of fossil
and recent holothuroids, the study of fossil groups has
burgeoned, largely under the leadership of Gilliland (1992,
1993) and Reich (2002, 2004). Gilliland (1993) provides a
comprehensive summary of the state of knowledge of fossil
holothurians, and identifies areas where more research is
required. In several publications, Reich reviews European
and other fossil holothurians. Gilliland and Reich, and
Smirnov (1999), suggest divergence times for the major
holothurian taxa, and their conclusions are discussed by
Kerr & Kim (1999, 2001), and others.
3. CHARACTERISTICS OF EXTANT CLASSES
Crinoidea
Crinoids are the least understood ofliving echinoderms,
their skeletal remains are among the most abundant and
important of fossils. They appeared during the Lower
Ordovician and underwent several major radiations
during the Paleozoic Era. Crinoids were major carbonate
producing organisms during the Paleozoic and Mesozoic.
In many Paleozoic and Mesozoic settings entire carbonate
shelves were composed predominantly of crinoidal remains
24
(Ausich 1997). A persistent, traditional view treats living
crinoids as chiefly deep-sea organisms, relicts of their
opulent Paleozoic past, holding off final extinction in
remote abyssal habitats. This view is generally applied to
stalked crinoids, or sea lilies, as typical of the entire group,
because they most closely resemble their fossil forebears. It
is true that the approximately 80 extant species of stalked
crinoids are chiefly restricted to depths greater than 200m
(the shallowest occurs in 100m). However, 85% of extant
crinoids (approximately 540 named species) are unstalked
feather stars, or comatulids, the products of a continuing
post-Paleozoic radiation (Meyer & Macurda 1977). About
65% of living comatulids occur at shelf depths «200 m). All
crinoids are passive suspension feeders. They produce no
feeding/respiratory current but, rather, rely on extrinsic,
ambient water movement. In extant crinoids, the foodgathering apparatus functions as follows: each featherlike
arm that radiates from the central body bears an open
ambulacral groove bordered by triads of fingerlike podia,
or tube feet, which are terminal extensions of the water
vascular system. The longest tube foot in each triad, 0.430.85 mm in length, is held out at a right angle and flicks
passing food particles into the groove. After a food particle
is captured by a crinoid, the shortest tube foot wraps it in
mucous secretions; ciliary tracts on the groove floor then
transport it toward the mouth. In living crinoids, food
particle size ranges from about 50 to 400 /lm. Diets include
a variety of protists (e.g., diatoms and other unicellular
algae, foraminiferans, actinopods), invertebrate larvae,
small crustaceans, and detrital particles.
Crinoids are pentamerous, stalked echinoderms
with a cuplike body bearing five usually branched and
commonly featherlike arms. Most of a crinoid's body
25
consists of an endoskeleton composed of numerous
calcareous pieces, called plates or ossicles. The visceral
mass of the crinoid animal is encased in the aboral cup
that is typically composed of 2-3 circlets of plates. The
mouth and anus are on the upper or oral surface of the
animal. Additional circlets of fixed arm plates and fixed
interradial plates may occur above the aboral cup, making
a larger calyx. Five radial plates (the uppermost circlet
of aboral cup plates) are aligned with the radial water
vascular canals and give rise to five arms on the oral side
of the body. Each arm is an articulated series of ossicles
extending outward from the body. Arms contain extensions
of coelomic, nervous, water vascular, and reproductive
systems and bear an ambulacral groove bordered by
fingerlike tube feet, or podia. Arms may be non-branching
or branch in many different ways. All living crinoids are
pinnulate, that is, they bear a small side branch (pinnule)
on alternating sides of successive ossicles along the arm.
In living crinoids, the pinnules bear the food-gathering
tube feet. The crinoid stalk typically consists of numerous
discoidal skeletal pieces called columnals, held together
by ligaments and penetrated by a central canal containing
coelomic and neural tissue. In most species, the stalk
serves to anchor the animal permanently to the substrate
via one of a variety of terminal structures, e.g., a discoidal
or encrusting holdfast, rootlike radix, or grapnel. In
others, such as the living isocrinids, whorls of hooklike
cirri along the stalk allow the crinoid to release its hold
and crawl with its arms. Several crinoid groups, notably
the comatulids, which include the only living shallowwater crinoids, have lost the stalk. Comatulids anchor
via numerous cirri that arise from the retained topmost
columnal (the centrodorsal).
26
Asteroidea
The Asteroidea IS one of the largest and most
familiar classes within the Phylum Echinodermata. These
animals, commonly known as sea stars or starfishes, form
a diverse and speciose group. There are approximately
1600 extant species (Hyman 1955; Clark 1977; Clark and
Downey 1992) which are found throughout the world's
oceans. Like other echinoderms, asteroids are important
members of many marine benthic communities. They
can be voracious predators, having significant impacts
on community structure. Paine (1966) used Pisaster
ochraceus to illustrate his concept of the role keystone
species play in community ecology. The crown-of-thorns
starfish, Acanthaster pZanci, is particularly well-known
because it can cause extreme detrimental effects to coral
reefs, particularly during population outbreaks (Moran
1988).
The controversial Concentricycloidea (a proposed sixth
class of the Echinodermata; Baker et aZ. 1986, Rowe et aZ.
1988, Pearse and Pearse 1994) have been diagnosed as
unusual asteroids (Smith 1988, Belyaev 1990, Janies and
Mooi 1999). Their relationship to other asteroid taxa is not
well resolved, but alliances with species from the Vela tid a
and the Forcipulatida have been proposed. The unique
morphology of the concentricycloids makes it difficult to
assign this group to the recognized asteroid orders and is
cited as sufficient distinction for class recognition.
Like other asterozoans, asteroids have a characteristic
star-shaped body plan consisting of a central disc and
multiple (typically 5) radiating arms. Asteroids are
most easily distinguished from other asterozoans (the
Ophiuroidea) by the structure of the arms. In asteroids,
skeletal support for the arms is provided by the ossicles
27
of the body wall, which merge with those of the central
disc, giving the arm a very broad based attachment
to the disc. This skeletal arrangement allows for the
extension of a comparatively large coelomic cavity from
the central disc into the arms, which serves to hold some
of the animal's organ systems, namely the gonads and
pyloric caeca. Additionally, this skeletal arrangement
also limits lateral flexion of the arms. Locomotion
by asteroids is accomplished almost exclusively by
means of the podia of the water vascular system.
Taxonomy of asteroids usually is based on externally
observable characteristics of the skeleton, particularly
the primary ossicular series which define the body wall
(ambulacrals, adambulacrals, marginals, terminals,
actinals, abactinals), as well as secondary ossicles such
as spines, spinelets and pedicellariae. Perhaps the most
important ossicular series defining the Asteroidea is the
ambulacral column, found along the oral surface of the
disk and radiating arms and associated with two or four
rows of podia. The asteroid ambulacrum is distinguished
by erect ambulacral ossicles arranged in series along the
length of the ambulacral column. According to the EAT,
the ambulacral and terminal ossicles of asteroids are
axial elements. These ossicles are formed according to
the Ocular Plate Rule (OPR) and are associated with the
developing water vascular system during ontogeny as are
the axial ossicles of other echinoderms. The remaining
asteroid ossicle series are extraxial elements, which can
be added during ontogeny without any particular ordering
system (although secondarily ordered serial homologous
elements are common in the asteroids, e.g. adambulacrals
and marginals). In comparison to axial elements, extraxial
ossicles are prone to much more evolutionary lability
(Mooi and David 1997).
28
Ophiuroidea
Ophiuroids are a large group (over 1600 species) of
echinoderms that includes the brittle stars (Ophiurida) and
basket stars (Euryalida). The more familiar Ophiurida,
or brittle stars, usually have five arms and superficially
resemble true starfish (Asteroidea). However, brittle stars
have long, flexible arms (hence the other common name
for ophiuroids, "snake stars" and a central, armored, diskshaped body that is clearly demarcated from the arms.
Instead of crawling on hundreds of tube feet like starfish,
brittle stars move fairly rapidly by wriggling their arms.
These fragile arms are supported by an internal skeleton
of calcium carbonate plates that superficially look like
vertebrae, and that are in fact called vertebral ossicles.
These are moved by a system of muscles and linked
together by ball-and-socket joints. The body and arms are
also protected by calcium carbonate plates, and the arms
generally bear delicate spines. Basket stars (Euryalida)
have a similar structure to brittle stars, although they
are usually larger. However, their arms are very highly
forked and branched, and even more flexible than those
of brittle stars.
Ophiuroids can be found in most parts of the world,
from the Arctic and Antarctic to the tropics. Ophiuroids
are common in many shallow-water marine habitats, and
include a few species which can adapt to brackish water,
which is quite unusual for echinoderms. Ophiuroids are
dominant in many parts of the deep sea, where in certain
regions the bottom may swarm with brittle starts. Basket
stars also tend to live in deeper water. Most ophiuroids
are scavengers and detritus feeders, although they also
prey on small live animals such as small crustaceans and
worms. Some, in particular the basket stars, filter-feed on
plankton with their arms.
29
Echinoidea
Echinoids are one of the more diverse and successful
echinoderm groups today, including familiar echinoderms
such as the sea urchins and sand dollars. The roe (egg
mass) of some species, notably certain sea urchins, is
eaten in some cultures, notably in Japanese sushi; as a
result, certain echinoid species are commercially fished.
The larval development of echinoids has also been studied
extensively, and many discoveries in developmental
biology have been made using echinoids. In echinoids, the
skeleton is almost always made up of tightly interlocking
plates that form a rigid structure or test in contrast with
the more flexible skeletal arrangements of starfish, brittle
stars, and sea cucumbers. Test shapes range from nearly
globular, as in some sea urchins, to highly flattened, as
in sand dollars. Living echinoids are covered with spines,
which are movable and anchored in sockets in the test.
These spines may be long and prominent, as in typical
sea urchins. In sand dollars and heart urchins, however,
the spines are very short and form an almost belt-like
covering. The mouth of most echinoids is provided with
five hard teeth arranged in a circlet, forming an apparatus
known as Aristotle's lantern.
Echinoids are classified by the symmetry of the test,
the number and arrangement of plate rows making up
the test, and the number and arrangement of respiratory
pore rows called petals Traditionally, echinoids have
been divided into two subgroups: regular echinoids,
with nearly perfect pentameral (five-part) symmetry;
and irregular echinoids with altered symmetry. Regular
echinoids include the Cidaroida (pencil urchins) and
Echinoida (sea urchins, including the long-spined sea
urchin). The Clypeasteroida (sand dollars and sea biscuits,
30
above center), the Spatangoida (heart urchins), and the
Cassiduloida, a somewhat sand-dollar-like group whose
members are rare today, make up the irregular echinoids.
Holoth uroidea
The Holothuroidea, or sea cucumbers, are an abundant
and diverse group of worm-like and usually soft-bodied
echinoderms. They are found in nearly every marine
environment, but are most diverse on tropical shallowwater coral reefs. They range from the intertidal, where
they may be exposed briefly at low tide, to the floor of the
deepest oceanic trenches. Considerable diversification has
occurred since then with about 1400 living species in a
variety of forms. Some of these are about 20 cm in length,
though adults of some diminutive species may not exceed
a centimeter, while one large species can reach lengths
of 5 m (Synapta maculata). Several species can swim
and there are even forms that live their entire lives as
plankton, floating with the ocean currents.
The most important feature distinguishing the sea
cucumbers is a calcareous ring that encircles the pharynx
or throat. This ring serves as an attachment point for
muscles operating the oral tentacles and for the anterior
ends of other muscles that contract the body longitudinally.
Sea cucumbers are also distinct as echinoderms in having a
circlet of oral tentacles. These may be simple, digitate (with
finger-like projections), pinnate (feather-like), or peltate
(flattened and shield-like). A third key feature, found in
90% of living species, is the reduction of the skeleton to
microscopic ossicles. In some species, the ossicles may
be enlarged and plate-like. As in other echinoderms, the
holothurian water vascular system consists of an anterior
ring canal from which arise long canals running posterior.
31
Despite their similarity to the radial canals of other
echinoderms, these latter structures arise embryologically
in a quite different Mannar. For this reason these canals
in holothurians have been recently renamed longitudinal
canals (Mooi and David 1997). In holothurians, the larval
structures that would form the radial canals in other
echinoderms instead become the five primary tentacles.
Also, holothurians with the exception of members in
Elasipodida have a madrepore that opens into the coelom
(body cavity). In contrast, elasipodans and nearly all other
echinoderms have a madrepore that opens externally.
Some sea cucumbers possess organs not found in other
invertebrates. In some Aspidochirotida, the respiratory
trees display Cuvierian tubules. In most species, these
are apparently defensive structures. They can be expelled
through the anus, whereupon they dramatically expand
in length and become sticky, entangling or deterring
would-be predators, such as crabs and gastropods. Many
forms, with the exception of members of Elasipodida and
Apodida, possess respiratory trees used in gas exchange.
These are paired, heavily branched tubes attached to the
intestine near the anus. This type of breathing ("cloacal
breathing") is also present in an unrelated group, the
echiuran worms.
4. STUDY AREAS
The Andaman and Nicobar group of Islands is located
the southeast of Bay of Bengal, between 6° -14 ° N
Latitude and 91°-94° E Longitude. They are the part of the
mountain chain and lie on a ridge that extends southward
from Irrawaddy delta of Burma, containing the trend of the
Arakan Yoma range (Venkataraman et aZ., 2003). There
are 572 islands in the chain, some of which are volcanic.
III
32
The islands occupy an
area of 8293 km 2 with
a coastline of 1962 km
and account for 30% of
the Indian Exclusive
Zone. There are 106
protected areas in these
islands, 96 designated
as wildlife sanctuaries,
9 national parks and
one biosphere reserve.
Among
9
national
parks, 2 are marIne
national parks which
have not yet inventoried
thoroughly. The coral
reefs of Andaman and
Nicobar Islands are the
biodiversity hot spot of
India
(Jeyabaskaran,
1999).
. .......
セ@
Ten llegIee
_..
.....
The names of the study areas are cited below with
Global Positioning System (GPS) coordinates.
S1.
Area surveyed
No.
GPS Coordinates
Latitude
I
Longitude
South Andaman
1.
Off Burmanella
11 ° 33.468'N
92° 43.873'E
2.
Off Rangachang
11° 34.350'N
92° 44.133'E
3.
Chidyatapu
11 ° 29.460'N
92° 42.530'E
4.
Pongibalu
11° 31.030'N
92° 39.159'E
33
GP8 Coordinates
81.
Area surveyed
No.
Latitude
I
Longitude
5.
North Bay
11 ° 42.068'N
92° 45.116'E
6.
Off Collinpur
11 ° 41.598'N
92° 37.035'E
7.
Off Kurmadera
11° 39.933'N
92° 35.903'E
Rutland Island
8.
Chain Nalah
12° 08.522'N
93° 06.551'E
9.
Padauk Dikri
12° 29.288'N
92° 40.141'E
10.
Surumai Dikri
11 ° 25.504'N
92° 40.301'E
11.
Komeo
11° 24.314'N
92° 39.780'E
12.
Mitta Nalah
11° 28.541'N
92° 40.371'E
13.
Arom Point
11° 30.541'N
92° 38.769'E
14.
Aam Dera
11 ° 24.664'N
92° 37.456'E
Mahatma Gandhi Marine National Park
15.
North Wando or
11° 37.270'N
92° 37.035'E
16.
Grub Island
11 ° 35.391'N
92° 35.637'E
17.
J oIly Buoy Island
11° 30.251'N
92° 32.591'E
18.
Tarmugli Island
11° 33.261'N
92° 36.809'E
Ritchie's Archipelago
34
19.
Ha velock Island
12° 00.005'N
92° 56.808'E
20.
Inglis Island
12° 08.639'N
93° 06.786'E
21.
Henry Lawrence
12° 05.000'N
Island
93° 06.312'E
81.
Area surveyed
No.
GP8 Coordinates
Latitude
I
Longitude
22.
John
Lawrence
12° 04.075'N
Island
93° 00.398'E
23.
Outram Island
12° 00.574'N
92° 56.808'E
24.
Sir William Peel
12° 03.315'N
Island
92° 59.929'E
25.
Nicolson Island
12 ° 06.739'N
92 ° 57.235'E
26.
South
Island
Button
12° 13.467'N
92° 01.334'E
27.
North
Island
Button
12° 18.974'N
92° 03.826'E
28.
Middle
Island
Button
12° 16.4 73'N
93° 01.334'E
29.
Wilson Island
12° 13.061'N
93° 15.207'E
Neill Island
30.
Lakshmanpur
11 ° 50.826'N
93° 00.554'E
31.
Hawrah Bridge
11 ° 49.727'N
93° 00.818'E
32.
Middle Point
11° 50.857'N
93° 00.554'E
33.
Ramangar
11 ° 48.400'N
93° 01.440'E
34.
Sunset point
11° 51.941'N
93° 00.667'E
35.
Little Neil Island
11° 47.063'N
93° 04.616'E
36.
Pearl Park Beach 11° 50.766'N
93° 00.795'E
Little Andaman Island
37.
Butler Bay
10° 40.232'N
92° 56.808'E
35
81.
Area surveyed
No.
GP8 Coordinates
Latitude
I
Longitude
38.
Kala Pathar
10° 39.558'N
92° 34.109'E
39.
Haminder Bay
10° 32.975'N
92° 32.651'E
40.
Off
Ramkrishnapur
10° 42.630'N
92° 33.066'E
41.
Sister Island
10° 55.830'N
92° 07.023'E
42.
Hut Bay
10° 35.419'N
92° 33.066'E
43.
Dugong Creek
10° 48.385'N
92° 64.000'E
44.
Off Light House
10° 30.734'N
92° 30.264'E
Middle Andaman
45.
North Reef
Island
12° 56.084'N
92° 57.345'E
46.
Interview Island
12° 59.125'N
92° 42.981'E
47.
Avis Island
12° 56.210'N
92° 33.066'E
48.
Sound Island
12° 56.084'N
92° 57.345'E
49.
Rail Island
12° 56.860'N
92° 54.620'E
50.
Karlo Island
12° 56.084'N
92° 53.378'E
51.
Karmatang
12° 51.322'N
92° 56.050'E
North Andaman
36
52.
Ross Island
13° 18. 167'N
93° 04.261'E
53.
Smith Island
13° 18. 406'N
93° 04.207'E
54.
Ariel Bay
13° 16. 093'N
93° 02.433'E
GP8 Coordinates
81.
Area surveyed
No.
Latitude
Lamia Bay
13° 24. 879'N
93° 05.516'E
55.
I
Longitude
Nicobar Islands
Car Nicobar Island
56.
Malacca
09° 10.490'N
92° 49.714'E
57.
Kakaana
09° 07.750'N
92° 48.678'E
58.
Tamoloo
09° 1l.350'N
92° 49.498'E
59.
Kimos
09° 07.587'N
92° 46.316'E
60.
Perka
09° 1l.203'N
92° 49.877'E
61.
Lapati
09° 13.978'N
92° 48.002'E
62.
Kamorta Island 12° 51.322'N
Bada Enaka
92° 56.050'E
63.
Champin Island
08° 01.670'N
93° 33.123'E
64.
Trinket Island
08° 02.806'N
93° 34.556'E
65.
Kamorta Island Kardip
08° 02.151'N
93° 33.182'E
66.
Kamorta Island Kakkana
08° 07.170'N
93° 31.606'E
67.
M unak Island
07° 59.813'N
93° 30.534'E
68.
Katchal Island
07° 58.952'N
93° 24.351'E
69.
Teressa Island
08° 13.686'N
93° 10.913'E
70.
Kundol Island
07° 10.023'N
93° 42.949'E
N ancowry Islands
37
81.
Area surveyed
No.
GP8 Coordinates
Latitude
I
Longitude
Great Nicobar Island
71.
Campbell Bay
06° 59.749'N
93° 56.718'E
72.
Off Laxman
Beach
07° 01.482'N
92° 37.456'E
Off Gandhi
Nagar
06° 50.496'N
73.
74.
Joginder Nagar
06° 57.226'N
93° 55.495'E
75.
Singam Basti
06° 58.307'N
93° 55.7 48'E
76.
Navy Dera
07° 07.571'N
93° 53.133'E
77.
Indira Point
06° 45.428'N
93° 49.541'E
78.
Kopen Heat
06° 50.923'N
93° 47.983'E
93° 53.680'E
5. MATERIAL AND METHODS
Several surveys were conducted during the period of
April, 2009 to July, 2010 at different sites of Andaman
and Nicobar Islands to assess the diversity of echinoderms
in coral reef enviornments by employing Self Contained
Underwater Breathing Apparatus (SCUBA) diving
and snorkeling. Line intercept transect (Bradbury and
Reichelt et.al., 1986), Quadrate methods (Endean and
Stablum, 1973), Photoquadrate and underwater video
transect method were applied to investigate the diversity
and distribution of the Echinoderms. Underwater video
sampling provides highly precise quantitative estimate
of echinoderms and abundance of common benthic taxa.
During SCUBA diving, species recording was made by
38
underwater digital photography (Sony - Cyber shot,
Model-T900, marine pack, 12.1 megapixels) for detailed
identification.
6. KEY CHARACTERS
Class: CRINOIDEA
Proximal pinnules very flexible and with some of the
terminal segments modified to form a comb; mouth near
the edge of the disc and anal tube approximately central
........................................................ Family: Comasteridae
The two ossicles of the IBr series and the first two
after most, if not all the axillaries united by syzygy,
Division series often alternating with arms ...................... .
..... ....... ...... ...... ...... ...... ...... ...... ....... ...... ..... Genus: Comaster
No dorsal process on the basal segments ofthe proximal
pinnules; if cirri are present at all then their distal
segments are smooth dorsally ............ Comaster schlgeli
Arms well over 100 in number (when fully grown) ....
............................................... Comaster multibrachiatus
The most external IIIbr series of each ray usually
two, the internal ones four; arms up to 200 ...................... ..
............................................................ Genus: Comanthina
Whole body black in color, central disc and proximal
portions of arms with black spots; some distal parts of
arms also with black spots but fewer; tips of pinnules
yellow ................................................ Comanthina nobilis
III Br series, when present, either all four or else four
and two irregularly; up to 120 arms but in most species
less than 60 ......................................... Genus: Comanthus
39
Cirri very reduced in size, often discontinuously
arranged around the edge of the centrodorsal, their distal
segments only slightly shorter than the proximal ones ....
..................................................... Comanthus parvicirrus
More than 60 arms; no dorsal processes on the distal
cirrus segments ......................... Oxycomanthus bennetti
Middle and distal cirrus segments with a pair of
dorsal spines or tubercles, one each side of the mid-line,
rarely a transverse rigid .......... Family: Colobometridae
Arms fewer <40, rarely >45 cirrus segments, 53-120
arms; division series all two; cirri with 41-80 segments;
all the pinnule present .............. Pontiometra andersoni
Outer pinnules (2) very stout, stiff and erect or even
recurved over the disc, its segments with conscipicously
flared and spinose distal ends, Arms: more than 10 .........
..... ....... ...... ...... ...... ...... ...... ...... ....... ........ Genus: Cenometra
Basal segments of none of the proximal pinnules
carina tes, A simple keel on the bases of the proximal pinnules
or else the edge rounded ........................ Cenometra bella
Basal segments of the proximal pinnules with knoblike rounded process forming a crest along the edge facing
the tip of the arm ........................ Cenometra emendatrix
Outer pinnules (2) markedly prismatic, often with the
angles conscipicously produced at the distal ends of the
segments ..................................... Oligometra serrapinna
All the pinnules prismatic; the distal pinnules, if not
all of them, flexible and not conscipicously stiffened .........
................................................... Family: Himerometridae
The first three external brachial pinn ules (P l' P 2'
P 3)' progressively diminishing in size, if any pinnules are
present on the division series they are largest of all .........
........................................................... Genus: Himerometra
40
Proximal pinnule up to 25, usually 20, segments,
tapering fairly abruptly near the tip, not flagellate,
Enlarged proximal pinnules with smooth segments
although their distal ends may be flared; proximal
brachials with smooth but hardly at all flared distal
edges .................................... Himerometra robustipinna
Rarely more than 10 arms but should this number
be exceeded than the IlBr series are two; the proximal
pinnules little different from the following ones .............. ..
............................................................ Genus: Amphimetra
Cirri with 30-35 segments and outer pinnules with
18-21 ................................................ Amphimetra molleri
Usually more than 10 arms with the IlBr series mostly
four but ifonly 10 arms are present then the proximal pinnules
are distinctly modified with a strong crest or the segments
have flared or spinose distal ends ....... Genus: Heterometra
Distal cirrus segments with distinct dorsal spines or
tubercles, Proximal pinnules appearing serrated in profile,
due to projections from the distal ends of the segments ...
...... ...... ...... ...... ...... ...... ....... ...... .... Heterometra crenulata
Arms often exceeding 20 in number than the IllBr series
usually with four ossicles .. ......... Heterometra philiberti
Always more than 10 arms ......................................... ..
...................................................... Family: Mariametridae
Second interradial arm is the largest proximal
pinnule .......................................... Genus: Lamprometra
Second inter radial arm either very stout or distinctly
carina te basally .......................... .Lamprometra palmata
Class: ASTEROIDEA
Skeleton of dorsal surface paxilliform or otherwise
tube feet with suckers ............................ Order: Valvatida
41
The quadrangular or crescentic abactinal plates more
or less obviously imbricating in the proximal direction
usually armed with fine spineletsor granules, rarely naked
except around the anus and madreporite, arms flattened
or rounded above, occasionally somewhat carinate but
then with no mid-radial series of conspicuous spines .......
