Professional Documents
Culture Documents
Tropical Pacific Invertebrata PDF
Tropical Pacific Invertebrata PDF
Invertebrates
A Field Guide to the Marine Invertebrates
Occurring on Tropical Pacific Coral Reefs,
Seagrass Beds and Mangroves
Patrick L. Colin
&
Charles Arneson
Tropical
Pacific
Invertebrates
i
ii
TROPICAL
PACIFIC
INVERTEBRATES
A Field Guide to the Marine Invertebrates
by
iii
Dedicated to
Lori Jane Bell Colin
and the memory of Charles E. Cutress
Published by:
Coral Reef Press
270 North Canon Drive, Suite 1524
Beverly Hills, California 90210
U.S.A
ISBN 0-9645625-0-2
Frontispiece- Reef building corals thrive in the clear, shallow waters of the tropical Pacific. This picture was taken in a
shallow pass through the barrier reef at Losap Atoll in the Federated States of Micronesia.
The Coral Reef Research Foundation was established in 1991 to promote research and education on coral
reefs and other tropical marine environments. This volume is a contribution by CRRF toward that goal. A
portion of the profits from this volume will support activities of the Coral Reef Research Foundation.
iv
Table of Contents
Introduction 1
Phylum Porifera 17
Cnidarians
Phylum Cnidaria 63
Ctenophores
Phylum Ctenophora 140
Worms
Phyla Platyhelminthes, Nemertea, Annelida,
Echiurida, Sipuncula and Hemichordata 143
Molluscs
Phylum Mollusca 157
Crustaceans
Phylum Arthropoda 201
Lophophorates
Phyla Ectoprocta, Phoronida and Brachiopoda 227
Echinoderms
Phylum Echinodermata 235
Ascidians
Phylum Urochordata 267
Index 290
v
Acknowledgements
Foundation by the U.S. National Cancer Institute. Our We would also like to acknowledge those governments,
goal in forming the Coral Reef Research Foundation was institutions, colleagues and coworkers who have had
to re-establish a marine research laboratory to support exceptional involvement with this book. These include,
basic research on atoll ecosystems in the central Pacific. but are not limited to, the following:
The idea was to fill the void left by the closure of the Mid Former and present Coral Reef Research
Pacific Marine Laboratory on Enewetak in 1983. In ful- Foundation staff: Lori J. Bell Colin, Larry Sharron,
fillment of that goal we now have two new research lab- Terry Frohm, Chester Gustaf, Peter Schupp, Ron
oratories, one in Chuuk and the other in Palau, which Sjoken, Don DeMaria, Hector Manglicmot, Matthew
serve as bases for further study on marine biodiversity by Mesubed, Emilio Basilius and Chuck Bowling.
us and scientists from all nations. The National Cancer Federated States of Micronesia: Chuuk State
Institute research project involves careful collection, Government, Marion Henry, Kimiuo Aisek, Gradvin
database development and identification of marine inver- Aisek, Lenny Kolczynski, and crew of the Truk
tebrate species for cancer and AIDS drug development. Aggressor. Republic of the Marshall Islands: The
This book is a result of our past and current studies and is people of Enewetak Atoll; Scott Johnson, Lisa Boucher
our first attempt at organizing field notes, photographs and Clark, Walt Chidsey and the crew of the Liktanur II.
identifications into a volume that can be used by everyone. Philippines: Gabriel S. Casals, Pedro Gonzales,
Domingo Madulid, Marivene Manuel, Hilconida
We have been fortunate over the last fifteen years to work Calumpong, Adrian Zecha and staff of Amanpulo.
and live in a number of locations in the western Pacific. Papua New Guinea: University of Papua New Guinea
The opportunity for us to study, identify and photograph (UPNG) Motupore Island Research Station,
the organisms illustrated here is the result of help by fel- Christensen Research Institute (CRI), Madang Dept. of
low scientists, friends, governments and various research Environment and Conservation, Max Benjamin, Alan
organizations. However, without the permission and Rabe, crew of the M/V Febrina, Kevin Baldwin, crew
interest of local peoples throughout the region, who of the M/V Tiata, Mike Huber, John Rewald, Sae Gwae,
allowed us to visit, dive and collect in their home waters, Matthew and Serena Jebb, Larry Orsak, Jim and Debbie
our work would not have been possible. While we can- Prescott. Republic of Palau: President Kuniwo
not identify all of these individuals separately, we would Nakamura, Division of Marine Resources, Department
first like to acknowledge their contributions and help dur- of Resources and Development; former President
ing the studies that led to the completion of this volume. Ngiratkel Etpison, the Governments of Koror,
Needless to say, this book would not be possible without Melekeok, Ngatpang, and Ngchesar States, Shallum and
their unselfish support and cooperation. Mandy Etpison, Kevin Davidson and crew of the Palau
Aggressor. Indonesia: Hanny and Ineke Batuna,
The formation of the Coral Reef Research Foundation Graham Usher, Roman Palete, crew of the M/V
and the establishment of the Chuuk Atoll Research Serenade, Peter Jennings. Hong Kong: Brian Morton,
Laboratory helped make most of this work possible and Yvonne Sadovy, George Mitcheson, staff of the Swire
we would like to thank those who helped us early on. Marine Laboratory. Bahrain: Jassim al Qasser, Roger
These friends include: Donald Benjamin, Mardi Bren, Uwate, staff of the Directorate of Fisheries.
vi
United States: Bruce Carlson, Marjorie Awai, Phil Scott Johnson- pages: 140, 160, 161a,b, 243c, 244c; pic-
Helfrich, Jack Randall, Chuck Nicklin, Bonnie Pelnar, tures: 655, 633, 634, 674, 682, 743, 744, 954, 955, 971,
Ali Baradar, Nick Holland, Bill Hamner, Fritz Schmitz, 998, 1006, 1062, 1067, 1071, 1166, 167, 1169, 1171,
John Faulkner, Mike Cameron, Lolita De Palma, Bertha 1174, 1179-84.
Cutress and Steven Cross. Guam: Valerie Paul, Gustav
Pauley, Charles Birkeland and Terry Donaldson. Mike Severens- page 206d; pictures: 1053, 1060, 1061,
1065, 1066, 1068, 1070, 1230.
Many of the identifications in this book were made from
specimens or photographs by specialist taxonomists. We would also like to thank Mandy Etpison, who drew
Without their efforts, no order or understanding of the the line illustrations for this book.
organisms in the sea would come from the chaos super-
ficially apparent when so many forms of life co-exist. Bonnie Pelnar showed us how to print this book in the
We thank the following for their help and identifica- United States using digital images. Without her advice
tions: and assistance this project might never have been com-
Sponges- Michelle Kelly-Borges, Ph.D. and John pleted and we owe her our sincere thanks. John Hull,
Hooper, Ph.D. Dan Weber, Arlene Mercado, Clayton Watson and the
Hydroids and Jellyfish- Dale Calder, Ph.D. and Ron staff of Mybar Printing, Irvine, California contributed
Larson, Ph.D. significantly to the planning, quality and completion of
Soft corals- Katherine Muzik, Ph.D. (Octocorals), Gary this book. Steve Miller of Miller Imaging, Santa
Williams, Ph.D. (Sea Pens) Monica, California helped us scan the original photo-
Hard corals- Gustav Pauley, Ph.D. graphic slides to Kodak Photo CD. The entire book was
Sea Anemones- Hajo Schmidt, Ph.D. printed from digital images using the latest printing
Flat Worms- Leslie Newman, Ph.D. techniques. The traditional printing process is notorious
Polychaetes- Leslie Harris, Ph.D., Kirk Fitzhugh, Ph.D. for producing waste that is harmful to the environment.