............................................................ Family: Asterinidae
Body usually stellate but if nearly pentagonal then
the margin is thin and pliable, often curling upwards in
preserved specimens. Radius from centre to arm exceeding
20mm, Abactinal armament usually distinctly spiniform
and sometimes fine .................................. Genus: Asterina
Abactinal armament very delicate, consisting of fine
minute hyaline spinelets, easily rubbed off, the petaloid
radial popular areas linked only to the area in the centre of the
disc, body markedly flattened .............. Asterina sarasini
Marginal plates large, forming a conscipicous side-wall
to the body, the upper surface almostflat, rarely somewhat
convex; no papillae on the lower side; interradial arcs
rounded .......................................... Family: Goniasteridae
Radial areas, if not the whole upper side, distinctly
convex; supero-marginals narrower; arms slender;
granules around the papulae usually similar to the rest..
................................................................. Genus: Stellaster
Not more than one infero-marginal spine, sometimes
none .................................................... Stell aster equestris
Three series of spines on the ambulacral plates; arms
usually triangular or tapering to a blunt tip; carina I rows of
tubercles not conscipicously enlarged ...... Genus: Anthenea
Armament of the distal plates not dissimilar to that of
the other abactinalm plates, though the tubercles may be
somewhat enlarged and more numerous than proximally;
primary aboral tubercle stout, flat-topped and widely spaced
with more than one tubercle ....... Anthenea tuberculosa
42
Form massive, adults with major radius often well in
excess of lOOmm, interradial areas extensive, the arms
tapering and stellate or short, body almost flat below but
usually markedly convex above, often highest at the five
primary radial plates which may each be crowned by a high
conical prominence, marginal plates well developed but not
very conscipicous in aboral view ........ Family: Oreasteridae
Marginal plates concealed by thickened skin, some
enlarge tubercles present on the upper side, pore areas
usually rather irregular and sometimes indistinct or more
or less continuous ....................................... Genus: Culcita
A distinct pore free area at least towards the lower
side at the margins as well as more or less extensive
reticular areas on the upper side, though some of the pore
areas may be somewhat confluent, some larger, usually
spaced tubercles often also present, No spines or spinelets
on the pore areas ........................ Culcita schemideliana
Actinal granulation including more or less numerous
coarse, often polygonal granules, often in groups but
generally with some fine granules among the coarse ones,
pores area somewhat irregular and often only narrowly or
incompletely separated ................... Culcita noveguineae
At least the distal marginal plates and the convex
part of the larger abactinal plates covered with asmooth
plastering of unequal polygonal flattened granules, dorso
lateral areas of the arms rarely with any convexities .......
...... ...... ...... ...... ....... ...... ...... ...... ...... ..... Genus: Protoreaster
A few of the distal supero-marginal plates bearing
laterally projecting, usually conscipicous, tapering spines
or at least knobs ................................ Protoreaster lincki
Disc markedly elevated, some of the carina I plates
with very conscipicous, more or less high, rounded or
conical elevations, particularly huge on the five primary
radial plates, the consecutive carinal tubercles usually
43
spaced from each other and not broadened; pore areas
confluent ....................................... .. Protoreaster nodosus
Only the primary plates of the upper side with elevations
and these tending to form regular longitudinal series;
pore-areas usually well defined ......... Genus: Pentaceraster
No spines on the first two or four supero-marginal pIa tes
in each interradial angle; some of the interradial superomarginals with spines; dorso-Iateral elevations or spines
developed along the arms ............ Pentaceraster regulus
The distalmost supero marginals not conscipicously
different from the other marginal plates, arms usually well
developed but some genera stellate or even pentagonal or
circular in outline ................................ Genus: Choriaster
Entire surface covered by opaque smooth skin,
interrupted only by the rela tively small adam bulacral spines
and the popular areas, arms well-developed but short and
broadly rounded at the tip ............ Chorister granulatus
Edge of the body defined by the two series of marginal
plates, the supero-marginals sometimes smaller than the
infero-marginals but always conscipicously different from
the paxillae ................................ Family: Astropectinidae
Periphery fringed with conscipicous large spines
......................................................... Genus: Astropecten
Arms more or less blunt at the tip, the paxillar areas
ending abruptly; ventral sides of the infero-marginal
plates usually with few spines among the small rounded
scales, sometimes only on the interradial plates .............. .
........................................................... Astropecten indicus
Supero-marginal plates relatively narrow, the paxillar
area at the base of the arm distinctly more than half the
total arm breadth, usually abut two-thirds; ventral side od
the infero-marginal superficially appearing very smooth,
44
being covered with short, rounded more or less apressed
squamules .............................. Astropecten monacanthus
Body with cylindrical arms or sometimes cushion
like rounded ventro- laterally; aboral skeleton usually
reticular with relatively large space between the plates
often small. ...................................... Family: Asteropsidae
Body covered with smooth skin obscuring the nonimbricating oval or circular abactinal plates which are
nearly all quite naked, though bearing crystal-bodies
embedded in their surface; the edge of the body formed
by supero marginal plates with prominent conical single
.
Ast
OO
01'.
spInes..............................................
・イッーウセ@
」。イセョQ@
era
Armament predominantly granuliform, usually
continuous, though sometimes increasing in size or
modified into tubercles, basal as well as distal marginals
bearing granules, intermarginal plates present and
then only basally ....................... Family: Ophidiasteridae
Plates rounded and completely granule-covered, not
markedly swollen, pores in groups, rarely occurring on the
lower side .................................................... Genus: Linckia
Subambulacral spines are in two series, those of each
plate contiguous with each other and with the furrow
spines but aligned slightly obliquely so as to give a herring
bone pattern to the underside of each arm ...................... ..
..... ...... ....... ...... ...... ...... ...... ...... ....... ........ Linckia guildingi
Subambulacral spines or tubercles very low and
surrounded by the granulation, usually only a single
series present, granules extending down between the
furrow spines, especially numerous in large specimens,
arms normally five in number and madreporite single.
Arms fairly stout and blunt at the tips, body colour blue .
............................................................... Linckia laevigata
Arms often irregular in length and number, normally
two madreporites; Arms more slender and attenuated
45
towards the tips; colour in life often variegated, purpulish,
reddish, brownish or khaki coloured with yellowish,
sometimes more nearly uniform ........... Lincki multifora
Papular pores present on the oral side, no pores in the
infero-marginals, pores single, form more or less flattened,
the marginals usually defining the edge of the body ........ .
...................................................................... Genus: Fromia
Carinal plates not conspicuously enlarged and widely
separated from the supero-marginals. Denuded plates with
bumpy surface due to embedded crystal bodies, actinal
pores few or absent on the disc, abactinal granulation is
very fine .................................................... Fromia monilis
Abactinal plates markedly unequal in size, usually
two longitudinal series of distinctly enlarged plates on
each arm ..................................................... Fromia indica
Tube feet tapering to a rounded or conical knob, no
terminal disc; tube feet cylindrical and with a terminal
disc; edge of the body defined by the large infero- marginal
plates alone, the supero-marginals indistinguishable from
the paxillae ............................................. Family: Ludiidae
No large bivalve pedicillarie at the apics of the jaws
close to the mouth; most of the arm breath taken up
by the larger, quadrangular lateral paxillae, upto five
longitudinal rows of which each side also tend to form
transverse rows, often many of these bearing single
enlarged spines ............................................ Genus: Luidia
No conscipicous single paxillar spines, though the
smaller proximal paxillae may have alarge blunt central
spinelet about twice as high as the peripheral spinelets;
some or many ofthe lateral paxillae armed with large sharp
single spines; seven to nine arms ........ Luidia maculata
Ten to twenty arms and numerous madreporites,
aboral armament consisting of large conical isolated
46
spines mounted singly on stalklike pedicle ...................... ..
................................................... Family: Acanthasteridae
Usually five to six arms and only one madreporite,
sometimes two. Armament variable but if conical
spines are present then these are mounted only von low
eminences ......................................... Genus: Acanthaster
Aboral spines are long and conscipicous, commonly
15-30mm, long when major radius exceeds 100mm,
Pedicellariae slender ........................ Acanthaster planci
Conscipicous marginal plates forming a broad
vertical edge to the arm usually wanting, Aboral skeleton
reticulate or imbricate, tubefeet in two rows with suckers,
Pedicellariae rarely present.. ............. Order: Spinulosida
No enveloping aboral granulation, Spines sometimes
conscipicous but more often diminutive, adambulacral
spines few, usually about 3 in number .............................. .
...................................................... Family: Echinasteridae
Skeletal reticulum irregular aborally, no well defined
longitudinal and transverse series of plates ...................... .
...... ...... ...... ...... ...... ...... ...... ....... ...... ........ Genus: Echinaster
Five or six rays, occasionally seven, autonomous with
single separated arms regenerating to produce comet forms
but apparently not fissparous, arms slender and cylindrical,
often relatively long ....................... Echinaster luzonicus
Class: OPHIUROIDEA
No dorsal process of the arm bases, both tooth papillae
and oral papillae present, the former usually numerous
but occasionally few ..................... Family: Ophiocomidae
Disc covered, at least dorsally, with a dense coat of
rounded granules, the marginal ones either similarly
spherical or else very slightly elongated ............................ .
...... ...... ...... ...... ...... ...... ....... ...... ...... ....... Genus: Ophiocoma
47
Colour uniformly dark, above and below, Two tentacles
scales, Colour paler on the underside, variegated or
sometime uniformly dark above ..... Ophiocoma erinaceus
Colour of the disc either uniformly dark or variegated,
reticulated or spotted with dark markings ........................ .
...................................................... ...... Ophiocoma dentata
Disc granulation coarse, 3-6 granules/mm. length;
uppermost arm spines usually thickened and cigar sha ped or
cylindrical, rarelytapering ..... Ophiocoma scolopendrina
Disc completely naked, uppermost spines similar to
each other, none abruptly and markedly clavate though
some spines may be slightly so ...... Genus: Ophiarthrum
Disc beautifully patterned with grey, yellow and
white, at least in life, Arms with a dusky longitudinal
stripe dorsally ................................ Ophiarthrum pictum
No oral papillae, each jaw crowded with a more or
less compact cluster of apical tooth papillae .................... ..
..................................................... .Family: Ophiotrichidae
Radial shields with some or many thorny granules or
short stumps similar to, but shorter than, the stumps on
the rest of the rather puffy disc, which completely obscure
the countless underlying scales, Arm spines usually
transparent .................................. Genus: Macrophiothrix
Longer arm spines with the sides parallel or only
slightly divers gent distally and finely thorny for most of their
length, not at al clavate, Radial shields granules covered;
colouration spotted .............. Macrophiothrix longipeda
Ventral arm plates hexagonal, narrower at the
proximal end, the distal edge slightly concave .................. .
.............................................. Macrophiothrix propinqua
Disc densely granulated, also the oral plates,
sometimes even the oral and adoral shields .................... ..
................................................... Family: Ophidermatidae
48
Arms spines long and flaring, all of them easily exceeding
the segment in length; apical oral papillae sometimes rather
irregular, stimulating the tooth papillae of ophiocomids,
though much numerous .................. Genus: Ophiarachna
Arms spines four, rarely five basally; size very large,
colour in life including extensive areas of uniform green,
becoming yellow or buff in preservation and running along the
arms above and below, contrasting with interradial areas of
black -ringed light sports forming a reticula tion and with the
annulated arm spines ............. Ophiarachna incrassata
Class: ECHINOIDEA
Primary spines large, widely separated, contrasting
markedly with numerous, small secondary spines .......... ..
................................................................. Order: Cidaroida
Interambulacral plates high, each with a single
massive primary spine, usually ringed by much smaller,
often spatulate secondary spine, ambulacral plates
simple so that the pore-pairs form single vertical series,
sometimes sinuous ................................ Family: Cidaridae
In the apical system the ocular plates usually nearly
all insert, large globiferous Pedicellariae usually with a
limb or frill of rods on the stalk close to the head ............ .
.......................................................... Genus: Prionocidaris
Primary spines distinctly verticillate, with three or
four spaced complete thorny whorls along their lengths ..
...... ...... ...... ...... ...... ...... ....... ..... Prionocidaris verticillata
Test very flexible, in life more or less hemispherical
in shape but usually collapsing into a flat pancake on
preservation ......................................... Family: Arbaciidae
The ambulacral and interambulacral tubercles
developed throughout the adapical and adoral sides .........
.................................................................... Genus: Arbacia
49
Plates of abactinal system and upper interambulacral
plates coarsely granular, not marked with deep red in contrast
with the ground colour .................... Arbacia punctulata
Epiphysis of the Aristotle's lantern not fused across the
top of each pyramid, Spines lack a cortex and are solid or
provided with a narrow lumen ...... Order: Diadematoida
Primary tubercles perforate and often also crenulate,
Spines usually hollow, long, cylindrical and very slender
and breakable ................................ Family: Diadematidae
Ambulacral spines of the aboral side very fine and
needle like with backwardly-directed barbs near the tip,
contrasting with the other spines ..... Genus: Echinothrix
Ambulacra distinctly bulging aborally, with naked
interambulacral areas between them apically, Naked test
usually greenish in colour ...... .... Echinothrix calamaris
Ambulacra not distinctly bulging and no naked
interambulacral areas aborally, Cavity of primary spines
very small, their surface with fine longitudinal ridges
only , No green colour on the naked test ............................ .
........................................................ Echinothrix diadema
No spines on the buccal plates, no globiferous
pedicellariae ............................................ Genus: Diadema
Large tridentate pedicellariae mostly with narrow
blades only meeting at the tip, A red ring around the
anus ...................................................... Diadema setosum
Tridentate pedicellariae leaf or spoon shaped, tapering
slightly to the rounded distal end ...... Diadema savignyi
Spines are solid with or without cortex teeth
unkeeled .................................... Order: Phymosomatoida
Stirodonts with: apical disc hemicyclic; pore-pairs in
strongly offset arcs at ambitus and adapically; Primary
tubercles imperforate and non-crenulate .......................... ..
...................................................... Family: Stomechinidae
50
No complete bridges across the V-shaped space at
the upper end of each of the five pyramids forming frame
of the Aristotle's lantern, Ambulacral plates doubly
compound so that at the ambitus one very large tubercle
correspondence to three to six arcs each of three pore
pairs ....................................... Stomopneustus variolaris
Epiphysis of the Aristotle's lantern not fused across
the top of each pyramid ............ Order: Temnopleuroida
Tubercles usually crenulate; test usually with distinct
pits, troughs or pores at the angles of the sutures, or the pIa tes
more extensively sculpture ..... Family: Temnopleuridae
Crenulations conscipicous, Pore pairs more or less
distinctly in arcs of three .............. Genus: Temnopleurus
Spines not banded but often greenish basally becoming
purple distantly, or else uniform in colour- green, purple
or white ..................................... Temnopleurus alexendri
Tubercles distinctly crenulate, pores distinct; no
extensive bare areas on the test.. .......... Genus: Salmacis
Base of spines green ...................... ...... Salmacis belli
Ten conspicuous abruptly spineless vertical areas on
the test covered with extremely numerous globiferous
pedicellariae, forming a dense carpet, their valves with
the blades bearing two or three lateral teeth each side ....
...... ....... ...... ...... ...... ...... ...... ....... ...... ...... ...Mespila globulus
Globiferous pedicellariae less conscipicously developed
the blade fairly narrow and bearing not more than one
lateral tooth each side, tubercles indistinctly crenulate;
sutural pores very small and inconscipicous; naked areas
conspicuous aborally ........................ Genus: Macrocyphus
Test patterned with olive-green; spines banded with
reddish brown and white ......... Microcyphus ceylanicus
Gill slits sharps and deep, test circular or pentagonal,
as viewed from above ............... Family: Toxopneustidae
51
Globiferous
pedicillarie
not
enlarged
but
inconscipicous; only one in three or four ambulacral plates
with a primary tubercle; test high almost globular; porepairsin horizontal arcs and spaced to form three distinct
vertical series in larger specimens; preserved material
often with a conscipicous dark vertical stripe down each
inter ambulacrum contrasting with the light ambulacral
areas and white spines .................... Tripneustes gratilla
Lantern and teeth present throughout life; test usually
more or less flattened with a low sometimes acute margine,
rarely the test ovate ...................... Order: Clypeasteroda
PIa tes of the petals all alike and running across half
the width of the petal; aboral miliary spines ending in
a crown or a glandular bag; five genital pores; periproct
below the margin ........................ Family: Clypeasteridae
No interradial projections around the peristome ........
................................................................ Genus: Clypeaster
Generally smaller; test flat orally; frontal petals
closed .................................................. Clypeaster humilis
Test always very flat and perforated by two or more
lunules; a pair of interambulacral plates heading the
double series apically, miliary spines ending in a glandular
bag ................................................. Family: Astriclypeidae
Only two lunules, situates in the posterior paired
ambulacra ......................................... Genus: Echinodiscus
Lunules open distally ............. Echinodiscus auritus
Primary and secondary spines not markedly
contrasting in size .................................. Order: Echinoida
Shape either circular or more or less ovate, Periproct
with multiple plates; globiferous pedicellariae with an
unpaired lateral tooth .............. Family: Echinometridae
Test usually ovaL ..................... Genus: Echinometra
52
Long aXIS of the test through ambulacrum I and
interambulacrum 3, Spines slender and acute, Usually only
pore-pairs per arc aborally ........... Echinometra mathaei
Long axis; primary spine often massive and either
very long or truncated and very short, but even if they
are slender then the lower ones are more or less flattened
and some are angular, Eight or more pore-pairs per arc
aborally .......................................... Echinometra oblonga
Primary spines massive, all either rounded and are
very thick but somewhat flattened towards the tip, or
triangular in cross section and slightly tapering .............. .
...................................................... Genus: Heterocentrotus
Usually 15-16 pore-pairs in each arc at the ambital
region, primary ambulacral tubercles gradually decreasing
in size aborally and their spines correspondingly only
gradually shorter and not truncated .................................. .
...... ...... ...... ...... ...... ...... ........ Heterocentrotus trigonarius
Class: HOLOTHUROIDEA
Podia (Pedicels and papillae) present, body usually
stout, body wall more or less thick and muscular, usually
1-15mm thick, dominant spicules in the form of tables,
perforated plates, buttons, cups, rods or rosettes present
in the body wall .......................... Order: Aspidochirotida
Gonads in a single tuft to the left of the dorsal
mesentery, Spicules diverse in form and combination, 'S'
or 'C' shaped rods not present ..... Family: Holothuriidae
Spicules: tables nearly always present buttons, rods,
perforated plates and rosettes present or absent, minute
dichotomously branched or lobed rods rarely present and if so
then only in combination with tables,Anus guarded by five
calcified papillae, 20-30 tentacles ...... Genus: Actinopyga
Colour completely black ........... Actinopyga miliaris
53
Colours brown or brown and white, Colour brown on
upper side and white on lower side; often found near low
water mark ................................ Actinopyga mauritiana
Colour completely brown with often sand deposits on
upper side of the body, mostly found din deeper waters ...
......................................................... .Actinopyga echinites
Body colour: light grey or brown; Spicules composed of
small rods ....................................... Actinopyga lacanora
No calcified anal teeth present though five groups of
papillae may be evident, presence of 20 tentacles ............ .
..... ....... ...... ...... ...... ...... ...... ...... ....... ...... Genus: Bohadschia
Colour black or brown with distinct eye like spots all
over the body ....................................... Bohadschia argus
Colour variable, usually light brown with black spots
................................................. ... Bohadschia marmorata
Body colour: pale cream with brown speckling and low
papillae, spicules resemble a somewhat tack-like bundle
of spinose rods described as racquet-like .......................... ..
.......................................................... .Bohadschia graeffei
Spicules: variously developed and III varIOUS
combinations,Calcareous ring never ribbon-like, radial
plates either as long as broad or longer, intertidal plates
usually half as long as broad but never curved, Body wall
variously developed often rather thick. Body form showing
a wide range but pedicles usually irregularly arranged
on a more or less flattened ventral 'sole' and papillae
irregularly arranged on the arched dorsal surface .......... ..
............................................................... Genus: Holothuria
Body tubular, body wall not very thick. No lateral
projections in the living condition, Body completely black
in colour, red colour comes off when live specimens are
handled .................................................... Holothuria atra
54
Yellow transverse band on the upper side of the body,
lower side white with a number of black dots .................. .
..... ....... ...... ...... ...... ...... ...... ...... ....... ...... .Holothuria scabra
Body like a loaf with very thick body wall. In the living
condition about six pairs of lateral teat-like projections
are seen, Body with black or white patches .................... ..
...... ...... ...... ...... ...... ...... ....... ...... ...... ...... .Holothuria nobilis
Buttons of the spicules have large holes, In the spicules
tables stout with a cluster of short spines at the top fugitive
form skin sandy to touch ........ ...... Holothuria impatiens
Spicules: tables not stout and with a few spines at the top
skin smooth and soft, fugitive form ....... Holothuria hilla
Spicules: buttons with small holes burrowing form
with often red spots ...................... Holothuria arenicola
Body large and snake like, Spicules: tables with
complete or incomplete discs often reduced to four holes,
buttons may sometimes be asymmetrical, tentacles- 20 ...
...... ...... ...... ...... ...... ...... ....... ...... ... Holothuria leucospilota
Tentacles large, sub-globose when fully expanded.
Pedicles arranged in three rows. Papillae irregular,
Spicules: tables and rods present, Rods simple and
granulated ............................... ... Holothuria cinerascens
Body colour pink with varying degrees of black
pigment, numerous pedicles on the ventral side, Spicules:
Tables and buttons present, discs of tables is narrower
than the top of the spire ..................... Holothuria edulis
Body colour brownish pink with some of the tubercles
blackish brown, Spicules consist of only tables in the body
wall ..................................................... .... Holothuria pyxis
Papillae scattered on the dorsal side, Cuvierian
tubules are thick, Body colour brown with 5-7 honey
coloured transverse bands of different widths. Spicules:
55
incomplete and oblong rods with lateral projections
resembling narrow rosettes, tables sub circular ................ .
..... ...... ....... ...... ...... ...... ...... ...... ....... ... Holothuria pervicax
Body colour brownish black with grey ventral side,
Cuvierian tubules large. Button has two narrow slit-like
holes and one or two pairs of minute holes at each end ..
..... ...... ....... ...... ...... ...... ...... ...... ... Holothuria fuscocinerea
Body
colour
black
with
white
papillae,
Spicules include spinose 'cup and saucer' tables and
rosettes ............................................. Holothuria coluber
Spicules: rods only, usually dichotomously branched or
lobed, tables and buttons never present, Calcareous ring
very stout, both radial and interradial plates with their
anterior margin scalloped and sometimes with their suture
indistinct, radial plates usually about twice as large as the
interradials and possessing median anterior ampullary
notch, Tentacles: 20-30, Body wall very thick and muscular,
usually about 5mm. thick, pedicels and papillae small and
numerous, indistinguishable from each other, scattered
ventrally and dorsally .......... Family: Labidodematidae
Spicules: tables scattered, variously developed, either
with disc reduced and spire low and ending in a row of
spines or else disc well developed and spinose, with spire
of moderate height and usually very spinose, Buttons
when present very smooth and irregular, often incomplete
or deformed and suggesting clumsy 'C' shaped body,
Body cylindrical or vermiform with pedicels and papillae
confined to the five ambulacral areas, size moderate,
Cancerous ring ribbon like, radial plates usually shorter
than broad, intertidal plates also shorter than broad
tending to be curved ...................... Genus: Labidodemas
White, almost translucent sea cucumber, with
yellow tube feet and a dark posterior end, Spicules are
spinose tables and sometimes buttons and rods .............. ..
............................................. Labidodemas semperianum
56
Gonads in two tufts, one each side of the dorsal
mesentery, Spicules: primarily tables, branched rods and
'S' or 'C' shaped rods, the latter very rarely absent, or
slender dichotomously branched rods, buttons rarely, if
ever, present ............................... Family: Stichopodidae
Spicules reduced: grains and dichotomously- branched
rods, pedicles crowded ventrally, papillae resembling large
leaf shaped structures dorsally ............. Genus: Thelenota
Very massive form with numerous large pointed teats
in groups of two or three all over the upper surface ........
.............................................................. Thelenota ananas
Spicules: tables, branched rods and Sand C shaped
rods (the latter absent only in some growth stages of
Stichopus horrens, H.L.Clark, 1922) buttons rare ............ .
..... ....... ...... ...... ...... ...... ...... ...... ....... ...... .... Genus: Stichopus
Body quadrangular with four rows of large finger like
processes, Body colour dark green, appearing almost black
in some shades of light.. ............. Stichopus chloronatus
Body massive and loaf-like with irregular brown patches
on yellow grey background ............ Stichopus variegatus
Body colour grey to green-black with dark patches,
smooth tegument but large and irregular papillae, Spicules
are tables and large "C" bodies .......... Stichopus horrens
The body wider in the middle, tapering at both ends,
covered with a bright smooth skin, upper surface uniformly
brown, zigzag bands at all over the body, spicules consists
of tables, branched rods and'S' and 'C' shaped rods .........
................................................................. Stichopus vastus
Tentacles bush ortree shaped, dendritic. Anterior end in
the form of a thin walled introvert capable of retraction
by special muscles ...................... Order: Dendrochirotida
57
Body cylindrical or fusiform, without a well defined
ventral 'sole', no conscipicous dorsal plates, Mouth and
anus terminal in position, Tentacles 10 ............................ .
....................................................... .Family: Cucumariidae
Spicules: small nodular buttons with few holes,
usually four, sometimes larger perforated plates present .