Molluscs- Lori J.B. Colin, Terry Gosliner, Ph.D. and Mybar Printing is to be commended for its environmen-
Scott Johnson, Roger Hanlon, Ph. D. tally safe printing procedures. They utilize a number of
Arthropods- Lou Eldredge, Ph.D., Marivene Manuel and techniques to reduce many hazardous by-products of the
William Newman, Ph.D. printing process: soy based inks, alcohol-free printing
Lophophorates- William Banta, Ph.D. (Bryozoa) and aqueous coating. Soy-based oils are renewable,
Echinoderms- Loisette Marsh, Ph.D., Charles Messing, biodegradable and have lower solvent emissions than
Ph.D., Gustav Pauley Ph.D., Gordon Hendler, Ph.D. petroleum inks. Alcohol-free printing allows the water
Urochordates- Francoise and Claude Monniot, Ph.D.s used in the printing process to be recycled. Unlike var-
nish, aqueous coating is biodegradable, renewable, non-
We would like to thank the following for their review of volatile and non-hazardous.
the manuscript: Nick Holland, Bill Hamner, Jim Miller,
Gale Anne Hurd, Michelle Kelly-Borges (sponges), Lori We welcome notification of errors detected in this book.
J.B. Colin (molluscs) and Charles Messing (crinoids).
vii
Introduction
“The only solid piece of scientific truth about which
I feel totally confident is that
we are profoundly ignorant about nature.”
This book covers the tropical Pacific, that vast region running from
Hawaii and French Polynesia in the east to the Philippines, Papua New Guinea and Indonesia
in the west. It includes, in addition, all of Micronesia (Marshall Islands, Gilbert Islands,
Caroline Archipelago, Nauru and the Marianas), the remainder of Melanesia (Fiji, Solomon
Islands, Vanuatu and others) and the remainder of tropical Polynesia (Samoa, Tonga and oth- 1
ers). This is the area of the highest marine biodiversity on earth and that diversity is greatest
in the shallow waters of the region. It is also an area where new scientific discoveries occur
almost every day. The last great undefined marine faunas (primarily invertebrates) can be
found in the western Pacific and Indian Ocean, what is collectively called the Indo-west
Pacific. Species still undescribed to science can occur in snorkeling depths literally right off
the dock or beach.
Above- The Pacific Ocean covers nearly one third of the globe as is apparent in this view centered overthe Marshall Islands. The
approximate limits of the tropics, where watertemperature is not below 68oF (20oC) forprolonged periods, are indicated by dashed
lines. The directions of major currents are indicated by arrows. Equatorial currents run east to west approximately ten degrees
above and below the equator, while equatorial countercurrents run west to east nearthe equator. This diagram shows warm water
circulation in the central/Western Pacific.
Current runs east to west south of the equator, and turns Above- This aerial view of Pakin Atoll shows the elements of
and splits when it reaches the Papua New Guinea- a coral atoll, the ring of barrier reef with islands surrounding
a lagoon and the deep ocean outside. Pakin lacks a deep-water
Solomon Islands region. Part of this water is directed
pass.
back east at the equator to join the Equatorial Counter
Current, while part flows south along the east coast of
Australia.
3
To complete this generalized current pattern for
the region, many other lesser currents occur on the
periphery of the wide Pacific. Water moves east
through the Straits of Malacca, between the Malay
Peninsula and western Indonesia (Sumatra), turning
north between Borneo and Sulawesi. Similarly, water
moves eastward through southeastern Indonesia and
passes through the Torres Strait between Australia and
New Guinea before reaching the Coral Sea. Currents
are quite complicated within the northern islands of
Papua New Guinea and still poorly understood. In
addition to the wind driven currents, tidal fluctuations
also play a major role in local current patterns.
Above- Onang Island in Chuuk is a typical low coral island on the barrier reef of a coral atoll. Such islands
are no more than several feet above sea level and are made of reef rubble and sand thrown up by storms. Soils
on such islands are usually poor, but the coconut palm thrives in this environment.
The calcareous algae Halimeda produces large amounts of calci- This photo of three species of hermatypic corals growing right
um carbonate in the form of plate-like skeletal material. These next to each otherillustrates the intense competition for space on
plates are a major component of reef sediment in the tropics. the reef. The corals maintain these boundries either through use
of chemicals or nematocysts.
are adapted to filter feeding and have excellent means also produce large amounts of calcium carbonate on or
for attachment inhabit these areas of heavy current. near reefs. The green calcareous algae Halimeda pro-
duces huge amounts of flake carbonate material which
When most divers or snorkelers think of the forms its own habitat, “Halimeda beds”, in and around
tropical Pacific, coral reefs immediately come to mind. reefs. The coralline red algae are generally pink in
In ecological terms, coral reefs are referred to as a com- color and they are important binders of fine reef sedi-
munity. Communities are an assemblage of plant and ment at all depths. In very shallow water coralline red
animal populations occupying a given area. Marine algae form a ridge of calcareous material which helps to
communities of the shallow tropical Pacific can be break the force of waves on the reef. Calcium carbon-
grouped into several different types, based either on a ate is produced in lesser amounts by shelled creatures,
5
dominant organism (seagrass bed, coral reef) or some such as molluscs and echinoderms.
conspicuous non-biological component within that
community (rocky shore, sand slope, mud flat). These The development of a coral reef is a constant
somewhat arbitrary divisions are a simplistic but useful interplay between growth and destruction of the reef
way to group communities. Still it is important to structure. Growth rates of reefs are the rate at which the
remember that such labels give no indication of the calcium carbonate matrix of the overall reef increases
overlaps, relationships and differences which occur towards the surface. Such rates have been estimated,
among the organisms identified with these particular from a variety of sources, to arrive at a generalized fig-
communities. ure on the order of one half of an inch per year.
Individual corals can grow at rates much higher than
Among shallow tropical communities, coral that figure, but these rates do not apply over large areas
reefs are the most complex and species diverse. The of reef. The fine branching Acropora corals may
corals that build reefs (called hermatypic or stony increase the length of their branches as much as 10
corals) are generally colonies of hundreds or thousands inches (25 cm) a year while a table-like coral Acropora
of indvidual coral polyps. Such corals produce the hyacinthus grows outward about six inches (15 cm) a
framework of the reef. These corals thrive in relatively year. Reef growth is not just a factor of how fast corals
shallow, clear water that allows light penetration. The can grow, it is controlled by many factors; for example,
light enables photosynthesis by the symbiotic algae death of individual coral polyps, storms, predation by
(zooxanthellae) contained within the coral polyps. The other animals (e.g. crown of thorns starfish) and coral
algal cells provide essential nutrients to the polyps that boring organisms which weaken the calcium carbonate
help the coral polyps to grow and deposit calcium car- matrix of individual coral colonies.
bonate in sufficient quantity to build the massive skele-
tal material that is the reef structure. There are other There is fierce competition for space on most
corals, known as ahermatypic or non-reef building coral reefs, both among the corals themselves and with
corals, which lack zooxanthellae and do not produce a other attached reef organisms. Stony corals fall into
significant amount of calcium carbonate. Ahermatypic three general types; the branching corals, massive
corals are usually found in shaded habitats where they corals and plate corals, each with particular advantages
still play an important role in the reef ecosystem. Other when it comes to occupying space on the reef. The
organisms such as calcareous algae and the foraminifera branching corals, typified by the genus Acropora, are
Above- Idealized cross section of a typical Pacific Ocean barrier reef, near a high island, from shore to the drop off. There are dis-
tinct zones along this transect, with mangroves and beach inshore, a lagoon with seagrass coral patches and pinnacles, then the bar-
rier reef with reef flat and fore reef slope.
the fastest growing, but are relatively fragile, prone to
destruction by storm waves and surge. The plate corals,
found in many genera, are slower growing, but their flat-
tened or convoluted plate structure allows them to capture
light in deeper areas and to grow over nearby corals, cut-
ting off their competitors’light. Some, such as species of
Turbinaria, are well adapted to low light and high levels
of siltation, and the lushness and density of their growth
6 in murky water is often stunning. The massive corals, the
brain and star corals, form large heads and clumps reach-
ing ten feet high, but these are the slowest growing. They
have the advantage, though, of being the strongest and
most resistant to storm damage. In clear tropical water
corals dominate other sessile invertebrates and cover
large areas of available substrate.