........................................................................ Genus: Stolus
Body colour dark, purple-black and brown or grey colour,
Spicules: small nodular buttons .............. Stolus buccalis
Podia absent, though warty prominences may be
present, Body either vermiform, body wall thin, thick
or less often translucent, with smooth, rough or warty
surface, or body stout and sausage-shaped with a caudal
appendage ................................................ Order: Apodida
Spicules: anchors and anchor-plates, rods and granules,
never wheels or sigmoid particles, rarely spicules wanting,
Tentacles pinnate or digitate ............ Family: Synaptidae
Anchor-plates not abruptly contracted at the posterior
end but with a large hole on each side, Calcareous ring without
conscipicous anterior projections, Stone canals always
few in number (one to three) ..................... Genus: Euapta
Body colour: mottled cream-white with grey,
spicules are anchors with tiny knobs on the vertex and pIa tes
with large posterior holes .................. .Euapta godeffroyi
Spicules: characteristically large, Anchor plates sub
rectangular or irregular broad posterior with numerous
smooth holes .............................................. Genus: Synapta
Tentacles- 15; Body colour: (brown-yellow) with broad
longitu dinal stripes and large dark patches .................... .
...................................................... ........ Synapta maculata
58
7. DESCRIPTIONS
Class CRINOIDEA Miller, 1821
59
Order : COMATULIDA A. H. Clark, 1908
Family: COMASTERIDAE Clark, 1908
Genus : Comanthus
1. Comanthus parvicirrus (Muller, 1841)
Description: Cirri are very short in size. It is
discontinuously arranged around the edge of the
centrodorsal and their distal segments only slightly shorter
than the proximal ones. Arms are 10-63 in number. Cirri
are formed with 11-16 segments. Distal segments bear
a sharp aboral transverse bar or tubercle. Centrodorsal
are small in size discoidal in shape. Radials are easily
visible.
Colour: Brownish black.
Habitat: Benthic, inshore in habitat and this species
is suspension feeder.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar; Elsewhere: Aldabra, Eastern Mrica,
Madagascar, Fiji, Great Barrier Reef, Indonesia, Marion
Reef, Mascarene Basin, Mozambique, New Caledonia,
Palau, Philippines, Seychelles, South China Sea, South
Japan, Thailand Exclusive Economic Zone, Tongan
Exclusive Economic Zone and West Indian Ocean.
Comanthus parvicirrus (Muller, 1841)
60
I Genus
: Comanthina I
2. Comanthina nobilis (P.R. Carpanter, 1884)
Description: Central disc and proximal portion of
arms have black spots. Tips of pinnules are yellow in
colour. Cirri are 0-5 in number. Proximal aboral surface is
heavily plated and smooth. Arms are 50-140 in number.
Combs of pinnules arising from brachials often with one
tooth per segment.
Colour: Body is uniformly black in colour.
Habitat: It is found over 8m depth ranging up to
92m.
Distribution: India: Andaman Islands; Elsewhere:
Coral Sea, Great Barrier Reef, Indonesia, Malaysia, New
Caledonia, Philippines and Sri Lanka.
Comanthina nobilis (P.H.Carpanter, 1884)
61
I Genus
: Com aster
3. Comaster schlegeli (P.R. Carpanter, 1881)
Description: This species bears 9-33 cirri which
consist of 11-17 segments. Proximal aboral area is relatively
smooth. Radial plates are narrowly visible. Arms are 1090 in number. Brachial pinnules (except possibly PI) with
a pair of more or less equally sized teeth.
Colour: Body colour- Grey.
Habitat: Found over 5 m depth.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar; Elsewhere: Fiji, Indonesia, Japan and
Kerama Group.
Comaster schlegeli (P.H.Carpanter, 1881)
62
4. Comaster multibrachiata
(P.R. Carpanter, 1888)
Description: Cirri are 21-50 in number. Each cirrus
is of 12-15 segments. Arms are more than 100 in number
with a maxiumum of 170.
Colour: Body colour is blue with orange colour tips of
the pinnules.
Habitat: It is found on reefs at variable depth.
Distribution: India: Andaman Islands; Elsewhere:
East Indies, China, South Japan and Philippines.
Comaster multibrachiata (P.R. Carpanter, 1888)
63
I Genus
Oxycomanthus I
5. Oxycomanthus bennetti (Muller, 1841)
Description: Arms are 31-120 in number. Anterior
arms not markedly longer than posterior arms. Cirri consist
of 23-35 segments (usually 25-29) and these segments
usually as broad as long and without aboral processes.
Centrodorsal is large and thick, covering radials.
Colour: Uniformly yellow in colour.
Habitat: This species is commonly found below 5m
depth. It mainly occurs on live coral specimen.
Distribution: India: Andaman Islands; Elsewhere:
East Indies, North Australia, Philippine, China, south
Japan and South Pacific Islands.
Oxycomanthus bennetti (Muller, 1841)
64
Family: HIMEROMETRIDAE
Genus : Himerometra
6. Himerometra robustipinna (P.R. Carpanter, 1912)
Description: This species have stout, enlarged
proximal pinnules that taper rapidly or gradually to the
tip. Almost all segments are broader than long with 20-42
in numbers. The distal end of the segments is unmodified
or swollen, but never spinous. Distal edges of proximal
brachial smooth or slightly produced. Arms 33-52 up to
200 mm long.
Colour: Grey to white in colour.
Habitat: Found with the reefs.
Distribution: India: Andaman Islands; Elsewhere:
Ashmore Reef, China, China Sea, East Indies, Great Barrier
Reef, Indonesia, Japan, Lady Musgrave Island,North
Australia, Northern Territory, Okinawa, Philippines,
South China Sea, South Japan, South Pacific and Sri
Lanka.
Himerometra robustipinna (P.H. Carpanter, 1912)
65
Family: MARIAMETRIDAE
Genus : Heterometra
7. Heterometra philiberti (Muller, 1849)
Description: Arms are more than 20 in number. The
proximal pinnules are distinctly modified with a strong
crest or the segments have flared or spinose distal ends.
Colour: It is usually brown in colour. Sometimes it
can be seen in the form of brown-black specimen.
Habitat: Found in the rocky areas of reef slopes.
Distribution: India: Andaman Islands; Elsewhere:
East Indies.
Heterometra philiberti (Muller, 1849),
66
8. Heterometra crenulata (P.R. Carpenter, 1882)
Description: Proximal pinnules are large and strongly
triangular in cross section, strongly serrate in profile.
Outer portion of prismatic ridge on each segment raised
into conspicuous broad rounded triangular processes.
Distal cirrus segments occur with distinct dorsal spines
or tubercles.
Colour: Body colour Red with white ridges in arms.
Habitat: Found in the reef slopes.
Distribution: India: Andaman Islands; Elsewhere:
Cambodia, China, Double Island Point, East Indies,
Indonesia, Maldives, Monte Bello Islands, North Australia,
Philippines, Queensland, Singapore, South Japan, Vietnam
and West Australia.
Remark: New record to Andaman and Nicobar Islands
reported from Pongibalu, South Andaman.
Heterometra crenulata (P.R. Carpanter, 1882)
67
I Genus: Amphimetra I
9. Amphimetra molleri (AH. Clark, 1908)
Description: Usually 10 arms are present and rarely
more than 10 arms are found. Cirri are stout, curved and
formed of 24-50 short sub equal segments. First aboral
spine appears proximal to 8th cirrus. Arms are 150mm in
size. Outer pinnules are 18-21 in number.
Colour: Black in colour.
Habitat: Mainly found on shoreline to 50m depth
range.
Distribution: India: Andaman and Nicobar Islands
and Gulf of Mannar; Elsewhere: China, East Indies,
Greater Sunda Islands, Gulf of Thailand, Maldives,
Philippines, South Japan and Sri Lanka.
Remark: New record to Andaman and Nicobar
Islands
Amphimetra molleri (AH.Clark, 1908)
68
I Genus: Lamprometral
10. Lamprometra palmata (Muller, 1841)
Description: Second interradial arm is stout or
distinctly carinate basally and has the largest proximal
pinnule. Arms more than 140 in number. This species is
commonly distributed in tropical Indo-Pacific Ocean.
Colour: Arms greenish black in colour with white
patches
Habitat: Benthic, inshore, continental
suspension feeder. Depth range: 0-51m
shelf,
Distribution: India: Gulf of Mannar, Andaman and
Nicobar Islands and Karwar Coast; Elsewhere: Aldabra,
Eastern Africa, Madagascar, Red Sea and Tanzania.
Lamprometra palmata (Muller, 1841)
69
Family: COLOBOMETRIDAE
Genus : Cenometra
11. Cenometra bella (Hartlaub, 1890)
Description: Outer pinnules (2) are very stout,
stiff and erect or even recurved over the disc. Arms are
more than 10 in number. Basal segments of none of the
proximal pinnules carinates. A simple keel on the bases of
the proximal pinnules forms rounded edge.
Colour: Central ridge of the arm is white and pinnules
are dark red in colour.
Habitat: Commonly found below 5m depths and it
lives with the association of encrusting and massive coral
speCIes.
Distribution: India: Andaman Islands; Elsewhere:
East Indies, Philippines, China, South Japan and South
Pacific Islands.
Remark: New report from India.
Cenometra bella (Hartlaub, 1890)
70
12. Cenometra emendatrix (Bell, 1892)
Description: The Basal segments of the proximal
pinnules have knob-like rounded process that forms a
crest along the edge. Arms are 10 in number.
Colour: Dark red in colour
Habitat: It occurs in Littoral to 55m depths.
Distribution: India: Andaman Islands; Elsewhere:
Aldabra, Madagascar, Mascarene Basin, Mauritius,
Seychelles and West Indian Ocean.
Remark: New record to India reported from Rutland
Island, South Andaman.
Cenometra emendatrix (Bell, 1892)
71
I Genus
Oligometra
I
13. Oligometra serripinna (P.R. Carpanter, 1881)
Description: Pinnules are variable in form with
most segments as long as, or longer than, broad, either
simply prismatic with only the dorsal keel well developed
or the segments more or less flared at the distal ends on
the angels of the longitudinal ridges, with spinose process,
so that the profile of the pinnule is more or less serrated.
Colour: Uniformly white in colour with reddish
patches throughout the body.
Habitat: Depth range 0-91m.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar, Lakshadweep and Andhra Pradesh coast;
Elsewhere: Aldabra, Eastern Africa, Madagascar,
Mascarene Basin, Mauritius, Red Sea, Seychelles, West
Indian Ocean.
Oligometra serripinna (P.R. Carpanter, 1881)
72
Family: PONTIOMETRIDAE
Genus : Pontiometra
14. Pontiometra andersoni (P.R. Carpanter, 1889)
Description: Middle and distal cirrus segments are
attached with a pair of dorsal spines or tubercles that
situated one each side of the mid-line. Arms are 53-120 in
number. Cirri are composed of 41-80 segments.
Colour: Uniformly red in colour.
Habitat: Depth range 0-91m.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: East Indies, Philippines, China, South Japan,
South Pacific Island.
Pontiometra andersoni (P.R. Carpanter, 1889)
73
Class: ASTEROIDEA de Bklainville, 1830
74
Order : VALVATIDA Perrier, 1884
Family : ASTERINIDAE
Genus : Asterina
15. Asterina sarasini (de Loriol, 1897)
Description: Body is usually stellate and pentagonal.
The margin of the periphery is thin and pliable. Abactinal
armament is composed of fine minute hyaline spinelets.
Body is markedly flattened and the petaloid radial
popular areas are linked only to the area in the centre of
the disc.
Colour: Mter preservation the speCImen becomes
white in colour. Live specimens are uniformly grey or
variegated colour.
Habitat: This species is commonly found in intertidal
rocky areas and shallow reef areas.
Distribution: India: Andaman and Nicobar Islands
and Gulf of Mannar; Elsewhere: Ceylon.
Asterina sarasini (de Loriol, 1897)
from left to right ventral and dorsal
75
Family: GONIASTERIDAE Forbes, 1841
Genus : Stellaster
16. Stellaster equestris (Retzius, 1805)
Description: Arms are slender and long. Granules
around the papulae are usually similar to the rest. Radial
areas on the upper side are distinctly convex. Only single
infer-marginal spine is present.
Body Colour: Live specimens are white or grey
colour.
III
Habitat: Rocky shore areas, benthic, sometimes found
in shallow reef environments.
Distribution: India: Gulf of Mannar, Andaman
Islands and West Bengal: Digha; Elsewhere: East Mrica,
Madagascar, Red Sea, S.E Arabia, Persian Gulf, Ceylon,
East Indies, North Australia, Philippines, China and
South Japan.
Stellaster equestris (Retzius, 1805)
76
Family : OREASTERIDAE
Genus : Culcita
17. Culcita noveguineae (Muller and Troschel, 1842)
Common name: Pin Cushion Sea Star
Description: Actinal plates have polygonal granules
and granulation forms more or less numerous coarse.
Pores area is somewhat irregular and often only narrowly
or in completely separated. It is common in reef flats at
low tide and also inhabits in deeper waters to 30m.
Colour: Body colour is light greenish black.
Habitat: It feed on a variety of organism including
algae, bottom detritus and the polyps and some flesh of
corals.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Persian Gulf, East Indies,
Philippines, Northern Australia, China, South Japan and
South Pacific Island.
Culcita noveguineae (Muller and Troschel, 1842)
from left to right dorsal and ventral
77
18. Culcita schmideliana (Retzius, 1805)
Description: Presence of a distinct pore free area at
least towards the lower side at the margins as well as more
or less extensive reticular areas on the upper side. Some
of the pore areas may be somewhat confluent. Some pores
are larger in size. Spaced tubercles often also present.
There are no spines or spinelets on the pore areas.
Colour: Body colour is dark grey with small
irregular pink patches mostly adjacent to black tubercles.
Madreporite is some shade of orange being the same
colour as the intertubercular areas at the arm tips.
Habitat: It is commonly found in the Indo-Pacific
region. No spines or spinelets are present on the pore
areas. It mainly occurs in benthic, inshore and continental
shelf area. Depth range: 0-92m.
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep; Elsewhere: Western Indian Ocean,
Mascarene Islands, East Africa, Madagascar, Maldives
and Ceylon.
Culcita schmideliana (Retzius, 1805)
from left to right dorsal and ventral
78
I Genus
: Choriaster
19. Choriaster granulatus (Lutken, 1869)
Description: Body surface is entirely covered by
opaque smooth skin. The skin is often formed with
relatively small ambulacral spines. The pore areas and
arms are well-developed but short and broadly rounded
at the tip.
Colour: Body is uniformly red in colour.
Habitat: It is usually found at the reef slopes.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: East Africa, New Caledonia ,Palau Islands,
Red Sea, Seychelles, Sulawesi and West Indian Ocean.
Choriaster granulatus (Lutken, 1869)
from left to right dorsal and ventral
79
I Genus
: Anthenea
20. Anthenea tuberculosa (Gray, 1847)
Description: Ambulacral plates consist of three
series of spines. Armament of the distal plates is more
or less similar to the abactinal plates. Primary aboral
tubercle is stout, fiat-topped and widely spaced with more
than one tubercle.
Colour: Body colour is white.
Habitat: It is found in the sandy area.
Distribution: India: Andaman Islands and Orissa
coast; Elsewhere: South Japan
Anthenea tuberculosa (Gray, 1847)
80
I Genus
: Protoreaster
21. Protoreaster lincki (Blainville, 1830)
Description: Furrow spines are white with pink
tube feet. The distal marginal plates and the convex part
of the larger abactinal plates are covered with a smooth
plastering of unequal polygonal flattened granules. Dorso
lateral areas of the arms are rarely with any convexities.
A few of the distal supero-marginal plates laterally
projected. Those are usually conscipicous. The tapering
spines look like knobs.
Colour: Body Colour
background.
IS
red on pink or grey
Habitat: It is found almost all over the sandy-muddy
areas of reef slopes. The deapth ranges from 1-10 m.
Distribution: India: Andaman and Nicobar Islands
and Gulf of Mannar; Elsewhere: Aldabra, Cargados
Carajos, Eastern Mrica, Kenya, Madagascar, Mascarene
Basin, Mauritius, Mozambique, Red Sea, Seychelles,
Somalia, Tanzania
and West Indian
Ocean.
Protoreaster lincki (Blainville, 1830)
81
I Genus
: Protoreaster I
22. Protoreaster nodosus (Linneaus, 1758)
Description: Disc is markedly elevated. Carinal
plates are very conscipicous, more or less high, rounded
or conical elevations. Ambulacral spines are white, tube
feet pink with purple centre, particularly huge on the five
primary radial plates, the consecutive carinal tubercles
usually spaced from each other and not broadened; pore
areas confluent.
Colour: Tubercles are yellow with orange tips and
papular areas blue-grey with paulae green. Remainder of
aboral surface is deep green; marginals are pale yellow in
colour.
Distribution: India: Andaman and Nicobar
Islands; Elsewhere: Cargados Carajos, Eastern Africa,
Madagascar, Kenya, New Caledonia, Palau Islands,
Seychelles, Tanzania and West Indian Ocean.
Protoreaster nodosus (Linneaus, 1758)
82
I Genus
: Pentaceraster
I
23. Pentaceraster regulus (Muller and Troschel, 1842)
Description: 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. Dorso-Iateral elevations or spines are
developed along with the arms.
Colour: Body colour is yellow with reddish spine on
the upper side.
Habitat: Usually found in sandy slopes of the marine
water. Depth ranges 2-115.
Distribution: India: Gulf of Mannar, Lakshadweep
and Andman and Nicobar Islands; Elsewhere: New
Caledonia, Red Sea and West Indian Ocean.
Pentaceraster regulus (Muller & Troschel, 1842)
83
Family: ASTEROPSIDAE Muller and Troschel, 1840
Genus : Asteropsis
24. Asteropsis carinifera (Lamarck, 1816)
Description: Body covered with smooth skin
obscuring the non-imbricating oval or circular abactinal
plates which oval or circular in shapes which are nearly
all quite naked. Crystal-bodies embedded in their surface,
the edge of the body formed by supero marginal plates with
prominent conical single spines. The carinal plates formed
a mid radial ridge emphasized by a series of spines.
Colour: Body colour pale grey with irregular greenish
marking or sometimes uniformly grey in colour.
Habitat: It is commonly found in sandy and rocky
areas. Some times it can be seen under the coral reefs.
Depth range: 0-15m.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Aldabra, Chagos, Eastern
Mrica and Madagascar, Kenya, Mascarene Basin,
Mozambique, New Caledonia, Red Sea, Seychelles and
West Indian Ocean.
Asteropsis carinifera -ventral
84
Asteropsis carinifera -dorsal
Family : ACANTHASTERIDAE
Genus : Acanthaster
25. Acanthaster planci (Linneaus, 1758)
Description: This species usually bears 15 to 18 arms
and usually more madreporite. Armament is variable but
if conical spines are present then these are mounted only
on low eminences. Aboral spines are long and conscipicous,
commonly 15-30mm in size. Pedicellariae are slender.
Colour: Body colour is blue with black large spines
on the upper side.
Habitat: It lives mainly with the association of coral
beds and sometimes taking shelter under stones close to
the live colonies.
Distribution:
India: Andaman and Nicobar Island, Lakshadweep;
Elesewhere:
Western Indian Ocean, Mascarene Islands, East
Mrica, Madagascar, Red Sea, South East Arabia, Maldive
area, Bay of Bengal, East Indies, Philippines, Northern
Australia, China, South Japan, South Pacific Island and
Hawaii.
from left to right ventral and dorsal
85
Family: OPHIDIASTERIDAE Verrill, 1870
Genus : Fromia
26. Fromia monilis (Perrier, 1869)
Description: Carinal plates are not large in size. It
is widely separated from supero marginal. Actinal pores
are few or absent. Abactinal granulation is very fine. This
star is quite hard to the touch and its rigid skeleton does
not allow it to be bent without breaking.
Colour: The specimen is uniformly red in colour
although some specimens may have solid red patches on
the disc.
Habitat: It occurs in shallow water lagoon areas
among living coral reefs, and on channel slopes between
reefs and on drop offs in moderately deep water.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: East Indies, Philippines,
China, South Japan and South Pacific Islands.
Fromia monilis (Perrier, 1869)
-ventral
86
Fromia monilis (Perrier, 1869)
-dorsal
27. Fromia indica (Perrier, 1869)
Description: Abactinal plates are unequal III size.
Each arm bears two longitudinal series of two distinctly
enlarged plates. Papular pores are present on the oral
side. Single pore forms more or less flattened marginals
that usually define the edge of the body.
Colour: It is reddish-brown in colour.
Habitat: It is usually found on the rocky shores of
reef areas.
Distribution: India: Andaman and Nicobar Island,
Lakshadweep and Gulf of Mannar; Elsewhere: Maldives,
East Indies, Philippines, China, South Japan, South
Pacific Island and Ceylon.
Fromia indica (Perrier, 1869)
87
I Genus
: Linckia
28. Linckia guildingi (Gray, 1840)
Description: Plates are covered with granules and
round in shape. Pores are present in groups. Subambulacral
spines are in two series.
Those of each ambulacral
plate contiguous with each
other and with the furrow
spines but aligned slightly
obliquely so as to give a
herring-bone pattern to
the underside of each arm.
Colour: Body colour is
uniformly brown.
Habitat: This speCIes
is commonly found III
shallow water depths and
inhabits in rocky reefs and
rocky shore areas.
Distribution:
Andaman
and
Nicobar
Islands
and
Lakshadweep
Islands;
EIswhere:
Mascarene
Islands, East Africa, Red
Sea, Madagascar, Maldives,
South East Arabia, Ceylon,
Persian Gulf, East Indies,
Philippines,
Northern
Australia, China, South
Japan, South Pacific Island
and Hawaii.
India:
Linckia guildingi (Gray, 1840)
88
29. Linckia laevigata (Linneaus, 1758)
Description: Subambulacral spines are distinctly
placed from furrow spines. Usually only a single series
present, granules are extending down between the furrow
spines, especially numerous in large specimens. Arms are
normally five in number. Single madreporite is present.
Arms fairly stout and blunt at the tips.
Colour: Body colour is blue.
Habitat: This species is commonly found in shallow
water depths and inhabits in rocky reefs and rocky shore
areas.
Distribution:
India: Andaman and Nicobar Islands, Lakshadweep
and Gulf of Mannar.
Elsewhere: Northern Australian reefs, Western Indian
Ocean, Mascarene Basin, East Africa and Madagascar,
Maldives area, Ceylon, East Indies, Philippines, China,
South Japan, South Pacific Island and Hawaii.
Linckia laevigata (Linneaus, 1758)
from left to right ventral and dorsal
89
30. Linckia multifora (Lamarck, 1816)
Description: Arms are variable in length and
number. Normally two madreporites are present. Arms
are more slender and attenuated towards the tips.
Colour: Body colour purplish, reddish, brownish
coloured with yellow.
Habitat: This sea star is much smaller and rarely
occurs in the open on intertidal reef fiat. It is a very
common species and is abundant at depths of 5-30m and
is generally observed on bommies and underhangs.
Distribution: India: Andaman and Nicobar
Islands, Lakshadweep and Gulf of Mannar; Elsewhere:
Maldives, Aldabra, Comores, Eastern Africa, Kenya,
Madagascar, Mascarene Basin, Mauritius, Mozambique,
Red Sea Seychelles, Somalia, South Mrica, Tanzaniaan
and West Indian Ocean.
Linckia multifora (Lamarck, 1816)
90
Order : SPINULOSIDA Perrier, 1884
Family : ECHINASTERIDAE
Genus : Echinaster
31. Echinaster luzonicus (Gray, 1840)
Description: Arms are five or six in number,
sometimes 7 in number. Arms are slender, cylindrical
and stellatae. Skeletal reticulum is placed aborally,
longitudinal and transverse series of plates are not well
defined. This species is commonly distributed in Indian
Ocean region.
Colour: Body colour is reddish brown, sometimes
may be uniformly brown in colour.
Habitat: It is found in shallow reef environments,
usually below 3m depth.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Gulf of Kachchh; Elsewhere: East
Indies, Philippines, China, South Japan, South Pacific
Island and Northern Australia.
Echinaster luzonicus (Gray, 1840)
from left to right dorsal and ventral
91
Order : PAXILLOSIDA Perrier, 1884
Family: ASTROPECTINIDAE Gray, 1840
Genus : Astropecten
32. Astropecten indicus (Doderlein, 1888)
Description: It has pointed legs and with that many
legs, the paxillar areas are ended abruptly. Ventral sides
of the infero-marginal plates are usually with few spines
among the small rounded scales. Sometimes spmes are
also present on the interradial plates.
Colour: Live specimens are white or grey in colour,
sometimes with black patches on the upper side.
Habitat: It lives in the sandy shore areas, or on dead
corals. This species is a very good burrower and it is in
this domain that it hunts for its preys.
Distribution: India:
Gujarat,
Orissa, West
Bengal, Tamil Nadu and Andaman and Nicobar islands;
Elsewhere: South East Arabia,
Persian gulf, Pakistan, Maldives,
Ceylon and East Indies.
Remark:
92
New record to
Andaman
and
Nicobar
Islands
33. Astropecten monacanthus(Sladen, 1883)
Description: Peripheral margins of the body are
fringed with conscipicous 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. Actinal
surface of the infero-marginal plates is very smooth
and covered with short, rounded more or less apressed
squamules.
Colour: Body is purple or reddish yellow or light grey
in colour.
Habitat: This species are lying on the sand and feeds
on sea shells.
Distribution: India: Tamil Nadu, Orissa coast and
Andaman and Nicobar Islands; Elsewhere: East Mrica,
Eastern Mrica, Madagascar, Mozambique and Red Sea.