known as the “spur and groove” zone, a series of alter- At the point where the spur and groove meets
nating rocky fingers and channels perpendicular to the the reef flat on windward reefs, there is an exceptional-
reef face. The spur and groove zone is actually pro- ly interesting community, the coralline algal ridge. The
duced in response to wave action, both through active coralline algal ridge is an assemblage of various
8 growth of the reef in response to wave action and by coralline red algae. They occur in what is usually the
erosion from sand carried by the waves. Spur and most turbulent water found at coral atolls. The algae
groove formation serves to dissipate the energy of grow as a thin crust or as small fan-like forms. They
incoming ocean waves by breaking up the momentum deposit calcium carbonate which produces the slightly
of the water in the wave. It has been reported, for elevated algal ridge. A well developed algal ridge starts
example, that the spur and groove system on the wind- on the shallow tops of the spurs and continues onto the
ward (eastward) side of Bikini Atoll dissipates 95% of reef flat, with the various species of corallines having
wave energy, with the remaining 5% going to wave distinct zonation within the ridge area. The healthy
pumping and maintenance of water level on the reef coralline algal ridge produces a sponge-like rock struc-
flat higher than that of the ocean. ture with channels, cavities and chambers honeycombed
into the entire ridge. Animals found here are well adapt-
ed for hanging on in the face of extreme wave condi-
tions. Some, such as the spiny lobster Panulirus penni -
cilatus, may hide in the reef during the day and then
move onto the reef flat at night to feed. Others, such as
the sea urchin Echinometra mathei, dig deep grooves
into the rock on the spurs where they are protected from
wave action which would otherwise rip them away from
the bottom.
Tropical Pacific Invertebrates is designed to serve as a general field guide for those marine invertebrates seen or found by
divers, snorkelers, naturalists and others in the reefs and shallow water marine environments of the tropical Pacific. We hope
the book will prove useful to students of marine biology and ecology, particularly within the region we are covering. We have
attempted, within limits, to include most of the common organisms that would be encountered, plus a number of the rarer
species which are distinctive and interesting. We have tried to emphasize those groups of invertebrates, such as the sponges
and the ascidians, which, because of difficulties in identification, have been treated only superficially in previous popular guides
to the Pacific marine fauna. More familiar groups, such as the stony corals and gastropod molluscs, have been the subject of
other excellent publications (some of which are referenced at the end of this book) and we have not attempted to include all
possible species. For those groups we have chosen to present a selection of typical species which would enable the reader to
become familiar with the broadest possible range of tropical marine invertebrates.
Each major division of the animal kingdom, typically a phylum, is included as a separate section and these are presented in
approximate phylogenetic order. Each section has two parts: an introduction about the general features of the group, followed
by photographs and notes about selected species within that division. The introductory text describes important features of the
phylum or division, such as their form (morphology), diversity, reproduction and feeding. Various species are mentioned in
these introductory remarks and some photographs are included which illustrate characteristics described. Text remarks are not
designed to serve as all inclusive descriptions of each group, but rather to provide a brief introduction to animals concerned,
pointing out those things that an observant diver might notice. Readers can look for information at any level they desire in
order to gain additional insights into the organisms.
The photograph and notes section illustrates and where relevent, comments on the species selected within that group. Below
each photo is a reference number, the scientific name and the country where the photograph was taken. Detailed notes on the
taxonomy of organisms and locations of the photographs, indexed by the photo number, are included on the facing page. In
many cases the illustrated specimen is backed up by specimens preserved in museum collections and it is these specimens, along
with the study of such specimens by a taxonomist, which allow us to assign a scientific name, even if only a genus, to most of
the photographs. In some cases, however, it is impossible to determine what species, or even to what genus, the specimen prop-
erly belongs and this is noted by the absense of a generic or specific name. Quite a number of the species illustrated are not
yet described to science, although their existence is known to taxonomists and descriptions are in the process of being written.