Astropecten monacanthus (Sladen, 1883)
93
Family: LUIDIIDAE Verrill, 1899
Genus : Luidia
34. Luidia maculata (Muller and Troschel, 1842)
Description: Large bivalve Pedicellariae are absent
at the apics of the jaws close to the mouth. Single paxillar
spines are absent though the smaller proximal paxillae
may have a large blunt central spinelet about twice as
high as the peripheral spinelets. Arms are seven to nine
in number. Some or many of the lateral paxillae armed
with large sharp single spines.
Colour: Usually brown in colour, some times yellowish
with black patches.
Habitat: It is found in the depth of 1-10 m.
Distribution: India: Gulf of Mannar, Gulf of Kachchh
and Andaman Islands; Elsewhere: Western Indian
Ocean, East Mrica, Madagascar, Persian Gulf, Maldives,
Ceylon, East Indies, North Australia, Philippines, China
and South Japan.
Luidia maculata (Muller & Troschel, 1842)
94
Class OPHIUROIDEA Gray, 1840
95
Order : OPHIURIDA Muller and Troschel, 1840
Family: OPHIOCOMIDAE Lyman, 1865
Genus : Ophiocoma
35. Ophiocoma erinaceus (Muller
and Troschel, 1842)
Description: Arm length up to 12 cm, disc diameter
upto 20 mm. Spines on the arms is long, thick and shorter
towards the tips. Disc scales are covered by rounded
granules, extending into a V-shaped interradial area on
the ventral side of the disc. Dorsal arm plates broader
than long, narrower on one side. Arm spines alternating
three and four for greater part of the arm length.
Colour: Uniformly black in colour.
Habitat: Sand and rubble in shallow areas and
shallow sublittoral among coral; coral reef, beacon reef.
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep and Gulf of Mannar; Elsewhere: Aldabra,
Chagos, Comores,
Eastern
Africa,
Madagascar,
Kenya, Mascarene
Basin, Mauritius,
Mozambique, Red
Sea,
Seychelles, セi
エA セ セ@
Somalia,
South
Mrica, Tanzania セ As セ@
and West Indian
Ocean.
Ophiocoma erinaceus (Muller & Troschel, 1842)
96
36. Ophiocoma dentata (Muller and Troschel, 1842)
Description: Dorsal arm plates with a light border
along the edges and sometimes form a light central
region. Disc scales are with rounded granules, covering
the radial shields. Dorsal arm plates broader than long,
lateral angles rounded. Five arm spines, fiat, second from
above longer than the breadth of the segment.
Colour: Colour of
the disc uniformly dark
or variegated, reticulated
or spotted with dark
markings.
Habitat:
inshore.
Benthic,
Distribution: India:
Andaman and Nicobar
Islands and Lakshadweep
Islands;
Elsewhere:
Eastern
Africa,
Madagascar, Mascarene
Basin, Mozambique, Red
Sea and West Indian
Ocean.
Ophiocoma dentata (Muller
& Troschel, 1842) -dorsal
Ophiocoma dentata (Muller & Troschel, 1842) -ventral
97
37. Ophiocoma scolopendrina (Lamarck, 1816)
Description: Arm length up to 10 em, disc diameter
up to 20 mm. Disc scales are covered by granules, becoming
spiky towards the edges, higher than thick. Arms with
spines as long as the arm width. Disc mottled green, arms
distinctly banded. Only one tentacle scale at least beyond
the fifth segment.
Colour: Arms commonly black or green brown and
banded by darker areas but banding may be deep brown
or cream. Disc uniformly dull green to dark brown.
Habitat: It inhabits in shallow substrates; rocks,
coral reef, under weed.
Distribution: India: Lakshadweep; Andaman Islands;
Ophiocoma scolopendrina (Lamarck, 1816)
Elsewhere: Aldabra, Chagos, Eastern Africa,
Kenya,
Madagascar, Mascarene Basin, Mozambique, Red Sea,
Seychelles, Somalia, South Africa, Tanzania and West
Indian Ocean.
Remark: New record to Andaman and Niocbar
Islands
98
IGenus
: Ophiarthrum
38. Ophiarthrum pictum (Muller
and Troschel, 1842)
Description: Disc diameter 25 mm. Arms 6 times the
disc diameter in length. Disc is covered with thick skin.
Arm spines are blunt, twice as long as the segment with
dark rings or bands.
Colour: Disc patterned with grey, yellow and white.
Arms with dark longitudinal lines.
Habitat: Benthic, inshore.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: Red Sea, West Indian Ocean, East Indies,
North Australia and Philippines.
Ophiarthrum pictum (Muller & Troschel, 1842)
99
IGenus
: Ophiomastix
39. Ophiomastix annulosa (Lamarck, 1816)
Description: Disc scales with scattered blunt spines.
Ventral shields have concentric dark and light colouration.
Arm spines are claviform. Dorsal arm plates with light
coloured edges.
Ophiocoma scolopendrina (Lamarck, 1816)
Colour: Disc light red in colour. Arms are red
colour with white spines.
III
Habitat: Benthic, Inshore.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep. Elsewhere: Maldives, Ceylon, East
Indies, North Australia, Philippine, China, South Japan
and South Pacific Island.
100
Family: OPHIOTRICHIDAE
Genus : Macrophiothrix
40. Macrophiothrix propinqua (Lyman, 1861)
Description: Arm length up to 14 cm, disc diameter
upto 1 cm. Disc surface with or without scattered stumps.
Dorsal arm plates broader than long, distal side convex
or straight, laterodistal angles bent backwards, broadest
at about the middle of the plate. Distal edge of oral arm
plates straight. Arm spines as long as or slightly longer
than breadth of the segment. Colour variable, from evenly
dark bluish to banded orange and cream. Arms variegated
or uniformly blue with darker bands of 3-4 segments.
Colour: Disc black in colour. Arms are black with
white patches.
Habitat: It is found in rock crevices and among
coral.
Distribution: India: Andaman and Nicobar
Islands and Lakshadweep; Elsewhere: West Indian
Ocean, Mascarene Island, East Africa, Madagascar, Red
Sea, South East Arabia, Maldives, East Indies, North
Australia, Philippines, China, south Japan and South
Pacific Islands.
101
41. Macrophiothrix longipeda (Lamarck, 1816)
Description: Arm length 80 cm (about 20 times disc
diameter), disc diameter 1-4 cm. Disc scales are covered
by thorny stumps. Radial shields with rugose granule-like
stumps. Dorsal arm plates more than twice as broad as
long, distal edge with a straight median region, marked
laterodistal angles, broadest distally. Ventral arm plates
octogonal, distal edge straight. Up to ten thorny arm
spines, lowest one comb-like.
Colour: Disc colour blackish blue with dark blue
spots.
Macrophiothrix longipeda
-ventral
Macrophiothrix longipeda -dorsal
Habitat: Lower eulittoral, sublittoral, under coral
boulders or in burrow or a crevice.
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep Islands, Gulf of Mannar and West Bengal
coast; Elsewhere: Aldabra, Chagos, Eastern Mrica,
Madagascar, Mascarene Basin, Mauritius, Mozambique,
Red Sea, Seychelles, South Africa, Tanzania and West
Indian Ocean.
102
Family: OPHIODERMATIDAE Ljungman, 1867
Genus :Ophiarachna
42. Ophiarachna incrassata (Lamarck, 1816)
Description: Arms spines are 4-4 basally and are
long and flaring. Body colour is uniformly green, but it
appears yellow or buff in preservation and running along
the arms above and below. Interradial areas have black
ringed light spots forming a reticulation and with the
annulated arm spines.
Ophiarachna incrassata (Lamarck, 1816)
Colour: Green in colour.
Habitat: Large ophiuroid inhabits in coral reefs.
During the day time they hide below the coral slabs or in
creVIces.
Distribution: India: Andaman Islands; Elsewhere:
Eastern Mrica, Madagascar, Red Sea, Seychelles,
Tanzania and West Indian Ocean.
103
Class ECHINOIDEA Leske, 1778
104
Order : CIDAROIDA
Family: CIDAROIDAE
Genus : Prionocidaris
43. Prionocidaris verticillata (Lamarck, 1816)
Description: In the apical system the ocular plates
usually nearly all insert. Large globiferous Pedicellariae
usually with limb or frill of rods on the stalk close to
the head. Verticillate primary spines bear 3 or 4 thorny
whorls.
Colour: Uniformly greenish brown in colour.
Habitat: Benthic, inshore, continental shelf.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: West Indian Ocean,
Mascarene Island, East Africa, Madagascar, Bay of
Bengal, East Indies, North Australia, Philippine, China,
South Japan and South Pacific Island and Australia.
Prionocidaris verticillata (Lamarck, 1816)
105
Family : ARBACIIDAE
Genus : Prionocidaris
44. Arbacia punctulata (Lamarck, 1816)
Description:
Test diameter of
2 inches, with
total
diameter
reaching 4 inches.
Long and sharp
spmes generally
white in colour.
Tube feet are
InconSClplCOUS
and are generally
an olive colour.
Colour: Body
colour
ranges
Arbacia punctulata (Lamarck, 1816)
from black to
reddish brown and spines are white in colour, sometimes
it may brown.
Habitat: The speCIes inhabits in nearshore areas
including coral reefs, sea grasses beds, hard bottoms,
sand and shell areas. They are typically found from the
intertidal zone to depths of 250m but almost common at
53m.
Distribution: India: Andaman Islands; Elsewhere:
Capecod, Masachusetts, South to Cuba including Florida
and the Gulf of Mexico, Panamas, Barbados and French
Guiana.
Remark: New record to Indian waters reported from
Rutland Island, South Andaman.
106
Order : DIADEMATOIDA
Family: DIADEMATIDAE
Genus : Diadema
45. Diadema setosum (Leske, 1778)
Description: Globiferous pedicellariae and spines on
the buccal plates are absent. A red ring is present around
anus and white spot over each genital pore. This species
is commonly distributed in tropical Indo-Pacific region
and Indo-West Pacific Ocean. Tridentate pedicellariae are
Large in size and mostly form narrow blades meeting at
the tip.
Colour: Uniformly black in colour.
Habitat: Benthic, inshore, continental shelf.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Aldabra, Eastern Mrica,
Madagascar, Kenya, Mascarene Basin, Mauritius,
Mozambique, Red Sea, Seychelles, Tanzania and West
Indian Ocean.
Diadema setosum (Leske, 1778)
107
46. Diadema savignyi (Michelin, 1845)
Description: This sea urchin has a black anal cone
and no white on the body. Buccal plates do not bear any
spines. Pedicellariae which are leaf or spoon shaped and
is tapering to the rounded distal end. It is found in small
pockets in reefs or around rocks. It has a wide range
throughout Indo-west Pacific region.
Colour: Black in colour.
Habitat: It is found in the rocky shores.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Aldabra, Eastern Mrica,
Madagascar, Kenya, Mascarene Basin, Mauritius,
Mozambique, Red Sea, Seychelles, South Africa and West
Indian Ocean.
Diadema savignyi (Michelin, 1845)
108
IGenus
: Echinothrix
47. Echinothrix calamaris (Pallas, 1774)
Description: Naked test is green in colour. Ambulacra
distinctly bulged aborally. Inter ambulacral spines occur
with fine to broad transverse white bands. The banded
sea urchin should be avoided and certainly not touched
as its secondary spines are extremely fine and sharp and
capable of penetrating over the gloves. These spines are
highly venomous and cause painful wounds.
Colour: Body colour is purplish brown with white
strips like band.
Habitat: It is usually found in the slopes of coral reef
with rocky substances.
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep Islands and Gulf of Mannar; Elsewhere:
Aldabra, Chagos, Eastern Africa, Kenya, Madagascar,
Mascarene Basin, Mozambique, Red Sea, Seychelles,
Somalia, South Africa, Tanzania and West Indian Ocean.
Echinothrix calamaris (Pallas, 1774)
109
48. Echinothrix diadema (Linneaus, 1758)
Description: Spines are large and about 9 inch
long. Naked test is black in colour. Cavity of the primary
spines is very small. This species is commonly found in
Indian Ocean rather than Pacific reefs. The surface of the
primary spines has longitudinal ridges.
Colour: The body colour of the live speCImens IS
velvet-black or crimson red.
Habitat: The animal hides under ledges, and in caves
along the reef edges.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep Islands; Elsewhere: Maldives, Aldabra,
Chagos, Eastern Mrica, Madagascar, Kenya, Mascarene
Basin, Red Sea, Seychelles, Somalia and West Indian
Ocean.
Echinothrix diadema (Linneaus, 1758) - dorsal
110
Order : ECHINOIDA
Family : ECHINOMETRIDAE
Genus : Echinometra
49. Echinometra oblonga (de Blainville, 1969)
Description: Test is oval in shape. Primary spine is
often massive and either very long or truncated and very
short. If they are slender than the lower ones are more or
less flattened and some are angular. Eight or more porepairs per arc are present aborally.
Colour: Uniformly black in colour.
Habitat: Found on rocks, or on dead corals.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: Seychelles and Tanzania.
Echinometra oblonga (de Blainville, 1969)
111
50. Echinometra mathaei (de Blainville, 1825)
Description: Mathae's sea urchin is a cosmopolitan
rock boaring species that thrives in the rugged condition
of exposed intertidal situations. Spines of the sea urchin
are slender and acute. Long axis presents on the oval test
through ambulacra I and interambulacrum 3. This urchin
is herbivore, scraping algae from boulders and rubbles
close to its hollow at night and catching drifting algae.
Colour: Body colour may be brown, pink, green or
purple but the spines have usually white circle around the
base.
Habitat: It is common at the rocky shores
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep Islands and Gulf of Mannar; Elsewhere:
Western Indian Ocean, Mascarene Islands, East Mrica
and Madagascar, Red Sea, South-East Arabia, Maldives,
Ceylon, East Indies, Philippines, China, South Japan,
South Pacific Island, Hawaiian, Persian Gulf and North
Australia.
Echinometra mathaei (de Blainville, 1825)
112
IGenus
: Heterocentrotus
51. Heterocentrotus trigonarius (Lamarck, 1816)
Description: Primary spines are massive which
either rounded or are very thick but somewhat flattened
towards the tip. Ambital region contains 15-16 pore-pairs
in each arc. Primary ambulacral tubercles are gradually
decreasing in size aborally and their spines are short and
not truncated
Colour:
Uniformly mid brown.
Habitat:
shore areas.
Rocky
Distribution:
India:
Nicobar
Islands; Elsewhere:
Aldabra,
Eastern
Mrica,
Kenya,
Madagascar,
Mascarene
Basin,
Mauritius, Red Sea,
Seychelles,
Somalia
and Tanzania.
Heterocentrotus trigonarius (Lamarck, 1816)
113
Family: ASTRICL YPEIDAE
Genus : Echinodiscus
52. Echinodiscus auritus (Leske, 1778)
Description: Test is highly developed and oval III
shape. Lunules are two in number. Lunules open at the
distal end. This species has a bite marks at the posterior
margins of the test.
Colour: Uniformly brown in colour
Habitat: Sandy sea shore.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Mouth of Hugly River; Elsewhere:
Mascarene Island, East Africa, Maagascar, Red Sea,
South East Arabia, Persian Gulf, Ceylon, East Indies,
North Australia, Philippines, China and South Japan.
Echinodiscus auritus (Leske, 1778)
114
Order : TEMNOPLEUROIDA
Family: TEMNOPLEUROIDAE
Genus : Temnopleurus
53. Temnopleurus alexendri (Bell, 1884)
Description: Crenulations are conscipicous and
Pore pairs more or less distinctly in arcs of three. Spines
not banded but often greenish basally becoming purple
distantly. Globiferous pedicillaria does not bear lateral
teeth. The pores reach to the edge of the ambulacral areas.
Tubercles are absent at the outside of the pores.
Colour: Uniform in colour- green, purple or white
Habitat: This species inhabits in rocky substances
and found in shallow water depths.
Distribution: India: Tamil Nadu coast, Andaman
Islands; Elsewhere: East Indies and North Australia.
Temnopleurus alexendri (Bell, 1884)
115
IGenus
: Mespilia
54. Mespilia globulus (Linneaus, 1758)
Description: Ten conspicuous abruptly spineless
vertical areas on the test are covered with extremely
numerous globiferous pedicellariae. These pedicellariae
forms a dense carpet, their valves with the blades bearing
two or three lateral teeth each side.
Colour: Uniformly blue colour with reddish-brown
spmes.
Habitat: In warmer currents of shallow waters, on
hard bottom.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: Red Sea, West Indian Ocean, Red Sea, West
Indian Ocean, North Australia, Philippine, China, south
Japan, South Pacific Island and Australia.
Mespilia globulus (Linneaus, 1758)
116
IGenus
: Microcyphus
55. Microcyphus ceylanicus (Mortensen, 1925)
Description: Globiferous pedicellariae are less
conscipicously developed. The blade fairly narrow and
bearing not more than one lateral tooth each side. Tubercles
are indistinctly crenulate. Sutural pores are very small
and inconscipicous. Naked areas are conspicuous aborally.
Test patterned with olive-green.
Colour: Test is black in colour. Spines banded with
reddish brown and white.
Habitat: Inhabits in rocky substances and found in
shallow water depths.
Distribution: India: Andaman Islands; Elsewhere:
Ceylon.
Microcyphus ceylanicus (Mortensen, 1925)
117
IGenus
: Salmacis
56. Salmacis belli (Doderlein, 1902)
Description: Test is moderate in size that inflated in
profile with rounded ambitus. Apical disc is small, dicyclic,
with marginally positioned gonopores. Genital plates with
tubercles are developed around their inner edge forming
a perianal ring. Periproct is subcircular and central in
position. Spines are small, simple, without cortex.
Colour: Test black in colour, spines are whitish
brown in colour.
Habitat: Found in algal beds. It is a grazer and algae
eater.
Distribution: India: Nicobar Islands; Elsewhere:
East Indies and North Australia.
Remark: New record to Indian waters found in Great
Nicobar Islands.
Salmacis belli (Doderlein, 1902)
118
Family : TOXOPNEUSTIDAE
Genus : Tripneustus
57. Tripneustes gratilla (Linneaus, 1758)
Description: The pattern of the body spaces between
the spines is a reliable visual feature. The tube feet of the
large, tube-shaped, five body spaces and the five smaller
ones. Pore pairs are present horizontally on the arcs and
spaced to form three distinct vertical series in larger
speCImens.
Colour: It may be purple, black, brown or red and
even green.
Habitat: It is a grazer and algae eater and found in
the sandy reef slopes with muddy coverage.
Distribution: India: Andaman and Nicobar Islands,
Lakshadweep and Gulf of Mannar; Elsewhere: Aldabra,
Chagos, Eastern Mrica, Madagascar, Kenya, Mascarene
Basin, Mauritius, Mozambique, Red Sea, Seychelles,
Somalia, South Africa, Tanzania and West Indian Ocean.
Tripneustes gratilla (Linneaus, 1758)
119
Order : PHYMOSOMATOIDA
Family: STOMECHINIDAE (Clark and Rowe)
Genus : Stomopneustus
58. Stomopneustus variolaris (Lamarck, 1816)
Description: Complete bridges are absent across
the V-shaped space at the upper end of each of the
five
pyramids
forming
frame
of the Aristotle's
lantern.
Ambulacral
plates are doubly
compound so that
large
tubercle
correspondence
to three to six
arcs each of three
pore paIrs are
placed at the
ambitus.
Colo u r: Stomopneustus variolaris (Lamarck, 1816)
Uniformly black in colour.
Habitat: It is commonly found in rocky shore or
sometimes below the rubbles and dead coral patches.
Distribution: India: Andaman and Nicobar Island,
Lakshadweep, Gulf of Mannar and Andhra Pradesh Coast;
Elsewhere: Mascarene Island, East Africa, Madagascar,
South East Arabia, East Indies, Maldives, Ceylon, North
Australia, China, South Japan, South Pacific Island and
Australia.
120
Order : CLYPEASTEROIDA
Family : CLYPEASTERIDAE
Genus : Clypeaster
59. Clypeaster humilis (Leske, 1778)
Description: Interradial projections are not formed
around the peristome. Petals are generally smaller and
test fiat orally. Frontal petals are closed.
Colour: Uniformly white in colour sometimes may be
light yellowish in colour.
Habitat: Benthic, inshore, continental shelf.
Distribution: India: Gulf of Mannar, Kochi coast,
Lakshadweep and Andaman Islands; Elsewhere: Eastern
Mrica, Madagascar, Red Sea, South East Arabia, Persian
Gulf, Ceylon area, East Indies, North Australia and
Philippines.
Remark: Newly recorded in Andaman Islands.
Clypeaster humilis (Leske, 1778)
121
Class HOLOTHUROIDEA de Blainville, 1834
122
Order : ASPIDOCHIROTIDA Grube, 1840
Family: HOLOTHUROIDAE Ludwig, 1894
Genus : Actinopyga
60. Actinopyga mauritiana (Quoy and Gaimard, 1833)
Description: The shape is cylindrical with a fiat
underside; length up to 300 mm and live weight varies
from 0.5 to 1.0 kg. The tube feet are firmly attached to
rocks to prevent the animal from being washed away by
the waves.
Colour: Colour in living condition is brick red above
and white below.
Habitat: Usually found where the surf breaks on the
outside of the reef.
Distribution: India: Gulf of Mannar, Andaman and
Nicobar Islands and Lakshadweep; Elsewhere: Aldabra,
Chagos, Comores, Eastern Africa, Kenya, Madagascar,
Mascarene Basin, Mauritius, Mozambique, Red Sea,
Seychelles, Somalia, Tanzania and West Indian Ocean.
Actninopyga mauritiana (Quay and Gaimard, 1833)
123
61. Actinopyga miliaris
(Quoy and Gaimard, 1833)
Description: Length of the specimens ranged from
120 to 300 mm and the weight varied from 0.5 to 2 kg.
Massive cylindrical forms with rough surface. Anal teeth
very distinct.
Colour: Green and Black.
Habitat: Found mainly in waters less than 10 m
depth on pure sand. They also live on the reef fiats among
live corals and on algal beds.
Distribution: India: Gulf of Mannar, Andaman and
Nicobar Islands and Lakshadweep; Elsewhere: Aldabra,
Comores, Eastern Africa, Kenya, Madagascar, Mascarene
Basin, Mauritius, Mozambique Red Sea, Seychelles,
Tanzania and West Indian Ocean.
Actinopyga miliaris (Quay and Gaimard, 1833)
124
62. Actinopyga lecanora (Jaeger, 1833)
Description: The size of the specimens can be > 300
mm in length. The spicules are composed of small rods.
Colour: It is a light grey or brown sea cucumber, with
light speckled patches particularly around the somewhat
attenuated posterior end.
Habitat: It is often concealed amongst coral or rubble
of reefs.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: Djibouti (from synonym), East Mrica, Eastern
Mrica, Kenya, Madagascar, Mascarene Basin, Mauritius,
Mozambique, Somalia and Tanzania.
Actinopyga lecanora (Jaeger, 1833)
125
63. Actinopyga echinities (Jaeger, 1833)
Description: The body is wider in the middle and
tapers at both ends. The upper surface is wrinkled, often
with fine sand setteled over it. It attaches itself to rocks
with its tube feet.
Colour: Uniformly brown.
Habitat: It occurs at a depth of between 3-7m and
inhabits both hard (rocky area) and soft substratum
(sand).
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Lakshadweep. Elsewhere: Northern
Australian reefs, Western Indian Ocean, East Mrica and
Madagascar, South-East Arabia, Ceylon, East Indies,
Philippines, China, South Japan and South Pacific Island.
Actinopyga echinities (Jaeger, 1833)
126
Order : ASPIDOCHIROTIDA Grube, 1840
Family: HOLOTHUROIDAE Ludwig, 1894
Genus : Holothuria
64. Holothuria cinerascens (Brandt 1835)
Description: Ranges in length from 30 to 200 mm.
robust, sub-cylindrical with dorsal and ventral sides
sharply differentiated. Dorsal surface covered with
uniformly distributed numerous papillae. Ventrally beset
with crowded robust pedicels. Tentacles 20 in number,
large and sub-globose when fully expanded. Mouth
ventral, posterior end of the body blunt. Anus surrounded
by papillae. Body wall thick and fairly smooth to touch.
Pedicels more or less arranged in three rows. Papillae
of dissimilar sizes. In the living condition the tentacles,
though peltate, appear to be slightly arborescent. The
collar surrounding the tentacles is inconspicuous. The
calcareous ring is of the usual type. There were four
polian vesicles of dissimilar size. On the right side of the
mesentery there is a single stone canal. Cuvierian tubules
are well developed. Longitudinal muscle bands are
thin. Spicules are of two types, namely tables and rods.
Holothuria cinerascens (Brandt 1835)
127
Simple and finely granulated rdos are the characteristics
of the species. They are either straight of curved with
the extremetes often branched or coarser tubercles.
Occasionally, triradiate and tetraradiate rods occur with
three or four ends considerably branched. The length of
the rods varies from 0.10 mm to 0.30 mm. Table simple
with the annual disc varying in size from 0.042 mm to
0.060mm. Four large holes at the centre and four large
holes at the margin in each disc of the table. The crown of
the tables are subquadrate, being 0.045 mm in diameter.
Colour: Colour in living condition is reddish-brown
with some of the papillae and pedicels yellowish in
colour.
Habitat: This species IS found in rocky shores.
Individuals were often found attached firmly at the rock
edges by the three rows of pedicels on the ventral side.
Distribution: India: Gulf of Mannar, Andaman
and Nicobar Islands and Lakshadweep; Elsewhere:
Western Indian Ocean, Mascarene Islands, East Mrica
and Madagascar, Red Sea, South-East Arabia, Maldive
area, Ceylon area, East Indies, Philippines, China, South
Japan, South Pacific Island and Hawaii.