The name, taxonomy and locality is followed by information on natural history, geographic ranges, similar species or other
interesting notes. Descriptive notes for photographs vary greatly in length, largely related to the amount of interesting relevent
knowledge of the organism. Zoogeographic range information is included when it is of interest, but in many cases we can pro-
14 vide no further information regarding the geographic distribution of a species beyond the site records provided by the pho-
tographs. We hope the photo notes are an interesting blend of scientific information and personal experience. Each picture has
its story, and some of these are included in this section.
In general, most species have been illustrated with a single photograph. Many invertebrates, such as the sponges, show con-
siderable individual and environmental variation and certain examples may look considerably different from the ones we have
illustrated. Our photographs attempt to show a typical individual, group or colony in its natural habitat; in a limited number of
cases we have provided more than one photo to show radically different forms, or changes during growth, of a single species.
Scientific names are binomials, or two names, written in italics. The first is the genus (the plural is “genera”), always capital-
ized, while the second is the specific name, which is not capitalized. The genus is a general group into which one or more
species fall; a genus may have only a single species within it (monotypic) or many species, but each species within the genus
has in common the various characteristics which define the genus.
Some of the species included here are common and well known, their scientific names dating back to Linnaeus and other later
authors. Ascribing what we believe to be the correct scientific names to such species is simply a matter of referring to readily
available scientific literature which contains authoritative treatment of the particular group in which the species occurs. Other
species are much more difficult to identify. Where the letters “cf.” (from the Latin confer) appear between the generic and spe-
cific names, this means the species looks like, but is not necessarily identical with, the particular species named.
Even with the best taxonomy and taxonomists available, there are still many questions regarding the identification of inverte-
brates pictured. Many of the species illustrated here will prove to be undescribed to science, i.e. they have never been given a
scientific name with an adequate description of the species. For others, even a specialist cannot be absolutely certain what a
particular specimen is, even with color photographs and a specimen in hand. They may feel it is close to a particular species,
but are uncertain whether it is absolutely the same.
Relatively few Pacific marine invertebrates have widely accepted common names. Unless a species has a well known and wide-
ly accepted common name, we have avoided using any common names. Many of the common names come from the marine
aquarium trade and these are used whenever possible. In a few cases, we have used new common names where they were felt
to be particularly appropriate.
Photographs were generally taken using 35 mm single lens reflex cameras in underwater housings with submarine strobes for
illumination. In nearly all cases the photographs are of undisturbed animals taken against their natural backgrounds. We have
Scientific Naming and Taxonomy
When described (named), each species of animal or plant is given a two part scientific name. This consists of a genus and specific name, such
as Conus geographus, and is italicized in print. Species are placed into taxonomic categories, each a subunitof the category above it. The major categories
break down as follows: Phylum - Class - Order - Family - Genus - Species, and there can be intermediate categories of these, such as subclass or super-
order. The idea is to make natural and logical divisions between categories which reflect real biological differences
Taxonomy is the science of identifying, naming and studying the relationships between organisms. Ideally taxonomy creates a system which
mirrors the relationships between organisms and how they evolved. Many taxonomists work for museums or universities. While the public is familiar with
the exhibit sections of large museums, such as the Smithsonian Institution, that is only a small part of what these institutions do. They also maintain (curate)
large research collections, which are usually closed to public access. It is these collections and the experience and training of the taxonomists that allows
them to determine if an organism is already known, its proper name and to prepare a scientific description of the organism, if it does not already have a sci-
entific name. The descriptions of new organisms are usually published in specialty scientific journals or in books covering a particular group of animals.
In addition to their general collections, museums also maintain collections of type specimens-those specimens on which the original description of a given
species was based.