128
65. Holothuria (Thymiosycia) hilla (Lesson, 1830)
Description: Length from 50 mm to 200 mm. body
long and cylindrical with blunt ends, body wall soft. Dorsal
and ventral sides demarcated in the living condition.
Papillae sparsely arranged and have expanded bases.
Ventral side has numerous pedicels arranged in three
rows. A small space at the anterior end near the collar
Holothuria (Thymiosycia) hilla (Lesson, 1830)
is free from pedicels. Each band of pedicels with five or
six tube feet arranged side by side. Mouth surrounded by
20 inconspicuous papillae. Tentacles are small. There are
ten anal papillae. The calcareous ring is of the usual type
with the radials longer than the interradials. The right
respiratory tree is long, extending up to the anterior end,
while the left end one is shorter and joins the viscera. Two
polian vesicles and a stone canal are present. Spicules
consist of a table and buttons. Tables possess smooth
rounded discs. Four large holes corresponding to the
four spires in addition to about fifteen peripheral holes.
Spire of the tables consists of four pillars and one cross
beam which terminates in twelve or more teeth. Buttons
oval, smooth and symmetrical with three or four pairs of
holes. Holes at either end generally elongate. Length of
the button varies from 0.017 to 0.028 mm. diameter of the
table varies from 0.031 mm to 0.038 mm.
129
Holothuria (Thymiosycia) hilla (Lesson, 1830)
Colour: In living condition, the colour of the specimen
is chocolate brown and the large specimen is golden brown
with a circular pale area around the appendages.
Habitat: It is a fungitive species always found under
coral stones.
Distribution: India: Palk Bay, Gulf of Mannar,
Andaman and Nicobar Islands and Lakshadweep;
Elsewhere: Western Indian Ocean, Mascarene Islands,
East Mrica, Red Sea, South-East Arabia, Maldives,
Sri Lanka, Bay of Bengal, Persian Gulf, East Indies,
Philippines, Northern Australia, China, South Japan,
South Pacific Island and Hawaii.
130
66. Holothuria (Thymiosycia) impatiens
(Forskal, 1775)
Description: Length from 60 to 240 mm. Body bottle
shaped with a long neck. Superficially the body can not
be differentiated dorsally and ventrally. Mouth and anus
is terminal. Tentacles about 20 crowded around the small
mouth. Body surface covered by well developed papillae
placed on low, round warts which are conspicuous by
their lighter column than the rest of the body. Papillae
sca ttered fairly evenly over the surface and not in series.
Skin usually sandy to touch. Radial piece of the calcareous
ring much longer than interradials and projects forward.
The rounded margins have a deep cancavity. Interradial
pieces with short teeth. A single stone canal and one or two
olian vesicles. Cuvierian tubules occur in relatively large
branches. Respiratory trees slender with few branches.
Longitudinal muscle bands very thick. Spicules consist
of tables, buttons and supporting plates. Tables arranged
in a crowded Mannar with the edges of the disc touching
Holothuria (Thymiosycia) impatiens (Forskal, 1775)
131
or overlapping each other on the outer layer. Each table
consists of four upright rods and two cross beams. Spire
robust and top of the spire with a number ofteeth which are
level with the upper cross beam. Disc subquadrate usually
provided with nine holes forming three rows, central hole is
larger than the other two holes. Diameter of the table disc
is 0.10 mm. Spire 0.09 mm high and 0.05 mm in diameter.
Buttons oval in shape with mostly three pairs of holes,
smooth and with slightly undulating margins and obtuse
ends. Very rarely, with more than three holes on each
side. Length of the button varies from 0.084mm to 0.10
mm and breadth from 0.040 mm to 0.049 mm. supporting
rods slightly curved. Central portion dilated like a ring
and has invariably two holes. Tips slightly expanded with
one to four holes which are generally smaller than those
found at the middle. Sometimes the tip of the rods in the
papillae is not perforated.
Colour: In living condition the general body colour
is light brown with 4 to 5 dark brown transverse bands
on the dorsal side at the anterior end. A few dark brown
blotches are also found on the dorsal side on the rest of the
body. The ventral side is uniformly light brown with three
dark bands of the dorsal side extending to the ventral side
near the anterior end.
Habitat: It is often found under the dead coral
stones.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Lakshadweep; Elsewhere: Western
Indian Ocean, Mascarene Islands, East Africa, Red Sea,
Arabia, Maldives, Sri Lanka, Persian Gulf, East Indies,
Philippines, Northern Australia, China, South Japan,
South Pacific Island and Hawaii.
132
67. Holothuria atra (Jaeger, 1833)
Description: Length from 90-500 mm but known to
grow up to 600 mm. body elongate, sub-cylindrical and
capable of considerable extension. Posterior end is blunt.
Mouth is in the form of a transverse slit and surrounded
by a conspicuous papillose collar. There are 20 tentacles
Holothuria atra (Jaeger, 1833)
in double row. Pedicels numerous and crowded on the
ventral side. Papillae rather thicker than the pedicels
and sparsely arranged. Peristome rather thick, tough and
lethery in consistency. Anus terminal. The calcareous ring
is not large. The radial pieces extend rather forward than
interradials. Radials square shaped, the anterior edge of
each radial has a round incision while each interradial
piece has an anterior tooth. Posterior margin of the
interradial arched. There are four polian vesicle and 18
stone canals. The right respiratory tree extends forward
to the calcareous ring and is firmly attached to the body
wall and the left one which is shorter and connected with
Holothuria atra (Jaeger, 1833)
133
Holothuria atra (Jaeger, 1833)
the extensive rete mirabilis of the intestine. Curvierian
tubules absent. The spicules consist of tables and rosettes.
Tables numerous but not crowded. Each table posses a
smaller annular disc and a robust spire composed of four
rods and one cross beam. Disc diameter 0.055 mm and
commonly consist s of a simple ring with perforation at
the base of each rod. Cross beam nearer to the disc than to
the crown. Spire surround by the eight robust horizontal
and four equally strong, sharp large vertical teeth. Central
hole of the spire varies from 0.06 mm to 0.08 mm and the
breadth of the crown is about 0.06 mm. rosettes small and
vary in size from 0.019 to 0.045 mm. pedicels have well
developed terminal plates. A few bilaterally symmetrical
fenestrated plates are present close to the terminal plates
of the pedicels. The papillae contain slightly curved smooth
or spinose rods, mostly with enlarged fenestrated ends.
Colour: In living condition the colour is black or
vey dark brown or reddish-brown. The tentacles and the
peristomes are dark brown.
Habitat: It is always found fully exposed in shallow
water on sandy bottom.
Distribution: India: Andaman and Nicobar Islands,
Palk Bay, Gulf of Mannar and Lakshadweep; Elsewhere:
Western Indian Ocean, Mascarene Islands, East Mrica,
Red Sea, Arabia, Maldives, Sri Lanka, Persian Gulf, East
Indies, Philippines, Northern Australia, South Japan,
South Pacific Island and Hawaii.
134
68. Holothuria (Acanthotrapeza) pyxis
(Selenka, 1867)
Description: The length of the specimens varied from
270-450 mm. The body is tubular. The posterior region is
bulged and blunt with the anterior end narrow. A number
of projections are found on the dorsal side. Some of them
are 20 mm in length. They are not arranged in order. On
the ventral side, there are four bands of pedicels. In each
band there are five or six pedicels arranged side by side.
Radials large with a deep notch at the anterior end, the
posterior end straight. Interradials half the height of the
radials and have a concavity at the posterior margin. A
single stone canal and a single polian vesicles is present.
The spicules consist of only tables in the body wall.
Margins of the tables spiny with four large holes at the
centre and a number of small holes around the margin.
Table short and end in about 10 short spines. Height of
the tables is 0.04 mm and the diameter of the disc of the
tables is 0.05mm.
Colour: Dorsal side is brownish-pink with some
papillae blackish brown. The ventral side is light brown,
with spaces between the bands of the pedicels yellow.
Habitat: The
species is always
found under the
large stones which
are well fixed to
the ground.
Distribution:
India: Andaman
Islands; Elsewhere:
East Indies
Holothuria (Acanthotrapeza) pyxis
(Selenka, 1867)
135
69. Holothuria leucospilota (Brandt, 1835)
Description: Large and snake -like like forms with
leathery skin. The pedicels are large with well developed
sucking discs. The tentacles are 20 in number and are
ventrally placed. There is a well developed tentacles collar
with a fimbriated margin. In the calcareous ring each
radial is large and has a deep groove white the interradial
is a short, stump like projection. There is a single polian
vesicle and a single stone canal. The respiratory trees are
well developed. Cuvierian tubules are also well developed.
The spicules consist of an external layer of tables with a
complete or incomplete discs often reduced to four holes,
one at the base of each pillar. Spire is low and often
partly reduced, but when complete. It ends in a flattened
crown of eight or twelve teeth. Inner layer consist of
regular six holed buttons. The button may some times
to asymmetrical. Pedicels are with large end plates and
a few broad perforated plates with more or less slit like
holes. The length of buttons varies from 0.050mm to 0.063
Holothuria leucospilota (Brandt, 1835)
136
mm, and the breadth varies from 0.025 mm to 0.033 mm.
the height of the table is 0.042 mm and the diameter of
the disc varied from 0.037 mm to 0.054 mm.
Colour: The colour is reddish-brown in live condition
about looks black on contraction.
Habitat: The species has the peculiar habit of tucking
its posterior end under a stone. Some times can be seen
under coral stones.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Lakshadweep; Elsewhere: Western
Indian Ocean, Mascarene Islands, East Africa, Red Sea,
Arabia, Maldives, Sri Lanka, Persian Gulf, East Indies,
Philippines, Northern Australia, China, South Japan,
South Pacific Island and Hawaii.
137
70. Holothuria (Mertensiothuria) fuscocinerea
(Jaeger, 1833)
Description: The length of the specimen is 120
mm, the body is long and tubular with 20 large ventral
tentacles. Tentacle on ventral side are arranged in three,
though not distinct bands. The papillae on the dorsal side
are scattered. There is a single polian vesicle and large
stone canal. Cuvierian tubules are large. Usually button
has two narrow slit like holes and one or two pairs of
minute holes at each end. The length of the button varies
from 0.01 mm to 0.04 mm.
Colour: Colour in living condition is brownish, more
or less mottled, ventral side is pale gray.
Habitat: It is usually found sandy as well as rocky
areas.
Distribution: India: Andaman Islands; Elsewhere:
Sri Lanka, Philippines, Japan, Fiji, Samoa, Australia and
Celebes.
Holothuria (Mertensiothuria) {uscocinerea (Jaeger, 1833)
138
71. Holothuria (Metriatyla) scabra (Jaeger, 1833)
Description: The length of the species is 300-400 mm
the body is robust with both the ends blunt. The dorsal
side is convex and the ventral side is fiat. The skin is very
thick and slimy to touch. On the dorsal side, there are
many small papillae which are mainly scattered and often
inconspicuous. On the ventral sides the pedicels are densly
distributed without any arrangement. Each dark spot on
the ventral side represents one pedicel. There are two
polian vesicles and a single stone canal. The calcareous
ring is of the usual type. The left respiratory tree is much
larger than the right. The paired radial muscles are not
in firm contact with the body wall. The spicules consist
of tables and buttons. All the buttons are knobbed and
with holes and with irregular perforated plates.the tables
are short and the margins are not quite rounded. Each
table has a few to many holes. The tables are short with
a horizontal cross bar and a crown of spine at the top,
Holothuria (Metriatyla) scabra (Jaeger, 1833)
139
which are visible in lateral VIew. In the apical VIew, 8
outwardly pointed spines are seen. The spire consists of
four vertical bars which terminate in a few spines. There
is a tier of cross bars in the spire. Buttons are small and
generally have three pairs of holes. The pedicels have
small terminal plates.
Colour: Colour in the living condition is grey to black
on the dorsal side, and white ventrally.
Habitat: It is found in muddy-sandy regIOns and
prefer less saline waters.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Mannar and Palk Bay; Elsewhere: Mascarene
Islands, East Africa, Red Sea, Arabia, Maldives, Sri
Lanka, Persian Gulf, East Indies, Philippines, Northern
Australia, China, South Japan, South Pacific Island and
Hawaii.
140
72. Holothuria (Halodeima) edulis (Lesson, 1830)
Description: Length from 90 mm to 300 mm, body
elongate, narrow at the anterior end and blunt at the
posterior end. Minute papillae found on the dorsal side of
the body. Numerous pedicels present on the ventral side.
There are 20 medium-sized tentacles surrounded by a rim
of black papillae. Skin is smooth and thin. The inner wall
of the cloaca is black in colour. The calcareous ring is of
moderate size. There are 37 stone canals and one polian
vesicle. Both the right and left branches of the respiratory
trees are large and of equal size. Spicules consisit of tables
and buttons. Disc of the tables reduced to a ring which is
narrower than the top of the spire. There is a horizontal
beam in the middle of the spire. The top of the spire is
expanded and bears four blunt spines on each side which
can be seen only in the lateral view. Height of the tables
varies from 0.052 mm to 0.066 mm and diameter of the
spire varies from 0.037 mm to 0.043 mm. small button
present in the inner layer. The number of the holes varies
from 3 to 10 and most of them are incomplete. Length of
the button varies from 0.026 mm to 0.058 mm and the
Holothuria (Halodeima) edulis (Lesson, 1830)
141
Holothuria (Halodeima) edulis (Lesson, 1830)
breadth from 0.017 to 0.031 mm. long supporting rods
which have expanded ends and three to four holes are
present in the pedicels.
Colour: Body colour is bright rose pink which may be
obscured by varying degrees of black pigments. The black
colour is well marked on the dorsal side where it varies
from grey to intense black and at the side it is replaced by
pink. On the ventral side there is no black colour.
Habitat: It is usually found in shallow deapth.
Distribution: India: Andaman Islands and Gulf of
Mannar; Elsewhere: East Mrica, Red Sea and SouthEast Arabia.
142
73. Holothuria (Microthele) nobilis (Selenka, 1867)
Description: The length varied from 250mm to
400 mm. the body is tubular and massive in shape. Live
weight varies from 2 to 3 kg in fresh condition. Body
wall is 10-15 mm in thickness. Pedicels and papillae
are indistinguishable. Dorsal papillae are more thinly
scattered than the ventral pedicles. Anus is surrounded
by five calcified papillae. Calcareous ring is massive
with distinctly scalloped anterior margin. The radial and
interradials are squarish. Radial are twice the length of the
interradials. Tentacular ampullae are very large. Spicules
consist of tables and buttons. The tables are robust with
smooth discs and the spires terminate in 15-20 small
spines. The diameter of the table varies from 0.06 to 0.08
mm. the disc of the tables is either irregularly rounded or
square shaped. The inner layer has closely packed hollow
fenestrated ellipsoids which are 0.07 mm in length. They
have four rows of holes. A few simple knobbed buttons are
also present.
Colour: This species can be seen in two colour forms,
white and black.
Habitat: The species lies freely in the lagoon in the
adult stage and often covered with a coat of sand. Young
white forms live among the algae. The white form is found
in more than 3 m dea pth.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep, Elsewhere: Aldabra, Chagos, Comores,
East Mrica, Eastern
Mrica,
Kenya,
Madagascar,
Mascarene
Basin,
Mauritius, Red Sea,
Seychelles, Somalia,
South
Africa,
Tanzania and West
Indian Ocean.
Holothuria (Microthele) nobilis
(Selenka, 1867)
143
74. Holothuria coluber (Semper, 1868)
Description: It is a black sea cucumber with
white papillae and 20 yellow tentacles. It is a long (500600 mm) species with a firm, thick body wall, and a
tough tegument. Spicules include spinose cup and saucer'
tables and rosettes.
Colour: Uniformly black in colour.
Habitat: This species is found on reefs, usually with
its posterior wedged below rocks on reef fiats.
Distribution: India: First time reported from
Andaman Islands; Elsewhere: East Indies, North
Australia, Philippines and South Pacific Islands.
Remark: New record to Indian waters reported from
Andaman Islands
Holothuria coluber (Semper, 1868)
144
75. Holothuria (Mertensiothuria) pervicax
(Selenka, 1867)
Description: The length of the specimens ranges
from 70 mm to 120 mm. they are sub-cylindrical in shape.
The dorsal and the ventral sides are well differentiated. 0
n the ventral side there are a number of pedicels arranged
closely without any evidence of band formation. Tye
papillae are scattered on the dorsal side. The tentacles are
definitely ventral in position. Cuvierian tubules are thick.
Calcareous ring is ofthe usual type. There are a large single
polian vesicles and a single stone canal. Spicules consist
of incomplete and oblong rods with lateral projections
resembling narrow rosette. The disc of the table is usually
sub-circular. Each disc has a fairly big hole at the base of
each slender sire. Frequently supplientray holes are also
present. The edge of the disc is smooth. The diameter of
the discs varies from 0.03 mm to 0.05 mm. the spire has a
cross beam and is frequently incomplete and ends in four
simple teeth. In some cases, the spire is rudimentary and
Holothuria (Mertensiothuria) pervicax (Selenka, 1867)
145
the crowns have no transverse pieces. The rosette vary
in size from 0.023 mm to 0.069 mm in length. They are
irregular and smooth. The pedicels have well developed
plates, but in the papillae they are rudimentary. The
pedicels and papillae, in addition to long curved rods
with short irregular processes, have bilateral fenestrated
plates. These plates vary in length from 0.30 mm to 0.36
mm. Those plates which are neighborhood of the terminal
plates of the pedicels are formed by the branching and
joining of the lateral processes of the supporting rods.
Colour: In living condition the dorsal side IS
brown with 5 to 7 honey coloured transverse bands of
differentiated width. The ventral side is lighter, mottled
with white and light violet on a brown background. The
cloacal opening is surrounded by a dark violet ring with
some some portion of the inner cloacal wall of the same
colour.
Habitat: It is usually found in sandy areas.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Western Indian Ocean,
Mascarene Islands, East Africa, Red Sea, South East
Arabia, Maldive area, Sri Lanka, Persian Gulf, East
Indies, Philippines, Northern Australia, South Pacific
Islands and Hawaii.
146
76. Holothuria arenicola (Semper, 1868)
Description: Body is slender and vermiform. Mouth
is small and surrounded by tentacles ventrally. Pedicles
small and not conscipicous and arranged in three bands
ventrally. Anus is terminal in position and surrounded
by five groups of four to six short papillae. The spicules
consist of tables, buttons and supporting plates. Button
smooth and regular with six holes with edges regularly
indented between each pair of holes.
Colour: Body colour is white and on the dorsal side,
three pairs of reddish-brown spots are present.
Habitat: It inhabits in the sand, under the stones and
it is very difficult to take out the specimen completely.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: West Indies, Zanzibar,
Fiji, Tahiti, Galapagos and East Coast of Australia.
Holothuria arenicola (Semper, 1868)
147
I Genus
: Bohadschia
77. Bohadschia marmorata (Jaeger, 1833)
Description: Body short and thick with the lower
surface slightly flattened. It grows to a large size of
400mm.
Colour: Colour highly variable. Colour is golden
brown with small brown dots. Sometimes the colour is
yellowish brown with black spots.
Habitat: Occurs on coarse coarl sand at depths 2-6 m.
it is also seen in the intertidal region covered by a coating
of fine mud.
Distribution: India: Gulf of Mannar, Andaman and
Nicobar Islands and Lakshadweep; Elsewhere: Aldabra,
East Africa, Eastern Africa and Madagascar, Kenya,
Madagascar, Mascarene Basin, Mozambique, Red Sea and
Seychelles.
Bohadschia marmorata (Jaeger, 1833)
148
78. Bohadschia argus (Jaeger, 1833)
Description: Body is cylindrical with very smooth
surface. At the slightest disturbance the sticky threads are
thrown out. It grows to a large size of 600 mm in length.
Live weight is 1-2 kg. Distinct eye like spots are found all
over the body which are encircled with light yellow, white
grey colours. The eye spots are situated at a particular
angle.
Colour: Colour in living condition is brown or black.
Habitat: Occurs on coarse sand in 2-6 m depth. A few
pieces of shell and coarse sand usually stick to body.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Western Indian Ocean,
Ceylon, East Indies, Philippines, Northern Australia,
South Pacific Islands, China and South Japan.
Bohadschia argus (Jaeger, 1833)
149
79. Bohadschia graeffei (Semper, 1868)
Description: The length of the species is variable
from 15 cm to 18 cm. Underside is grainy. Upper side
of the species is rough in structural conformation. Body
thickness is 4mm. Papillae are low. Spicules resemble a
somewhat tack-like bundle of spinose rods described as
racquet-like.
Bohadschia graeffei (Semper, 1868)
Bohadschia graeffei (Semper, 1868)
150
Bohadschia graeffei (Semper, 1868)
Colour:
speckling.
Body
colour:
pale
cream
with
brown
Habitat: Reef slopes, close to the coast. Abundant on
bottom of mixed corals and calcareous red algae. Depth
range- 0-25 m but it is mostly found in 0-10 m.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep. Elsewhere: Red Sea.
151
Order : ASPIDOCHIROTIDA Grube, 1840
Family: LABIDODEMATIDAE
Genus : Labidodemas
80. Labidodemas semperianum (Selenka, 1867)
Description: The length of the species ranges between
85-250 mm. Spicules are spinose tables and sometimes
buttons and rods.
Colour: It is a white, almost translucent sea cucumber,
with yellow tube feet and a dark posterior end.
Habitat: This species can be found buried in sand
below rocks of the outer coral reef fiat.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: East Indies, Philippines, Northern Australia
and South Pacific Island.
Labidodemas semperianum (Selenka, 1867)
152
Order : DENDROCHIROTIDA Grube, 1840
Family: CUCUMARIIDAE Ludwig, 1894
Genus : Stolus
81. Stolus buccalis (Stimpson, 1856)
Description: It is small in size ranges from 70-110
mm, firm and cylindrical species, with a smooth tegument.
Spicules are small nodular buttons. It is found in coastal
habitats south to temperate regions, below stones on
intertidal fiats. It is mostly distributed in tropical, Indowest Pacific Ocean.
Colour: Specimen is dark purple- black in colour with
red colour tentacles.
Habitat: Usually found concealed under rock or
narrow creVIces
III
Distribution: India: Gulf of Mannar, Andhra
Pradesh, Palk Bay, Gulf of Kachchh and Andaman and
Nicobar Islands; Elsewhere: East Africa and Madagascar,
South East Arabia, Persian Gulf, West Indies, Ceylon,
East Indies, North Australia, Philippines, China and
South Japan.
Remark: New record to Andaman Islands reported
from Burmanella, South Andaman.
Stolus buccalis (Stimpson, 1856)
153
Family: STICHOPODIDAE Haeckel, 1896
Genus : Thelenota
82. Thelenota ananas (Jaeger, 1833)
Description: This species grows to a massive size
700 mm in length. The weight of a live specimen varies
from 3 to 6 kg. It is seen in massive form with numerous
large pointed teats in groups of two or three all over the
upper surface.
Thelenota ananas (Jaeger, 1833)
Colour: Colour of the sample in live condition is
reddish-orange on the upper side. Tube feet are bright
orange colour on the lower surface.
Habitat: It is found on clean sandy bottom at a depth
of 2-30 m.
Distribution: India: Andaman and Nicobar
Islands and Lakshadweep; Elsewhere: Maldives, East
Indies, North
Australia,
China, South
Japan
and
South Pacific
islands.
Thelenota ananas
(Jaeger, 1833)
154
セ@
..
セ
M
セM
セM
セ@
I Genus
: Stichopus
83. Stichopus horrens (Selenka, 1867)
Description: It is a medium-sized species (to 300
mm) with a smooth tegument but large and irregular
papillae. The big tubercles and irregular body form give
an "irregular, soft and almost repulsive" appearance.
Spicules are tables and large "C" bodies.
Colour: This species is a variable, grey to greenlblack
sea cucumber. It is often variegated with dark patches.
Habitat: S. horrens may be found on reefs, below
rocks on fiats.
Distribution: India: Andaman and Nicobar Islands;
Elsewhere: East Indies, North Australia, China, South
Japan, South pacific Islands, Philippines and Hawaii.
Stichopus horrens (Selenka, 1867)
155
84. Stichopus chloronotus (Brandt, 1835)
Description: The maximum length of the body is 300
mm. Body quadrangular in shape with four rows of large
finger like processes.
Colour: Green in colour. The tip of the finger like
processes is orange in colour. The tentacles and the tube
feet are ash-coloured and the stalks of tentacles are
white.
Habitat: Usually found only beyond low water mark.
Distribution: India: Andaman and Nicobar
Islands, Lakshadweep and Gulf of Mannar; Elsewhere:
Western Indian Ocean, Mascarene Islands, East Mrica,
Madagascar, Red Sea, Maldives, Ceylon, Bay of Bengal,
Persian Gulf, East Indies, Philippines, Northern Australia,
China, South Japan, South Pacific Islands and Hawaii.
Stichopus chloronotus (Brandt, 1835)
156
85. Stichopus hermanni (Semper, 1868)
Description: The maximum length of the species is
900 mm. Body massive and loaf like in appearance. The
tubercles are prominent.
Stichopus hermanni (Semper, 1868)
Colour: In living condition it is dark yellow with
irregular brown patches and pink tube feet.
Habitat: Occurs on algal beds and clean sand bottoms
between depths 3-30 m.
Distribution: India: Gulf of Mannar, Andaman and
Nicobar Islands, Lakshadweep and Palk Bay; Elsewhere:
Aldabra, East Mrica, Eastern Mrica, Kenya, Madagascar,
Mascarene Basin, Mozambique, Red Sea, Seychelles,
South Africa, Tanzania and West Indian Ocean.
Stichopus hermanni (Semper, 1868)
157
86. Stichopus vastus (Sluiter, 1887)
Description: This sea cucumber can grow up to a
length of 500 mm and reach maximum weight of 2.5 kg.
The body is wider in the middle, tapering at both ends
and covered with a bright smooth skin. The upper surface
is uniformly brown with zigzag bands at all over the body.
Body spicules consists of tables, branched rods and '8' and
'C' shaped rods.
Colour: Uniformly brown
patches.
III
colour with black
Habitat: It is commonly found in the depth of I-3m.
It occurs in sand and rocky substrata.
Distribution: India: Andaman Islands; Elsewhere:
North Australia and Maldives.
Stichopus vastus (Sluiter, 1887)
158
Order : APODIDA Brandt, 1835
Family : SYNAPTIDAE Ostergren, 1898
Genus : Synapta
87. Synapta maculata (Chamisso
and Eysenhardt, 1821)
Description: The structure of the body is like a
snake. It can reach more than 2 m in length. Presence of
15 tentacles.
Colour: It is tan to brown in colour with black
margins. The whole body surface has small white rings
which are closely arranged.
Habitat: It is usually found on the reef fiat.
Distribution: India: Gulf of Mannar, Andaman and
Nicrobar Islands and Lakshadweep; Elsewhere: Western
Indian Ocean, Mascarene Islands, East Africa, Red Sea,
Madagascar, Maldives, South East Arabia, Persian Gulf,
East Indies, Philippines, Northern Australia, China,
South Japan and South Pacific Island.
159
I Genus
: Euapta
88. Euapta godeffroyi (Semper, 1868)
Description: It is a medium to large species (to 400
mm in length), with 15 tentacles, each with numerous
pairs of digits. Its spicules are anchors with tiny knobs on
the vertex and plates with large posterior holes.
Colour: Body colour is creamy white with grey, with
green or brown longitudinal stripes.
Habitat: It is found in intertidal pool, in sand,
beneath stones. This species is found on reefs, concealed
among rubble of reef fiat or slope.
Distribution: India: Andaman and Nicobar Islands
and Lakshadweep; Elsewhere: Mascarene Island, Red
Sea, Maldives, East Indies, North Australia, Philippines
and South Pacific Island.
Euapta godeffroyi (Semper, 1868)
160
Order : MOLPADIIDA Miiller, 1850
Family: CAUDINIDAE Heding, 1931
Genus : Acaudina
89. Acaudina molpadioides (Semper, 1868)
Descripition: This sea cucumber is loaf shaped and
its maximum length is about 300mm. Body surface is
thick and covered with a fine coat of sand and is brown
coloured with black patches.
Colour: Uniformly brown in colour.
Habitat: This species is commonly found in shallow
reef environments. Depth range 2-15m.
Distribution: India: Andaman and Nicobar Islands,
Gulf of Kachchh, Andhra Pradesh, West Bengal and Gulf
of Mannar. Elsewhere: Northern Australia.
Acaudina molpadioides (Semper, 1868)
161
8. CHECKLIST OF ECHINODERMS
OF INDIA
Class: CRINOIDEA
Order: COMATULIDA
Family : Comasteridae
1. Capillaster mariae (AH. Clark, 1907)
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
Capillaster multiradiatus (Linneaus, 1758)
Comanthina nobilis (P.H. Carpanter, 1884)
Comanthina schlegeli (P.H. Carpanter, 1881)
Comanthus parviciirus (Muller, 1841)
Comanthus samoan us (AH. Clark, 1909)
Comanthus wahlbergi (Muller, 1843)
Comaster gracilis (Hartlub, 1890)
Comaster multibrachiata (P.H. Carpanter, 1888)
Comaster multifidus (J. Muller, 1841)
Comaster parvus (AH. Clark, 1909)
Comatella maculata (P.H. Carpanter, 1888)
Comatella nigra (P.H. Carpanter, 1876)
Comatella stelligera (P.H. Carpanter, 1880)
Comatula brevicirra (Bell, 1834)
Comatula micraster (AH. Clark, 1909)
Comatula pectinia (Linneaus, 1758)
Oxycomanthus bennetti (Muller, 1841)
Family: Himerometridae
19. Amphimetra molleri (AH. Clark, 1908)
20. Craspedometra acuticirra (P.H. Carpanter, 1882)
21. Craspedometra anceps (P.H. Carpanter, 1888)
22. Heterometra bengalensis (Hartlaub, 1890)
162
23.
24.
25.
26.
27.
28.
Heterometra compta (A.H. Clark, 1909)
Heterometra philiberti (Muller, 1841)
Heterometra reynaudi (Muller, 1846)
Heterometra crenulata (P.H. Carpenter, 1882)
Himerometra magnipinna (A.H. Clark, 1908)
Himerometra robustipinna (P.H. Carpanter, 1912)
Family : Mariametridae
29.
30.
31.
32.
Dichrometra ciliata (A.H. Clark, 1912)
Dichrometra protectus (P.H. Carpanter, 1879)
Lamprometra palmata (Muller, 1841)
Selenemetra aranea (A.H. Clark,1909)
Family : Stephanometridae
33. Stephanometra coronata (A.H. Clark, 1909)
34. Stephanometra indica (Smith, 1876)
35. Stephanometra monacantha (Hartlaub, 1890)
Family: Colobometridae
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
Cenometra herdmani (A.H. Clark, 1909)
Cenometra bella (Hartlaub, 1890)
Cenometra emendatrix (Bell, 1892)
Colobometra brevicirra (A.H. Clark, 1912)
Colobometra discolor (A.H. Clark, 1909)
Cotylometra gracilicirra (A.H. Clark, 1908)
Decametra brevicirra (A.H. Clark, 1912)
Decametra moebiusi (A.H. Clark, 1911)
Iconometra inter media (A.H. Clark, 1912)
Oligometra imbricata (A.H. Clark, 1909)
Oligometra serripinna (P.H. Carpanter, 1881)
163
Family : Pontiopmetridae
47. Pontiometra andersoni (P.H. Carpanter, 1889)
Family : Antedonidae
48. Andrometra indica (AH. Clark, 1909)
49. Euantedon sp.
50. Dorometra nana (Hartlaub, 1890)
51. Mastigometra micropoda (AH. Clark, 1909)
52.
53.
54.
55.
56.
Psathyrometra inusitata (AH. Clark, 1912)
Psathyrometra mira (AH. Clark, 1909)
Sarametra nicrobarica (AH. Clark, 1929)
Trichometra plana (AH. Clark, 1912)
Trichometra obscura (AH. Clark, 1909)
Family : Tropiometridae
57. Tropiometra carinata (Lamarck, 1816)
Family : Calometridae
58. Neometra spinosissima (AH. Clark, 1909)
Family : Thalassometridae
59.
60.
61.
62.
63.
Crotalometra rustica (AH. Clark, 1909)
Crotalometra sentifera (AH. Clark, 1909)
Stiremetra carinifera (AH. Clark, 1912)
Thalassometra peripolos (AH. Clark, 1929)
Thalassometra sp.
Family : Eudiometridae
64. Eudiocrinus minor (AH. Clark, 1919)
65. Eudiocirnus ornatus (AH. Clark, 1919)
164
Family : Zygometridae
66. Zygometra andromeda (AH.Clark, 1912)
Family: Chartiometridae
67.
68.
69.
70.
Glyptometra crassa (AH.Clark, 1912)
Glyptometra invenusta (AH. Clark, 1909)
Glyptometra macilenta (AH. Clark, 1909)
Perissometra occidentalis (AH. Clark, 1929)
Family : Pentametrocrinidae
71. Pentametrocrinus varians (P.H. Carpanter, 1882)
72. Decametrocrinus sp.
73. Thaumatocrinus investigatoris (AH. Clark, 1912)
Order: BOURGUETICRINIDA
Family : Bourgueticrinidae
74. Bathycirrus woodmasoni (AH. Clark, 1909)
Family: Hintacrinitidae
75. Comastrocinus ornatus (AH. Clark, 1909)
76. Comastrocrinus springeri (AH. Clark, 1909)
Class: ASTEROIDEA
Order: P AXILLOSIDA
Family: Luidiidae
77.
78.
79.
80.
81.
82.
Luidia denudata (Koehler, 1910)
Luidia hardwicki (Gray, 1840)
Luidia integra (Koehler, 1910)
Luidia limbata (Sladen, 1889)
Luidia maculata (Muller and Troschel, 1842)
Luidia savignyi (Audouin, 1826)
165
Family : Astropectinidae
83. Astropecten bengalensis (Doderlein, 1917)
84. Astropecten euryacanthus (Lutken, 1805)
85. Astropecten greigi (Koehler, 1909)
86. Astropecten hemipechi (Muller and Troschel, 1841)
87. Astropecten indicus (Doderlein, 1888)
88. Astropecten inutilis (Koehler, 1910)
89. Astropecten monacanthus (Sladen, 1883)
90. Astropecten polycanthus (Muller and troschel, 1841)
91. Astropecten tamilicus (Doderlein, 1888)
92. Astropecten vappa (Muller and Troschel, 1841)
93. Astropecten sp.
94. Craspidaster hesperus (Muller and Troschel, 1840)
95. Dipsacaster pentagonalis (Alcock, 1894)
96. Dipsacaster sladeni (Alcock, 1893)
97. Dytaster insignis (Perrier, 1884)
98. Persephonaster coelochiles (Alcock, 1893)
99. Persephonaster croceus (Wood-Mason and Alcock,
1891)
100. Persephonaster gracilis (Sladen, 1889)
101. Persephonaster rhodopeplus (Wood-Mason
Alcock, 1891)
102. Persephonaster roulei (Koehler, 1909)
103. Psilaster agassizi (Koehler, 1909)
Family: Porcellanasteridae
104. Abyssaster tara (Wood-Mason and Alcock, 1891)
105. Porcellanaster caulifer (Sladen, 1883)
106. Porcellanaster ceruleus (Wyville Thomson, 1877)
107. Sidonaster vaneyi (Koehler, 1909)
166
and
108. Stryracaster armatus (Sladen, 1883)
109. Styracaster caroli (Ludwig, 1907)
110. Styracaster clavipes (Wood-Mason and Alcock, 1891)
Family : Goniopectinidae
111. Goniopecten sp.
Order: NOTOMYOTIDA
Family : Benthopectinidae
112.
113.
114.
115.
116.
117.
118.
119.
Benthopecten huddlestoni (Alcock, 1893)
Benthopecten indicus (Koehler, 1909)
Benthopecten semisquamatus (Sladen, 1889)
Benthopecten violaceus (Alcock, 1893)
Cheriaster pilosus (Alcock, 1893)
Cheriaster synderi (Fisher, 1906)
Cheriaster cribellum (Alcock, 1893)
Pectinaster mimicus (Sladen, 1889)
Order: VALVATIDA
Family: Chaetasteridae
120. Chaetaster vestitus (Koehler, 1910)
121. Chaetaster sp.
Family: Asterinidae
122. Anserpoda ludovici (Alcock, 1893)
123. Anserpoda pellucida (Alcock, 1893)
124. Asterina burtoni (Gray, 1840)
125. Asterina cepheus (Muller and troschel, 1842)
126. Asterina coronata (Von Martens, 1866)
127. Asterina lorioli (Koehler, 1897)
128. Asterina sarasini (de Loriol, 1897)
167
129.
130.
131.
132.
133.
134.
135.
Disasterina leptalacantha (H.L.Clark, 1916)
Disasterina spinosa (Koehler, 1910)
Disasterina spinulifera (H.L. Clark, 1938)
Paranepanthia brachiata (Koehler, 1910)
Patiriella pseudoexigua (Dartnall, 1910)
Tegulaster ceylanica (Doderlein, 1888)
Tegulaster emburyi (Livingstone, 1933)
Family : Archasteridae
136. Archaster angualtus (Muller and Troschel, 1842)
137. Archaster typicus (Muller and Troschel, 1840)
Family : Goniasteridae
138. Anthenoides sarissa (Alcock, 1893)
139. Astroceramus fisheri (Koehler, 1909)
140. Calliaster childreni (Gray, 1840)
141. Calliaster mamillifera (Alcock, 1894)
142. Ceramaster cueneti (Koehler, 1909)
143. Ceramaster mortenseni (Koehler, 1909)
144. Circeaster magdalenae (koehler, 1909)
145. Circeaster marcelii (Koehler, 1909)
146. Johannaster superbus (Koehler, 1909)
147. Lithosoma pentaphylla (Alcock, 1893)
148. Mediaster arcuatus (Salden, 1889)
149. Milteliphaster woodmasoni (Alcock, 1893)
150. Nymphaster moebii (Studer, 1884)
151. Nympahster nora (Alcock, 1893)
152. Ogmaster capella (Muller and Troschel, 1842)
153. Paragonaster tenuiradiis (Alcock, 1893)
154. Pentagonaster intermedius (Alcock, 1893)
168
155.
156.
157.
158.
159.
160.
161.
Pentagonaster pulvinus (Alcock, 1893)
Pentagonaster ctenipes (Sladen, 1891)
Pentagonaster sp.
Plinthaster doderleini (Koehler, 1909)
Plinthaster investigators (Alcock, 1893)
Pseudarchaster jordani (Fisher, 1906»
Pseudarchaster mozaicus(Wood Mason and Alcock,
1891)
162.
163.
164.
165.
166.
Pseudarchaster roseus (Alcock, 1891)
Rosaster confinis (Koehler, 1910)
Rosaster florifer (Alcock, 1893)
Stiraster tubercualtus (H.L.Clark, 1915)
Stellaster childreni (Gray, 1805)
Family : Asterodiscididae
167. Asterodiscides elegans (Gray, 1847)
Family : Oreasteridae
168. Anthenea pentagonalua (Lamarck, 1816)
169. Anthenea tuberculosa (Gray, 1847)
170. Choriaster granulatus (Lutken, 1869)
171. Culcita novaguineae (Muller and Troschel, 1841)
172. Culcita schmideliana (Retzius, 1805)
173. Goniodiscaster forficulatus (Perrier, 1875)
174. Goniodiscaster scaber (Mobius, 1859)
175. Goniodiscaster vallei (Koehler, 1910)
176. Halityle regularis (Fisher, 1913)
177. Pentaceraster affinis (Muller and troschel, 1841)
178. Pentaceraster gracilis (Lutken, 1871)
179. Pentaceraster multispinus (Von Martens, 1866)
169
180. Pentaceraster regulus (Muller and Troschel, 1842)
181. Pentaceraster westermanni (Lutken, 1871)
182. Poraster superbus (Mobius, 1859)
183. Protoreaster lincki (de Blainvillae, 1830)
184. Protoreaster nodosus (Linneaus, 1758)
Family : Asteropsidae
185. Asteropsis carinifera (Lamarck, 1816)
186. Valvaster striatus (Lamarck, 1816)
Family : Acanthasteridae
187. Acanthaster planci (Linneaus, 1758)
Family : Ophidiasteridae
188. Andora faouzii (Macan, 1938)
189. Certonardoasemiregularis (Muller and Troschel, 1842)
190. Cistina columbiae (Gray, 1840)
191. Dactylaster cylindricus (Lamarck, 1816)
192.
193.
194.
195.
196.
197.
198.
199.
200.
201.
202.
203.
170
Fromia armata (Koehler, 1910)
Fromia indica (Perrier, 1869)
Fromia milleporella (Lamarck, 1816)
Fromia monilis (Perrier, 1869)
Gomophia egyptiaca (Gray, 1840)
Heteronardoa carinata (Koehler, 1910)
Leiaster glaber (Peters, 1852)
Leiaster leachi (Gray, 1840)
Linckia guildingi (Gray, 1840)
Linckia laevigata (Linneaus, 1758)
Linckia multifora (Lamarck, 1816)
Nardoa frianti (Koehler, 1910)
204. Nardoa glatheae (Lutkan, 1865)
205. Nardoa lemonnieri (Koehler, 1910)
206. Nardoa novacaledoniae (Perrier, 1875)
207. Nardoa sp.
208. Neoferdina offreti (Koehler, 1910)
209. Ophidiaster armatus (Koehler, 1910)
210. Ophidiaster hemprichi (Muller and Troschel, 1842)
211. Paraferdina laccadivensis (James, 1976)
212. Paraferina sohariae (Marsh and Price, 1991)
213. Tamaria fusca (Gray, 1840)
214. Tamaria dubiosa (Koehler, 1910)
215. Tamaria hirsuta (Koehler, 1910)
216. Tamaria megaloplax (Bell, 1884)
Order: VELATIDA
Family : Pterasteridae
217. Eureaster cibrosus (von Martens, 1867)
218. Hymenaster alcocki (Koehler, 1909)
219. Hymenaster nobilis (Wyville Thompson, 1876)
220. Marsipaster hirsutus (Wood Mason and Alcock, 1891)
Order: SPINULOSIDA
Family : Echinasteridae
221. Dictyaster xenophilus (Wood Mason and Alcock, 1891)
222. Dictyaster woodmasoni (Alcock, 1893)
223. Echinaster callosus (von Marenzeller, 1895)
224. Echinaster luzonicus (Gray, 1840)
225. Echinaster purpureus (Gray, 1840)
226. Henricia mutans (Koehler, 1893)
171
Family : Metrodiridae
227. Metrodira subulata (Gray, 1840)
Order: FORCIPULATIDA
Family : Zoroasteridae
228. Cnemidaster squameus (Alcock, 1893)
229. Zoroaster adami (Koehler, 1909)
230. Zoroaster alfredi (Alcock, 1893)
231. Zoroaster angulatus (Alcock, 1893)
232. Zoroaster barathri (Alcock, 1893)
233. Zoroaster carinatus (Alcock, 1893)
234. Zoroaster gilesii(Alcock, 1893)
235. Zoroaster planus (Alcock, 1893)
236. Zoroaster zea (Alcock, 1893)
237. Zoroaster sp.
Family: Pedicellaster
238. Pedicellaster atratus (Alcock, 1893)
Family: Asteriidae
239. Scleraasterias mazophora (Wood-Mason and Alcock,
1891)
240. Sclerasterias nitida (Koehler, 1910)
Order: BRISINGIDA
Family: Brisingidae
241. Brisinga andamanica (Wood-Mason and Alcock, 1891)
242. Brisinga bengalensis(Wood-Masona and Alcock, 1891)
243. Brisinga gunni (Alcock, 1893)
244. Brisinga insularum (Wood-Masona and Alcock, 1891)
172
245. Brisinga panopla (Fisher, 1906)
246. Brisinga parallela (Koehler, 1909)
247. Stegnobrisinga gracilis (Koehler, 1909)
Family : Freyellidae
248. Freyastera benthphila (Sladen, 1889)
249. Freyastera tuberculata (Sladen, 1889)
Class: OPHIUROIDEA
Order: PHRYNOPHIURIDA
Family : Ophiomyxidae
250. Ophiomyxa australis (Lutken, 1869)
251. Ophiophrixus confinis (Koehler, 1922)
Family : Asteronychidae
252. Asteronyx loveni (Muller and Troschel, 1842)
Family : Euryalidae
253. Asteromorpha fosculus (Alcock, 1893)
254. Trichaster acanthifer (Doderlein, 1911)
Family : Gorgonocephalidae
255. Astroboa clavata (lyman, 1861)
256. Astroba nuda (Lyman, 1874)
257. Astrocladus exiguus (Lamarck, 1816)
258. Astrothrombus vacors (Koehler, 1904)
Family : Asteroschematidae
259. Asterschema subfastosum (Doderlein, 1930)
260. Ophiocreas sibogae (Koehler, 1904)
261. Ophiocreas sp.
173
Order: OPHIURIDA
Family : Ophi uridae
262. Ophioelegans cincta (Muller and Troschel, 1842)
263. Ophiolepis superba (H.L. Clark, 1842)
264. Ophiolypus granulatus (Koehler, 1897)
265. Ophimusium elegans (Koehler, 1897)
266. Ophimusium familiare (Koehler, 1897)
267. Ophimusium fimbriatum (Koehler, 1927)
268. Ophimusium lymani (Wyville Thompson, 1873)
269. Ophimusium relictum (Koehler, 1927)
270. Ophimusium scalare (Lyman, 1878)
271. Ophimusium simplex (Lyman, 1878)
272. Ophimusium validum (Ljungman, 1871)
273. Ophioplocus imbricatus (Muller and Troschel, 1841)
274. Ophiosphalma elegans (Koehler, 1897)
275. Ophiosphalma planum (Lyman, 1878)
276. Ophioteichus nodosa (Duncan, 1887)
277. Ophiotrochus panniculus (Lyman, 1878)
278. Ophiozonella bispinosa (Koehler, 1897)
279. Ophiozonella molesta (Koehler, 1904)
280. Ophiernus adspersus adspersus (Lyman, 1878)
281. Ophioleuce seminudum (Koehler, 1904)
282. Ophiopallas paradoxa (Koehler, 1904)
283. Ophiostratus bispinosus (Koehler, 1897)
284. Amphiophiura ornata (Lyman, 1878)
285. Amphiophiura paupera(Koehler, 1897)
286. Amphiophiura radiata (Lyman, 1878)
287. Amphiophiura sculptilis (Lyman, 1878)
288. Amphiophiura sordida (Koehler, 1897)
289. Amphiophiura stellata (Studer, 1882)
174
290. Homalophiura inflata (Koehler, 1897)
291. Ophiomastus tumidus (Koehler, 1897)
292.
293.
294.
295.
Ophiopyrgus alcocki (Koehler, 1897)
Ophiotypa simplex (Koehler, 1897)
Ophiura aequalis (Lyman, 1878)
Ophiura flagellata (Lyman, 1878)
296.
297.
298.
299.
Ophiura
Ophiura
Ophiura
Ophiura
forbesi (Duncan, 1879)
irrorata (Lyman, 1878)
kinbergi (Ljungman, 1867)
undulata (Lyman, 1878)
Family: Ophiocomidae
300. Ophiarthrum elegans (Peters, 1851)
301. Ophiarthrum pictum (Muller and Troschel, 1842)
302. Ophiocoma anglyptica (Ely, 1944)
303.
304.
305.
306.
Ophiocoma
Ophiocoam
Ophiocoma
Ophiocoma
brvipes (Peters, 1851)
dentata (Muller and Troschel, 1842)
doderleini (de Loriol, 1890)
erinaceus (Muller and Troschel, 1842)
307.
308.
309.
310.
Ophiocoma lubrica (Koehler, 1898)
Ophiocoma pica (Muller and troschel, 1842)
Ophiocoma pusilla (Brock, 1888)
Ophiocoma scolopendrina (Lamarck, 1816)
311.
312.
313.
314.
Ophiocoma valenciae (Muller and Troschel, 1842)
Ophiocomella sexradia (Duncan, 1887)
Ophiomastix annulosa (Lamarck, 1816)
Ophiopsila pantherina (Koehler, 1898)
Family: Ophionereidae
315. Ophiochiton ambulator (Koehler, 1897)
175
316.
317.
318.
319.
Ophiochiton modestus (Koehler, 1897)
Ophionereis andmanensis (James, 1987)
Ophionereia dubia (Muller and Troschel, 1842)
Ophionereis porrecta (Lyman, 1860)
Family: Ophiodermatidae
320.
321.
322.
323.
324.
Bathypectinura heros (Lyman, 1879)
Gymnopelta indica (Koehler, 1897)
Ophiarachna incrassata (Lamarck, 1816)
Ophiarachnella gorgonia (Muller and Troschel, 1842)
Ophiarachnella infernalis (Muller and Troschel,
1842)
325.
326.
327.
328.
329.
330.
331.
Ophiarachnella intermedia (Bell, 1888)
Ophiarachnella megaloplax (Bell, 1884)
Ophiarachnella sphenesci (Bell, 1894)
Ophiocormus compsus (A.M.Clark, 1968)
Ophiopeza custos (Koehler, 1897)
Ophiopeza fallax arbacia (A.M.Clark, 1968)
Ophiosammus yoldii (Lutken, 1856)
Family : Ophiacanthidae
332.
333.
334.
335.
336.
337.
338.
339.
340.
176
Ophiacantha abnormis (Lyman, 1879)
Ophiacantha compos ita (Koehler, 1897)
Ophiacantha indica (Ljungman, 1899)
Ophiacantha pentagona (Koehler, 1897)
Ophiacantha sociabilis (Koehler, 1897)
Ophiacantha vagans (Koehler, 1899)
Ophiacantha vestita (Koehler, 1897)
Ophiacantha vorax (Koehler, 1897)
Ophiocamax fasciculata (Lyman, 1883)
341. Ophiocamax rugosa (Koehler, 1904)
342. Ophiomitra integra (Koehler, 1897)
343. Ophioplinthaca rudis (Koehler, 1897)
344. Ophiotreta matura (Koehler, 1904)
Family : Hemieuryalidae
345. Ophiomoeris tenera (Koehler, 1897)
Family : Ophaictidae
346. Ophiactis acosmeta (H.L. Clark, 1971)
347. Ophiactis brachyura (Doderlein, 1971)
348. Ophiactis delagoa (Balinsky, 1957)
349. Ophiactis flexuosa (Koehler, 1897)
350. Ophiactis maculosa (von Martens, 1870)
351. Ophiactis modesta (Brock, 1888)
352. Ophiactis picteti (de Loriol, 1893)
353. Ophiactis savignyi (Muller and Troschel, 1842)
Family: Amphiuridae
354. Amphiodia caullery (Koehler, 1897)
355. Amphioplus intermedius (Koehler, 1983)
356. Amphioplus cyrtacanthus (H.L. Clark, 1915)
357. Amphioplus personatus (Koehler, 1971)
358. Amphioplus andreae (Lutken, 1971)
359. Amphioplus depressus (Ljungman, 1867)
360. Amphioplus gravelyi (James, 1927)
361. Amphioplus hastatus (Ljungman, 1867)
362. Amphioplus laevis (Lyman, 1874)
363. Amphioplus misera (Koehler, 1899)
364. Amphioplus squamata (Delle Chiaje, 1828)
177
365. Amphiura ambigua (Koehler, 1905)
366. Amphiura septemspinosa (H.L. Clark, 1915)
367. Amphiura tenius (H.L. Clark, 1938)
368. Amphiura dispar (Koehler, 1897)
369. Amphiura famula (Koehler, 1910)
370. Amphiura lorioli (Koehler, 1897)
371. Dougaloplus echinatus (Ljungman, 1867)
372. Histampica duplicata (Lyman, 1874)
373. Ophiocentrus dilatatus (Koehler, 1905)
374. Ophiocentrus verticillatus (Doderlein, 1971)
375. Ophiostigma formosa (Lutken, 1899)
Family : Ophiotrichidae
376. Gymnolophus obscura (Ljungman, 1969)
377. Macropohiothrix aspidota (Muller and Troschel, 1842)
378. Macropohiothrix demessa (Lyman, 1869)
379. Macropohiothrix galatheae (Lutken, 1872)
380. Macropohiothrix koehleri (A.M. Clark, 1905)
381. Macropohiothrix longipeda (Lamarck, 1816)
382. Macropohiothrix Propinqua (Lyman, 1861)
383. Macropohiothrix speciosa (Koehler, 1898)
384. Macropohiothrix variabilis (Duncan, 1887)
385. Macropohiothrix hirsuta (Muller and Troschel, 1842)
386. Ophiocnemis marmorata (Lamarck, 1816)
387. Ophiogymna elegans (Ljungman, 1971)
388. Ophiogymna lineata (H.L.Clark, 1969)
389. Ophiogymna pellicula (Duncan, 1887)
390.0phiolophus novarae (Marktanner-Turneretsher,
1887)
391. Ophiomaza cacaotica (Lyman, 1871)
178
392. Ophiopteron elegans (Ludwig, 1888)
393. Ophiothela danae (Verill, 1869)
394. Ophiothrix diligens (Koehler, 1898)
395. Ophiothrix proteus (Koehler, 1905)
396. Ophiothrix purpurae (von Martens, 1867)
397. Ophiothrix vigelandi (A.M. Clark, 1922)
398. Ophiothrix nereidina (Lamarck, 1816)
399. Ophiothrix aristulata (Lyman, 1879)
400. Ophiothrix ciliaris (Lamarck, 1816)
401. Ophiothrix exigua (Lyman, 1874)
402.0phiothrix foveolata (Marktanner-Turneretscher,
1887)
403.
404.
405.
406.
407.
408.
409.
410.
Ophiothrix savignyi (Muller and Troschel, 1842)
Ophiothrix trilineata (Lutken, 1869)
Ophiothrix variegata (Duncan, 1887)
Ophiothrix vitrea (Doderlein, 1896)
Ophiothrix fumaria (Muller and Troschel, 1842)
Ophiothrix striolata (Grube, 1868)
Ophiothrix accedens (Koehler, 1966)
Ophiothrix sp.
Class: ECHINOIDEA
Order: CIDAROIDA
Family: Cidariidae
411.
412.
413.
414.
415.
416.
Eucidaris metularia (Lamarck, 1816)
Histocidaris denticulata (Koehler, 1907)
Phyllacanthus forcipulatus (Mortensen, 1936)
Phyllacanthus imperialis (lamarck, 1816)
Prinocidaris purpurata (Wyville- Thompson, 1869)
Prinocidaris baculosa (Lamarck, 1816)
179
417.
418.
419.
420.
421.
Prinocidaris bispinosa (Lamarck, 1816)
Prinocidaris verticillata (Lamarck, 1816)
Stereocidaris alcocki (Anderson, 1894)
Stereocidaris indica (Doderlein, 1901)
Stylocidris albidens (H.L. Clark, 1925)
422. Stylocidaris brevicollis (de Meijere, 1904)
423. Stylocidaris lorioli (Koehler, 1927)
424. Stylocidaris tiara (Anderson, 1894)
425. Acanthocidaris maculicollis (Meijere, 1903)
Order: ECHINOTHURIOIDA
Family: Echinothuridae
426.
427.
428.
429.
430.
Hygrosoma luculentum (Agassiz, 1879)
Phormosoma bursarium (Agassiz, 1881)
Phormosoma verticillatum (Mortensen, 1904)
Phormosoma sp.
Sperosoma biseriatum (Doderlein, 1901)
Order: DIADEMATOIDA
Family : Diadematidae
431. Astropyga radiata (Leske, 1778)
432. Centrostephanus nitidus (Koehler, 1927)
433. Chaetodiadema granulatum (Mortensen, 1903)
434. Diadema savignyi (Michelin, 1845)
435. Diadema setosum (Leske, 1758)
436. Echinothrix calamaris (Pallas, 1774)
437. Echinothrix diadema (Linneaus, 1758)
Family: Aspidodiadematidae
438. Aspidodiadema nicobaricum (Doderlein, 1901)
180
Order: PEDINOIDA
Family : Pedinidae
439. Coenopedina depressa (Koehler, 1927)
Order: SELENIOIDA
Family : Saleniidae
440. Salenocidaris miliaris (Mortensen, 1939)
441. Salenia sculpta (Koehler, 1927)
Order: PHYMOSOMATIDAE
Family : Stomechinidae
442. Stomopneustes variolaris (Lamarck, 1816)
Order: ARBACIOIDA
Family: Arbaciidae
443. Arbacia punctulata (Lamarck, 1861)
444. Coelopleurus vittiatus (Koehler, 1927)
445. Pygmaeocidaris prionigera (Agassiz, 1879)
Order: TEMNOPLEUROIDA
Family : Temnopleuridae
446. Mespilia globulus (Linneaus, 1758)
447.
448.
449.
450.
451.
452.
453.
454.
455.
Microcyphus ceylanicus (Mortensen, 1925)
Paratrema doderleini (Mortensen, 1904)
Printechinus impressus (Koehler, 1927)
Prionechinus agassizi (Wood-Mason and Alcock,1891)
Salmaciella dussumieri (L. Agassiz and Desor, 1846)
Salmacis belli (Doderlein, 1902)
Salmacis bicolor (Agassiz, 1841)
Salmacis virgulata (Agassiz, 1846)
Salmacis belli (Doderlein, 1902)
181
456.
457.
458.
459.
460.
461.
Temnopleurus apodus (Agassiz and H.L. Clark, 1943)
Temnopleurus proctalis (Koehler, 1927)
Temnopleurus toreumaticus (Leske, 1778)
Temnopleurus alexandri (Bell, 1884)
Temnometra scillae (Mazetti, 1894)
Trigonocidaris versicolor (Koehler, 1927)
Family : Toxopneustidae
462.
463.
464.
465.
Gymnechinus robillaridi (de Loriol, 1883)
Pseudoboletia maculata (Troschel, 1869)
Toxopneustes pileolus (Lamarck, 1816)
Tripneustes gratilla (Linneaus, 1758)
466.
467.
468.
469.
470.
471.
Order : Echinoida
Family : Echinometridae
Colobocentrotus atratus (Linneaus, 1758)
Echinometra mathai (de Blainvillae, 1825)
Echinometra oblonga (de Blainvillae, 1969)
Echinostrephus molaris (de Blainvillae, 1825)
Heterocentrotus mammilatus (Linneaus, 1758)
Heterocentrotus trigonarius (Lamarck, 1816)
Order: HOLECTYPOIDA
Family : Echinoneidae
472. Echinoneus cyclostomus (Leske, 1778)
Order: CLYPEASTEROIDA
Family: Clypeasteridae
473. Clypeaster annandalei (Koehler, 1922)
474. Clypeaster fervens (Koehler, 1922)
475. Clypeaster humilis (Leske, 1778)
182
476. Clypeaster rarispinus (de Meijere, 1903)
477. Clypeaster reticulatus (Linneaus, 1758)
Family : Arachnoididae
478. Arachnoides placenta (Linneaus, 1758)
Family: Fibularidae
479.
480.
481.
482.
483.
Echinocyamus crisp us (Mazetti, 1893)
Echinocyamus sollers (Koehler, 1922)
Fibularia cribellum (de Meijere, 1904)
Fibularia oblonga (Gray, 1847)
Fabularia volva (Agassiz, 1847)
Family : Laganidae
484.
485.
486.
487.
Laganum
Laganum
Laganum
Laganum
decagonale (de Blainvillae, 1827)
depressum (Lesson, 1841)
laganum (Leske, 1778)
retinens (Koehler, 1922)
488.
489.
490.
491.
492.
493.
494.
495.
Laganum retinense mortenseni (Mortensen, 1948)
Laganum versatile (Koehler, 1922)
Peronella lessueri (Valenciennes, 1841)
Peronella macroproctes (Koehler, 1922)
Pronella oblonga (Mortensen, 1948)
Pronella orbicularis (leske, 1778)
Pronella rubra (Doderlein, 1885)
Pronella rutlandi (Koehler, 1922)
Family: Astriclypeidae
496. Echinodiscus auritus (Leske, 1778)
497. Echinodiscus bisperforatus (Leske, 1778)
183
Order: CASSIDULOIDA
Family: Echinolampadidae
498. Echinolampas alexandri (de Loriol, 1876)
499. Echinolampas castanea (Alcock, 1894)
500. Echinolampas ouata (Leske, 1778)
Order: SPATANGOIDA
Family : Hemiasteridae
501. Hemiaster uanus (Koehler, 1914)
Family : Palaeostomatidae
502. Palaeostoma mirabile (Gray, 1851)
Family : Pericosmidae
503. Pericosmus micronesius (Koehler, 1914)
Family : Schizasteridae
504.
505.
506.
507.
508.
509.
510.
511.
Brisaster indicus (Koehler, 1914)
Faorina chinensis (Gray, 1851)
Moira stygia (Lutken, 1872)
Schizaster kempi (Koehler, 1914)
Schizaster angulatus (Koehler, 1914)
Schizaster gibberulus (Agassiz, 1847)
Schizaster inuestigatoris (Koehler, 1914)
Schizaster compactus (Koehler, 1914)
Family : Brissidae
512.
513.
514.
515.
184
Brissopsis luzonica (Gray, 1851)
Brissopsis oldhami (Alcock, 1893)
Brissopsis parallela (Koehler, 1914)
Brissus latecarinatus (Leske, 1778)
516.
517.
518.
519.
520.
Gymnopatagus magnus (Agassiz and Alcock, 1907)
Gymnopatagus valdiviae (Doderlain, 1901)
Metalia latissima (H.L. Clark, 1925)
Metalia spatagus (Linneaus, 1758)
Rhynobrissus pyramidalis (Agassiz, 1872)
Family : Spatangidae
521. Maretia planulata (Lamarck, 1914)
522. Nacospatangus alta (Agassiz, 1863)
Family : Loveniidae
523.
524.
525.
526.
Breynia verdenburgi (Anderson, 1907)
Lovenia elongata (Gray, 1845)
Lovenia gregalis (Alcock, 1834)
Lovenia subcarinata (Gray, 1845)
Family : Asterostomatidae
527. Araeolampas glauca (Wood-Masona and Alcock, 1891)
528. Argopatagus vitreus (Agassiz, 1881)
529. Elipneustes denudatus (Koehler, 1914)
530. Elipneustes rubens (Koehler, 1914)
531. Heterobrissus hemingi (Anderson, 1899)
532. Linopneustes spectabilis (de Meijere, 1904)
533. Paleotrema ovatum (Koehler, 1914)
Class: HOLOTHUROIDEA
Order: ASPIDOCHIROTIDA
Family: Holothuriidae
534. Actinopyga mauritiana (Quoy and Gaimard, 1833)
535. Actinopyga lecanora (Jaeger, 1833)
536. Actinopyga echinities (Jaeger, 1833)
185
637.
638.
639.
640.
641.
642.
643.
644.
646.
646.
647.
648.
649.
660.
661.
662.
663.
664.
666.
666.
667.
668.
669.
660.
661.
662.
663.
664.
666.
666.
186
Aetinopyga miliaris (Quoy and Gaimard, 1833)
Bohadssehia argus (Jaeger, 1833)
Bohadsehia marmorata (Jaeger, 1833)
Bohadsehia graeffei (Semper, 1868)
Bohadsehia tenuissiama (Semper, 1868)
Holothuria pyxis (Selenka, 1867)
Holothuria inhabilis (Selenka,1867)
Holothuria rigida (Selenka, 1867)
Holothuria atra (Jaeger, 1833)
Holothuria edulis (Lesson, 1830)
Holothuria paradalis (Selenka, 1867)
Holothuria exilis (Koehler and Vaney, 1908)
Holothuria fuseoeinerea (Jaeger, 1833)
Holothuria leueospilota (Brandt, 1836)
Holothuria pervieax (Selenka, 1867)
Holothuria albiventer (Semper, 1893)
Holothuria oeellata (Jaeger, 1833)
Holothuria seabra (Jager, 1833)
Holothuria fuseogilva (Cherbonnier, 1908)
Holothuria nobilis (Selenka, 1867)
Holothuria diffieilis (Semper, 1868)
Holothuria erinaeeus (Semper, 1868)
Holothuria moebii (Ludwig, 1883)
Holothuria eineraseens (Brandt, 1836)
Holothuria prompta (Koehler and Vaney, 1908)
Holothuria kurti (Ludwig, 1908)
Holothuria spinifera (Theel, 1886)
Holothuria arenieola (Semper, 1868)
Holothuria graeillis (Semper, 1868)
Holothuria hilla (Lesson, 1830)
567.
568.
569.
570.
Holothuria
Holothuria
Holothuria
Holothuria
impatiens (Forskal, 1775)
remollescens (Lampert, 1908)
integra (Koehler and Vaney, 1908)
coluber (Semper, 1869)
Family : Labidodematidae
571. Labidodemas rugosum (Ludwig, 1875)
572. Labidodemas semperianum (Selenka, 1867)
Family : Stichopodidae
573. Aspostichopus japonicus (Selenka, 1969)
574. Stichopus chloronatus (Brandt, 1835)
575. Stichopus hermanni (Semper, 1868)
576. Stichopus horrens (Selenka, 1867)
577. Stichopus vastus (Sluiter, 1888)
578. Thelenota ananas (Jaeger, 1833)
Family: Synallactidae
579. Allopatides dendroides (Koehler and Vaney, 1905)
580. Bathyplotes assimilis (Koehler and Vaney, 1905)
581. Bathyplotes cinctus (Koehler and Vaney, 1905)
582. Bathyplotes crenulatus (Koehler and Vaney, 1905)
583. Bathyplotes pappillosus (Koehler and Vaney, 1905)
584. Bathyplotes profundus (Koehler and Vaney, 1905)
585. Bathyplotes variabilis (Koehler and Vaney, 1905)
586. Benthothuria cristatus (Koehler and Vaney, 1905)
587. Benthothuria distortus (Koehler and Vaney, 1905)
588. Mesothuria abbreviata (Koehler and Vaney, 1905)
589. Mesothuria incerta (Koehler and Vaney, 1905)
590. Mesothuria multipes (Ludwig, 1894)
187
591. Mesothuria squamosa (Koehler and Vaney, 1905)
592. Pleopatides gelatinosus (Walsh, 1891)
593. Pleopatides insignis (Koehler and Vaney, 1905)
594. Pleopatides modestus (Koehler and Vaney, 1905)
595. Pleopatides mollis (Koehler and Vaney, 1905)
596. Pleopatides ovalis (Walsh, 1891)
597. Pleopatides verrucosa (Koehler and Vaney, 1905)
598. Pseudistichopus occulatus (Marenzeller, 1893)
599. Synallactes horridus (Koehler and Vaney, 1905)
600. Synallactes pellucidus (Koehler and Vaney, 1905)
601. Synallactes rigidus (Koehler and Vaney, 1905)
602. Synallactes woodmasoni (Walsh, 1891)
Order: DENDROCHIROTIDA
Family : Psolidae
603. Psolidium rugosum (Koehler and Vaney, 1905)
604. Psolus mannarensis (James, 1927)
605. Psolus membranaceus (Koehler and Vaney, 1905)
606. Psolus sp.
Family: Phyllophoridae
607. Actinocucumis typicus (Ludwig, 1875)
608. Afrocucumis africana (Semper, 1868)
609. Oshimella ehrenbergi (Selenka, 1868)
610. Phyllophorus celer (Koehler and Vaney, 1905)
611. Phyllophorus intermedius (Koehler and Vaney, 1905)
612. Phyllophorus parvipedes (H.L.Clark, 1938)
613. Phyllophorus sp.
614. Phyllophorus cubuensis (Semper, 1868)
188
615. Phyllophorus brocki (Ledwig, 1888)
616. Phyrella fragilis (Ohshima, 1912)
617. Pseudocucumis acicula (Semper, 1868)
Family : Cucumariidae
618.
619.
620.
621.
622.
623.
624.
625.
626.
627.
628.
Aslia forbesi (Bell, 1884)
Althyone sp.
Cladolabes acicula (Semper, 1983)
Cucumaria ardens (Koehler and Vaney,
Cucumaria ariana (Koehler and Vaney,
Cucumaria frauenfeldi (Ludwig, 1971)
Cucumaria inflexa (Koehler and Vaney,
Cucumaria turbinata (Hutton, 1988)
Havelockia versicolor (Semper, 1868)
Hemithyone semperi (Bell, 1884)
Leptopentacta bacilliformis (Koehler
1905)
1905)
1905)
and Vaney,
1905)
629.
630.
631.
632.
633.
634.
635.
636.
637.
638.
639.
640.
641.
Leptopentacta imbricata (Semper, 1868)
Leptopentacta japonicus (Sluiter, 1880)
Pentacta quadrangularis (Troschel, 1846)
Pseudocnus echinatus (von Marenzeller, 1882)
Pseudocolochirus tricolor (Sluiter, 1971)
Pseudocolochirus violaceus (Theel, 1882)
Stolus buccalis (Stimpson, 1855)
Stolus conjungens (Semper, 1868)
Stolus rapax (Koehler and Vaney, 1905)
Thorsonia investigatoris (Koehler and Vaney, 1905)
Thyone dura (Koehler and Vaney, 1905)
Thyone papuensis (Theel, 1886)
Trachythyone alcocki (Koehler and Vaney, 1905)
189
Order: DACTYLOCHIROTIDA
Family: Yapsilothuridae
642. Yapsilothuria bitentaculata (Ludwig, 1894)
643.
644.
645.
646.
647.
648.
Order: ELASIPODODA
Family : Deimatidae
Amphideima investigatoris (Koehler and Vaney,
1905)
Deima blakei (Theel, 1886)
Deima validum (Theel, 1882)
Oneiriphanta conservata (Koehler and Vaney, 1905)
Orphnurgus glaber (Walsh, 1891)
Orphnurgus invalidus (Koehler and Vaney, 1905)
Family : Laetmogonidae
649. Apodogaster alcocki (Walsh, 1901)
650. Laetmogone spongiosa (Theel, 1882)
651. Laetmogone violacea (Theel, 1882)
Family : Psychropotidae
652. Benthodytes glutinosa (Perrier, 1902)
653. Benthodytes typica (Theel, 1882)
654. Filithuria elegans (Koehler and Vaney, 1905)
Family: Pelagothuriidae
655. Euriplaster obscura (Koehler and Vaney, 1905)
Order: APODIDA
Family : Synaptidae
656. Anapta gracilis (Semper, 1868)
657. Chondrocolea baselii (Jaeger, 1833)
190
658.
659.
660.
661.
662.
663.
664.
665.
666.
667.
668.
669.
670.
671.
672.
673.
674.
675.
676.
677.
Euapta godeffroyi (Semper, 1868)
Labidoplax sp.
Leptosynapta sp.
Opheodesoma grisea (Semper, 1868)
Patinapta oollax (von Marenzeller, 1882)
Protankyra conferta (Koehler and Vaney, 1905)
Protankyra denticulata (Koehler and Vaney, 1905)
Protankyra errata (Koehler and Vaney, 1905)
Protankyra innominata (Koehler and Vaney, 1905)
Protankyra pseudodigitata (Semper, 1868)
Protankyra similes (Semper, 1868)
Protankyra timida (Koehler and Vaney, 1905)
Protankyra tristis (Koehler and Vaney, 1905)
Protankyra tuticorensis (Jaeger, 1833)
Protankyra sp.
Psamothuria ganapati (Rao, G.C.1968)
Synapta maculata (Chamisso and Eysenhardt, 1821)
Synapta sp.
Synaptulam recta (Semper, 1868)
Synaptula striata (Sluiter, 1888)
Family: Chiridotidae
678. Polycheira rufescens (Brandt, 1835)
679. Trochodota havelockensis (Rao, G.C.1975)
Family : Myriotrochidae
680. Ankyloderma brevicaudatum (Koehler and Vaney,
1905)
681. Ankyloderma contortum (Koehler and Vaney, 1905)
682. Ankyloderma intermedim (Koehler and Vaney, 1905)
683. Ankyloderma danielseni (Theel, 1886)
191
684. Ankyloderma musculus (Risso, 1826)
685. Ankyloderma polymorphium (Koehler and Vaney,
1905)
686. Trochostoma albicans (Koehler and Vaney, 1905)
687. Trochostoma andamanensis (Walsh, 1891)
688. Trochostoma ecalcareum (Koehler and Vaney, 1905)
689. Trochostoma elegatum (Koehler and Vaney, 1905)
690. Trochostoma pauperum (Koehler and Vaney, 1905)
Order: MALPADIIDAE
Family: Caudiniidae
691. Acaudina leucoprocta (Clark, H.L.1938)
692. Acaudina malpadiodes (Semper, 1868)
693. Paracaudina australis (Semper, 1868)
Source: Sastry (2007)
9. ACKNOWDEDGEMENTS
The authors are grateful to the The Director, Zoological
Survey of India for facilities and Ministry of Environment
and Forests for providing financial assistance through
the projects of National Coral Reef Research Institute,
Zoologicial Survey of India.
10. REFERENCES
Ausich, W.I. 1997. Calyx plate homologies and early
evolutionary history of the Crinoidea. Paleont. Soc.
Papers, 3: 289-304.
Ausich, W. 1998. Early phylogeny and subclass division of
the Crinoidea (Phylum Echinodermata). Journal of
Paleontology, 72(3): 499-510.
192
Ausich, W. 1999. Origin of crinoids. In: Candia Carnevali,
M.D.C. and Bonasoro, F. (Eds.) Echinoderm Research
1998. Balkema, Rotterdam: 237-242.
Ausich, W. and Kammer, T.W. 2001. The study of crinoids
during the 20th century and the challenges of the 21st
century. Journal of Paleontology, 75(6): 1161-1173.
Baker, AN. 2003. Concentricycloidea (sea daisies). In:
Hutchins, M., Thoney, D.A and Schlager, D. (Eds.)
Grzimek's Animal Life Encyclopedia, 2nd edition.
Volume 1. Gale Group, Farmington Hills, Michigan:
381-386.
Baker, AN., Rowe, F.W.E. and Clark, H.E.S. 1986. A new
class of Echinodermata from New Zealand. Nature,
321: 862-864.
Barker, M. 2001. Echinoderms 2000. Balkema, Lisse,
590 pp.
Bather, F.A 1900. Part III The Echinoderma. In:
Lankester, E.R. (Ed.) A Treatise on Zoology. Adam
and Charles Black, London: 1-344.
Belyaev, G.M. 1990. Is it valid to isolate the genus
Xyloplax as an independent class of echinoderms?
ZoologicheskiiZhurnal, 69: 83-96.
Blake, D.B. 1987. A classification and phylogeny of postPaleozoic seastars (Asteroidea: Echinodermata).
Journal of Natural History, 21: 481-528.
Blake, D.B. 1989. Asteroidea: functional morphology,
classification and phylogeny. In: Jangoux, M. and
Lawrence, J.M. (Eds.) Echinoderm Studies 3.
Balkema, Rotterdam: 179-233.
193
Blake, D.B. 2000. The Class Asteroidea (Echinodermata):
fossils and the base crown group. American Zoologist,
40: 316-325.
Blake, D.B. and Elliott, D.R 2003. Ossicular homologies,
systematics, and phylogenetic implications of
certain North American Carboniferous asteroids
(Echinodermata).
Journal
of
Paleontology,
77(3): 476-489.
Blake, D.B. and Hagdorn, H. 2003. The Asteroidea
(Echinodermata) of the Muschelkalk (Middle
Triassic) of Germany. Palaontologische Zeitschrift,
77: 23-58.
Blake, D.B. and Hotchkiss, F.H.C. 2004. Recognition of the
asteroid (Echinodermata) crown group: implications
of the ventral skeleton. Journal of Paleontology,
78(2): 359-370.
Blake, D.B., Janies, D.A. and Mooi, R 2000. Evolution
of starfishes: morphology, molecules, development,
and paleobiology. Introduction to the symposium.
American Zoologist, 40: 311-315.
Bourlat, S.J., Juliusdottir, T., Lowe, C.J., Freeman, R,
Aronowicz,J., Kirschner,M.,Lander, E.S., Thorndyke,
M., Nakano, H., Kohn, A.B., Heyland, A., Moroz, L.L.,
Copley, RR and Telford, M.J. 2006. Deuterostome
phylogenyreveals monophyletic chordates and the
new phylum Xenoturbellida. Nature, 444: 85-88.
Bruguiere, J.G. 1791. Tableau encyclopedique et
methodique de trois regnes de la nature, vol. 7.
Contenant l'helminthologie, ou les vers infusoires, les
vers intestins, les vers mollusques and c. Panckoucke,
Paris.
194
Cameron, C.B., Garey, J.R. and Swalla, B.J. 2000.
Evolution of the chordate body plan: new insights
from phylogenetic analyses of deuterostome phyla.
Proceedings of the National Academy of Sciences
USA 97: 4469-4474.
Candia Carnevali, M.D. and Bonasoro F. 2001. Echinoderm
Research 1998: Proceedings of the Fifth European
Conference on Echinoderms, Milan, Italy, 7-12
September 1998. Balkema, Rotterdam: 550 pp.
Cisternas, P., Selvakumaraswamy, P. and Byrne, M. 2004.
Evolution of development and the Ophiuroidea revisited. In: Heinzeller, T. and Nebelsick, J.H.
(Eds.) Echinoderms: Munchen. Taylor and Francis,
London: 521-526.
Clark, A.M. and Rowe, F.W.E. 1971. Monograph of
sahallow-water Indo-West Pacific Echinoderms.,
Trustees of the British Museum (Natural History),
London.
Clark, A.M. and Downey, M.E. 1992. Starfishes of the
Atlantic. Chapman and Hall, London, 794 pages.
Cohen, B.L., Ameziane, N., Eleaume, M. and Richer de
Forges, B. 2004. Crinoid phylogeny: a preliminary
analysis
(Echinodermata:
Crinoidea). Marine
Biology, 44(3): 605-617.
Cuenot, L. 1948. Anatomie, ethologie et systematiques
des echinoderms. In: Grasse, P.P. (Ed.) Traite de
Zoo logie, vol. XI. Masson, Paris: 3-275.
David, B., Lefebvre, B., Mooi, R. and Parsley, R. 1999.
Homalozoans in the light of the extraxial-axial
theory of skeletal homologies. In: Candia Carnevali,
195
M.D.C. and Bonasoro, F. (Eds.) Echinoderm research
1998. Balkema, Rotterdam, p. 319.
David, J., Roux, M., Messing, C.G. and Ameziane, N. 2006.
Revision of the pentacrinid stalked crinoids of the
genus Endoxocrinus (Echinodermata, Crinoidea),
with a study of environmental control of characters
and its consequences for taxonomy. Zoo taxa,
1156: 1-50.
Dean, J. 1999. What makes an ophiuroid? A morphological
study of the problematic Ordovician stelleroid
Stenaster and the paleobiology of the earliest
asteroids and ophiuroids. Zoological Journal of the
Linnean Society, 126: 225- 250.
Dominguez-Alonso, P. 1999. The early evolution of
echinoderms: the class Ctenocystoidea and its closest
relatives revisited. In: Candia Carnevali, M.D.C.
and Bonasoro, F. (Eds.) Echinoderm research 1998.
Balkema, Rotterdam: 263-268.
Fell, H.B. 1948. Echinoderm embryology and the origin
of chordates. Biological 'Reviews of the Cambridge
Philosophical Society, 23: 81-107.
Fell, H.B. 1962. Evidence for the validity of Matsumoto's
classification of the Ophiuroidea. Publications of the
Seto Marine Biological Laboratory, 10: 145-152.
Fell, H.B. 1963. The evolution of echinoderms. Annual
Report of the Smithsonian Institution for 1962:
457-490.
Fell, H.B. and Pawson, D.L. 1966a. Order Diadematoida.
In: R.C. Moore (Ed.) Treatise on Invertebrate
Paleontology. Part U, Echinodermsta 3(2). University
of Kansas Press, Lawrence: 350-365.
196
Fell, H.B. and Pawson, D.L. 1966b. Echinacea. In: RC.
Moore (Ed.) Treatise on Invertebrate Paleontology.
Part U, Echinodermsta 3(2). University of Kansas
Press, Lawrence: 367-440.
Feral, J.-P. and David, B. 2001. Echinoderm Research
2001. Balkema, Rotterdam, 337 pp.
Forey P.L., Fortey, RA., Kenrick, P. and Smith, A.B.
2004. Taxonomy and fossils: a critical appraisal.
Philosophical Transactions of the Royal Society.
Series B. Biological Sciences, 359: 639-653.
Gale, A.S. 1987. Phylogeny and classification of the
Asteroidea (Echinodermata). Zoological Journal of
the Linnaean Society, 89: 107-132.
Gilliland, P. 1992. Holothurians in the Blue Lias of
southern Britain. Palaeontology, 35(1): 159-210.
Gilliland, P. 1993. The skeletal morphology, systematics
and evolutionary history of holothurians. Special
Papers in Palaeontology, 47: 1-147.
Haude, R 2004. Mode of life of ophiocistioids (Echinozoa)
according to plated and "naked" forms in the Rhenish
Devonian. In: Heinzeller, T. and Nebelsick, J.H.
(Eds.) Echinoderms: Munchen. Taylor and Francis,
London: 417- 420.
Heinzeller, T. and Nebelsick, J.H. 2004. Echinoderms:
Munchen. Balkema, Leiden, 633 pp.
Herringshaw, L.G., Smith, M.P. and Thomas, A.T.
2007. Evolutionary and ecological significance
of Lepidaster grayi, the earliest multiradiate
starfish. Zoological Journal of the Linnean Society,
150(4): 743-754.
197
Hess, H., Ausich, W., Brett, C.E. and Simms, M.J. 2003.
Fossil crinoids. Cambridge University Press,
Cambridge, UK, 292 pp.
Hotchkiss,
F.H.C.
1977.
Ophiuroid
Ophiocanops
(Echinodermata) not a living fossil. Journal of
Natural History, 11: 377-380.
Hotchkiss, F.H.C. 1993. A new Devonian ophiuroid
(Echinodermata: Ophiuroidea) from New York State
and its bearing on the origin of ophiuroid upper
arm plates. Proceedings of the Biological Society of
Washington, 106(1): 63- 84.
Hotchkiss, F.H.C. 1995. Loven's law and adult ray
homologies in echinoids, ophiuroid, edrioasteroids,
and an ophiocistioid (Echinodermata: Eleutherozoa).
Proceedings of the Biological Society of Washington,
108(3): 401-435.
Hotchkiss, F.H.C. 2000. On the number of rays
starfish. American Zoologist, 40: 340-354.
III
a
Hrincevich, AW., Axayacatl, R.-O. and Foltz, D.W. 2000.
Phylogenetic analysis of molecular lineages in a
species-rich subgenus of sea stars (Leptasterias
American
Zoologist,
subgenus
Hexasterias).
40: 365-374.
Hyman, L.H. 1955. The invertebrates: Echinodermata.
McGraw-Hill, New York, 763 pp.
Jagt, J.W.M. 1999. Ophiuroid diversity in the type area
of the Maastrichtian Stage. Geologie en Mijnbouw,
78(2): 197-206.
James, D.B. 1987. Research on Indian Echinoderms- A
review. J. Mar. Boil. Ass. India, 25 (1983): 91-108.
198
Jangoux, M. and Lawrence, J.M. 2001. Echinoderm
Studies 6. Taylor and Francis, London, 285 pp.
Janies, D. 2001. Phylogenetic relationships of extant
echinoderm classes. Canadian Journal of Zoology,
79: 1232-1250.
Janies, D. and Mooi, R 1999. Xyloplax is an asteroid.
In: Candia Carnevali C. and Bonasoro F. (Eds.),
Echinoderm Research 1998. Balkema, Rotterdam:
311-316.
Jefferies, RP.S., Brown, N.A. and Daley, P.E.J. 1996. The
early phylogeny of chordates and echinoderms and
the origins of chordate left-right asymmetry and
bilateral symmetry. Acta Zoologica, 77: 101-122.
Kasyanov, V.L. 2001. Reproductive strategy of marine
bivalves and echinoderms. Science Publishers,
Enfield, New Hampshire, 240 pp.
Kerr, A.M. 2001. Phylogeny of the apodan holothurians
(Echinodermata) inferred from morphology. Zoological
Journal of the Linnean Society, 133: 53-62.
Kerr, A.M. and Kim, J. 1999. Bi-penta-bi-decaradial
symmetry: a review of evolutionary and developmental
trends in Holothuroidea (Echinodermata). Journal
of Experimental Zoology, 285: 93-103.
Kerr, A.M. and Kim, J. 2001. Phylogeny of Holothuroidea
(Echinodermata) inferred from morphology. Zoological
Journal of the Linnean Society, 133: 63-81.
Kerr, A.M., Janies, D.A., Clouse, RM., Samyn, Y., Kuszak,
J., Kim, J. 2005. Molecular phylogeny of coral reef
sea cucumbers (Holothuriidae: Aspidochirotida)
199
based on 16S mitochondrial ribosomal
sequence. Marine Biotechnology, 7: 53-60.
DNA
Klein, J.T. 1734. Naturalis disposition Echinodermstium.
Accesseit Lucubratiuncula des aculeis echinorum
marinorum, cun Spicilegio de belemnitis. Liutteris
Schreiberianis, Gedani, 78 pp.
Knott, K.E. and Wray, G.A 2000. Controversy and consensus
in asteroid systematics: new insights to ordinal
and familial relationships. American Zoologist,
40: 382-392.
Kroh, A 2004. First fossil record of the family Euryalidae
(Echinodermata: Ophiuroidea) from the Middle
Miocene of the central Mediterranean. In: Heinzeller,
T. and Nebelsick, J.H. (Eds.) Echinoderms: Munchen.
Taylor and Francis, London: 447-452.
Lacey, K.M.J., McCormack, G.P., Keegan, B.F. and Powell,
R. 2005. Phylogenetic relationships within the class
Holothuroidea, inferred from 18S rRNA gene data.
Marine Biology, 147: 1149-1154.
Lafay, B., Smith AB. and Christen, R. 1995. A combined
morphological and molecular approach to the
phylogeny of asteroids (Asteroidea: Echinodermata).
Systematic Biology, 44(2): 190-208.
Lamarck, J.B.P. 1801. Systeme des animaux sans vertebres.
Maillard, Paris, 432 pp.
Lawrence, J.M. 2001. Edible sea urchins: biology and
ecology. Elsevier, Oxford, UK, 432 pp.
Lawrence, J.M. 2006. Edible sea urchins: biology and
ecology, 37. Elsevier, Oxford, UK, 380 pp.
200
Lefebvre, B. 2007. Early Palaeozoic palaeobiogeography
and palaeoecology of stylophoran echinoderms.
Palaeogeography, Palaeoclimatology, Palaeoecology,
245: 156-199.
Le Gac, M., Feral, J. P., Poulin, E., Veyret, M. and
Chenuil, A 2004. Identification of allopatric clades
in the cosmopolitan ophiuroid species complex
Amphipholis squamata (Echinodermata). The end
of a paradox? Marine Ecology Progress Series, 278:
171-178.
Lee, Youn-Ho. 2003. Molecular phylogenies and divergence
times of sea urchin species of Strongylocentrotidae,
Echinoida. Molecular Biology and Evolution, 20(8):
1211-1221.
Leuckart, R. 1854. Bericht tiber die Leistungen in der
naturgeschichte der niederen Thiere wahrend der
Jahre 1848- 1853. Archiv fur Naturgeschichte, 20(2):
289-473.
Linnaeus, C. 1758. Systema naturae. 10th edition. Lucae,
Juntiniana: 3-376.
Littlewood, D.T.J. 1995. Echinoderm class relationships
revisited. In: Emson, R., Smith, AB. and Campbell,
A (Eds.) Echinoderm Research 1995. Balkema,
Rotterdam: 19-28.
Littlewood, D.T.J. and Smith, AB. 1995. A combined
morphological and molecular phylogeny for sea
urchins (Echinoidea: Echinodermata). Philosophical
Transactions of the Royal Society of London. Series
B. Biological Sciences, 347: 213-234.
201
Littlewood, D.T.J., Smith, A.B., Clough, K.A. and
Emson, R.H. 1997. The interrelationships of the
echinoderm classes: morphological and molecular
evidence. Biological Journal of the Linnean Society,
67: 409-438.
Ludwig, H. 1889-1907. Echinodermen. In: Bronn, H.G.
(Ed.) Klassen und Ordnungen des Tierreichs, 2(3).
Winter, Leipzig, 1602 pp.
Mah,
C.L. 2000. Preliminary phylogeny of the
forcipulatacean Asteroidea. American Zoologist,
40: 375-381.
Mah, C.L. 2006. A new species of Xyloplax (Echinodermata:
Asteroidea: Concentricycloidea) from the northeast
Pacific: comparative morphology and a reassessment
of phylogeny. Invertebrate Biology, 125(2): 136-153.
Mah,
C.L. 2007. Phylogeny of the Zoroasteridae
(Zorocallina: Forcipulatida). Zoological Journal of
the Linnean Society, 150: 177-210.
Matranga, V. 2005.
Berlin, 277 pp.
Echinodermsta. Springer-Verlag,
Matsubara, M., Komatsu, M., Araki, T., Asakawa, S.,
Yokobori, S.-I., Watanabe, K. and Wada, H. 2005.
The phylogenetic status of Paxillosida (Asteroidea)
based on complete mitochondrial DNA sequences.
Molecular Genetics and Evolution, 36: 598-605.
McEdward, L.R. and Miner, E.G. 2001. Larval and lifecycle patterns in echinoderms. Canadian Journal of
Zoology, 79: 1125-1170.
Messing, C.G. 1997. Living comatulids. In: Waters, J.A.
and Maples, C.G. (Eds.) Geobiology of echinoderms.
202
Paleontological Society papers 3. Paleontological
Society, Pittsburgh: 3-30.
Messing, C.G. 2007. The crinoid fauna (Echinodermata:
Crinoidea) of Palau. Pacific Science, 61(1): 91-111.
Meyer, D.L. and Macurda, Jr., D.B. 1977. Adaptive
radiation of the comatulid crinoids. Paleobiology,
3 : 74-82.
Mooi, R 2001. Not all written in stone: interdisciplinary
syntheses in echinoderm paleontology. Canadian
Journal of Zoology, 79: 1209-1231.
Mooi, Rand B. David. 1997. Skeletal homologies of
echinoderms. The Paleontological Society Papers,
3: 305-335.
Mooi, R and David, B. 2000. What a new model of
skeletal homologies tells us about asteroid evolution.
American Zoologist, 40: 326-339.
Mooi, R, Rowe, F.W.E., and David, B. 1998. Application
of a theory of axial and extraxial skeletal homologies
to concentricycloid morphology. In: Mooi, Rand
Telford, M. (Eds.), Echinoderms: San Francisco.
Balkema, Rotterdam: 61-62.
Mooi, R and Telford, M. 1998. Echinoderms: San
Francisco. Balkema, Rotterdam, 923 pp.
Moore, RC. 1966-1978. (Ed.) Treatise on Invertebrate
Paleontology. Part S Echinodermata 1, Volumes
1, 2 1967; Part-T, Echinodermsta 2, Volumes 1,2,3
1978; Part-u, Echinodermsta 3 Volumes 1, 2 1966.
University of Kansas Press, Lawrence.
203
Moran, P. J. 1988. The Acanthaster phenomenon.
Australian Institute of Marine Science Monograph
Series, Vol. 7, 78 p.
Nagabhushanam, Rand Rao, G.C. 1969. Preliminary
observation on a collection of shore fauna of Orissa
coast, India. Proc. Zool. Soc. Calcutta, 22: 67-82.
Nagabhushanam, Rand Rao, G.C. 1972. An ecological
survey of the marine fauna of Minicoy Atoll
(Laccadive Archipelago, Arabian Sea). Mitt. Zool.
Mus. Berlin, 48(2): 265-324.
O'Loughlin, P.M. and Waters, J.M. 2004. A molecular
and morphological revision of genera of Asterinidae
(Echinodermata: Asteroidea). Memoirs of the
Museum of Victoria, 61(1): 1-40.
Parsley, R 1999. The Cincta (Homostelea) as blastozoans.
In: Candia Carnevali, M.D.C. and Bonasoro, F. (Eds.)
Echinoderm Research 1998. Balkema, Rotterdam:
369-375.
Paul C.RC and Smith, A.B. 1984. "The early radiation
and phylogeny of echinoderms". Biol. Rev.,
59: 443-481
Pawson, D .L. 1966. Phylogeny and evolution ofholoth uroids.
In: RC. Moore (Ed.) Treatise on Invertebrate
Paleontology. Part- U, Echinodermata 3(2). University
of Kansas Press, Lawrence: 641-646.
Pawson, D.L. 1980. Holothuroidea. In: Broadhead, T.W. and
Waters, J.A. (Eds.) Echinoderms: Notes for a short
course. University of Tennessee Studies in Geological
Science 3. University of Tennessee, Knoxville:
175-189.
204
Pawson, D.L and Fell, H.B. 1965. A revised classification
of the dendrochirote holothurians. Breviora,
214: 1-7.
Pawson, D.L. and Kerr, A.M. 2001. Chitin in echinoderms?
Tentacle sheaths in the deep-sea holothurian
Ceraplectanatrachyderma
(Holothuroidea:
Molpadiida). Gulf of Mexico Science, 19(2): 192.
Pearse, V.B. and Pearse, J.S. 1994. Echinoderm phylogeny
and the place of concentricycloids. In: David, B.,
Guille, A., Feral, J.-P., Roux, M. (Eds.), Echinoderms
Through Time. Balkema, Rotterdam: 121-126.
Pearse, V.B., Pearse, J.S., Hendler, G. and Byrne, M.
1998. An accessible population of Ophiocanops off
NE Sulawesi, Indonesia. In: Mooi, R. and Telford,
M. (Eds.) Echinoderms: San Francisco. Balkema,
Rotterdam: 413-418.
Plancus, J and Gualtaire, N. 1743. De Stella Marina
Echinata Quindecim Radii insturcta Epistolae
binae. (In: James, 1987).
Rasmussen, H.W. and Sieverts Doreck, H. 1978. Articulata.
In: Moore, R.C. and Teichert, C. (Eds.) Treatise on
Invertebrate Paleontology Part T Echinodermsta 2 (3).
University of Kansas Press, Lawrence: T814-TI027.
Reich, M. 2002. Holothurien (Echinodermata) aus der
Oberkreide des Ostseeraumes: Teil1. Myriotrochidae
Theel, 1877. Neues Jahrbuch fur Geologie und
Palaotologie Abhandlungen, 224(3): 373-409.
Reich, M .. 2004. Fossil Holothuroidea (Echinodermata):
an overview. In: Heinzeller, T. and Nebelsick, J.H.
(Eds.) Echinoderms: Munchen. Taylor and Francis,
London, p.602.
205
Reich, M. and Haude, R 2004. Ophiocistioidea (fossil
Echinodermata): an overview. In: Heinzeller, T.
and Nebelsick, J.H. (Eds.) Echinoderms: Munchen.
Taylor and Francis, London: 489-494.
Rose, E.P.F. and Olver, J.P.S. 1988. Jurassic echinoids
of the family Menopygidae: implications for the
evolutionary interpretation and classification of
early Irregularia. In: Burke, RD., Mladenov, P.V.,
Lambert, P. and Parsley, RL. (Eds.) Echinoderm
biology. Balkema, Rotterdam: 149-158.
Roux, M., Messing, C.G. and Ameziane, N. 2002. Artificial
keys to the genera of living stalked crinoids
(Echinodermata). Bulletin of Marine Science, 70(3):
799-830.
Rowe, F.W.E., Baker, A.N. and Clark, H.E.S. 1988.
The morphology, development, and taxonomic
status of Xyloplax Baker, Rowe, and Clark, 1986
(Echinodermata: Concentricycloidea) with the
description of a new species. Proceedings of the Royal
Society of London. Series B. Biological Sciences, 233:
431-459.
Sastry, D.RK. 2007. Echinodermata of India: An
Annotated list. Rec. Zool. Surv. India, Occ. Paper
No., 271: 1-387.
Sastry, D.RK. 2005. Echinodermata of Andaman and
Nicobar Islands, Bay of Bengal: An Annotated list.
Rec. Zool. Surv. India, Occ. Paper No., 233: 1-207.
Shackleton, J.D. 2005. Skeletal homologies, phylogeny and
classification of the earliest asterozoan echinoderms.
Journal of Systematic Palaeontology, 3(1): 29-114.
206
Shu, D.G., Conway Morris, S., Han, J., Zhang, Z.F. and
Liu, J.N. 2002. Ancestral echinoderms from the
Chengjiang deposits of China. Nature, 430: 422-428.
Simms, M.J. 1988. The phylogeny of post-Paleozoic
crinoids. In: Paul, C.RC. and Smith, AB. (Eds.)
Echinoderm phylogeny and evolutionary biology.
Clarendon Press, Oxford: 269-284.
Simms, M.J., Gale, AS., Gilliland, P., Rose, E.P.F. and
Sevastopulo, G.D. 1993. Echinodermata. In: Benton,
M.J. (Ed.) The fossil record 2. Chapman and Hall,
London: 492-528.
Smirnov, AV. 1998. On the classification of the apodid
holothurians. In: Mooi, R and Telford, M. (Eds.)
Echinoderms: San Francisco. Balkema, Rotterdam:
517-522.
Smirnov, AV. 1999. The origin of the Order Apodida
(Holothuroidea) and its families. In: Candia
Carnevali, M.D.C. and Bonasoro, F. (Eds.)
Echinoderm research 1998. Balkema, Rotterdam:
393-395.
Smith, AB. 1984. Echinoid palaeobiology. George Allen
and Unwin, London, 190 pp.
Smith, AB. 1988a. Fossil evidence for the relationships
of extant echinoderm classes and their times of
divergence. In: Paul, C.RC. and Smith, AZ.B. (Eds.)
Echinoderm phylogeny and evolutionary biology.
Clarendon Press, Oxford: 85-97.
Smith, AB. 1988b. Phylogenetic relationship, divergence
times, and rates of molecular evolution for
camarodont sea urchins. Molecular Biology and
Evolution 5(4): 345-365.
207
Smith, A.B. 1997. Echinoderm larvae and phylogeny.
Annual Review of Ecology and Systematics, 28:
219-241.
Smith, A.B. 2004a. Deuterostome phylogeny and the
interpretation of problematic fossil echinoderms.
In: Heinzeller, T. and Nebelsick, J.H. (Eds.)
Echinoderms: Munchen. Taylor and Francis,
London: 543-546.
Smith, A.B. 2004b. Echinoderm roots. Nature, 430:
411-412.
Smith, A.B. 2007. Intrinsic versus extrinsic biases in
the fossil record: contrasting the fossil record of
echinoids in the Triassic and early Jurassic using
sampling data, phylogenetic analysis, and molecular
clocks. Paleobiology, 33(2): 310-323.
Smith, A.B., Lafay, B. and Christen, R. 1992. Comparative
variation of morphological and molecular evolution
through geologic time: 28s ribosomal RNA versus
morphology in echinoids. Philosophical Transactions
of the Royal Society of London. Series B. Biological
Sciences, 338: 365-382.
Smith, A.B., Paterson, G.L.J. and Lafay, B. 1995. Ophiuroid
phylogeny and higher taxonomy: morphological,
molecular
and
palaeontological perspectives.
Zoological Journal of the Linnean Society, 114:
213-243.
Smith, A.B., Pisani, D., Mackenzie-Dodds, J.A., Stockley,
B, Webster, B.L. and Littlewood, T.J. 2006. Testing
the molecular clock: molecular and paleontological
208
estimates oif divergence times in the Echinoidea
(Echinodermata). Molecular Biology and Evolution,
23(10): 1832-1851.
Solovjev, A.N. and Markov, A.V. 2004. The early
evolution of irregular echinoids. In: Heinzeller, T.
and Nebelsick, J.H.(Eds.) Echinoderms: Munchen.
Taylor and Francis, London: 551-556.
Soota, T.D., Mukhopadhyay, S.K. and Samanta, T.K.
1983. On some holothurians from the Andaman and
Nicobar Islands. Rec. zool. Surv. India, 80: 507-524.
Stockley, B., Smith, A.B., Littlewood, T., Lessios, H.A.
and Mackenzie-Dodds, J.A. 2005. Phylogenetic
relationships of spatangoid sea urchins (Echinoidea):
taxon sampling density and congruence between
morphological and molecular estimates. Zoological
Scripta, 34(5): 447-468.
Sumrall, C.D. and Wray, G.A. 2007. Ontogeny in the
fossil record: diversification of body plans and the
evolution of "aberrant" symmetry in Paleozoic
echinoderms. Paleobiology, 33(1): 149-163.
Vickery, M. and McClintock, J.B. 2000. Comparative
morphology of tube feet among the Asteroidea:
phylogenetic implications. American Zoologist, 40:
355-364.
Wada, H. and Satoh, N. 1994. Phylogenetic relationships
among extant classes of echinoderms, as inferred
from sequences of 18S rDNA, coincide with
relationships deduced from the fossil record. Journal
of Molecular Evolution, 38: 41-49.
209
Wagner, G.P., Lo, J., Laine, R. and Almeder, M. 1993.
Chitin in the epidermal cuticle of a vertebrate
(Paralipophrys trigloides, Blenniidae, Teleostei).
Cellular and Molecular Life Sciences, 49(4):317-319.
Wilson, N.G., Hunter, R.L., Lockhart, S.J. and Halanych,
K.M. 2007. Multiple lineages and absence ofpanmixia
III
the "circumpolar" crinoid Promachocrinus
kerguelensis from the Atlantic sector of Antarctica.
Marine Biology, 152(4): 895-904.
Yokota, Y., Matranga, V. and Smolenicka, Z. 2002. The sea
urchin: from basic biology to aquaculture. Balkema,
Rotterdam, 293 pp.
210