The scientific names of many reef organisms are occasionally changed. This is not due to the whim of the taxonomist, but because recent study
has shown that the species was given a scientific name by an earlier author. Unfortunately, some Pacific invertebrates have been described several times,
each subsequent author either not being aware of the previous description or thinking that their specimens represented a new species. In earlier times the
type specimens were often not adequately preserved, lost or not sufficiently large to make it easily apparent what the species named represented. Workers
in earlier centuries did not have the advantages of diving, photography, computers and well maintained collections to document their work and it is easy to
understand how confusion occurred with variable species. Where there is more than one scientific name for a species, the oldest name has priority and later
names become synonyms and are not used for the species.
tried to choose photographs which are not only a portrait of the species concerned, but which also display some aspect of the
biology of the creature. The notes included for each photograph attempt to describe the biological aspects illustrated in the
photographs.
Photographs were taken in many different areas, but readers will notice that a number of locality names appear regularly.
These are usually localities where we have been able to live or otherwise spend a considerable period of time, often due to
the presence of a field marine research facility nearby. These include Madang and Port Moresby in Papua New Guinea, Palau
and Chuuk in the Caroline Islands, and Enewetak Atoll in the Marshall Islands. Visits for shorter periods were made to other
locations, which because of their species richness were productive areas for photographing marine invertebrates. 15
One sidelight on the localities is that quite a number of the photographs included here were taken at Enewetak Atoll, the for-
mer nuclear test site, in the Marshall Islands. The term “former nuclear test site” has connotations which would imply that
the ocean bottom is a nuclear desert, devoid of life. Exactly the opposite is true, since Enewetak has been essentially unfished
and its marine life unexploited since the cessation of nuclear testing in 1958. All the organisms that would have been exploit-
ed as food by a local Marshallese population have flourished in the absence of human predators, and have approached levels
similar to what would probably occur if man did not exist at all.
It is an intended purpose of this book to discourage the thoughtless collection of invertebrates, or any other marine life. The
coral skeleton broken off today is usually a forgotten piece of trash a year later. Better to leave them where they are able to
grow and reproduce, to fulfill their part in the natural scheme of things. While we discourage frivilous collecting, individual
specimens are required in the pursuit of scientific studies. Such collections form the basis of much of our knowledge of
marine biological science and we owe a debt to the collectors of past centuries who have made much progress towards our
goal of understanding the world around us. These efforts are far from finished, as even a brief perusal of this book would
indicate, and there is still much left to be learned about the identification of most groups of marine organisms.
The Coral Reef Research Foundation (CRRF) is a non-profit organization founded in 1991 and dedicated to research and edu-
cation concerning coral reefs and other tropical marine environments. CRRF operates two small marine research laborato-
ries in Chuuk, Federated States of Micronesia and Koror, Republic of Palau. It also conducts basic marine research through-
out the western Pacific region. Laboratory projects include elucidation of the marine fauna and flora of the region, and base-
line monitoring and mapping of the marine environment.
For further information regarding CRRF and its programs please contact:
Sponges *
The biology of sponges is less well known than that of other organisms. Sponges lack the
muscles, nerves and the body organs with which we are all familiar. They have no digestive cavity or 17
mouth. Biological interactions in the sponge take place at the cellular, rather than the level of organs.
Sponges can be thought of as communal associations of cells, loosely arranged to form a net-
work of inhalent and exhalent canals. The inhalent canals originate as small pores (ostia) on the outer
surface of the sponge and lead to spherical chambers. These chambers are lined with choanocytes, cells
with whip-like flagella that beat in rhythmic waves to pump water through the body in one direction.
Water carried into the sponges is filtered for food particles and oxygen and is then expelled through
one or several exhalent pores (oscules). Complexity of the canal
structure most often increases with sponge size. Sponges vary
greatly in growth form and size from thin encrusting sheets a frac-
tion of an inch thick, to large barrels or vases, which may grow to
several feet in height and attain a volume of about two cubic yards
(about the size of a small cement mixer). From a rather simple
body plan, sponges have evolved myriad shapes, sizes, and colors.
Opposite- This blue vase sponge, Cribrochalina olemda, is common on reefs throughout much of the region covered
in this book. The sponge is frequently found on inshore patch reefs.
18
These four photographs illustrate some of the variation in growth form and habitat among the sponges. Above left - This encrust-
ing sponge Aplysilla sp. shows a beautiful pattern of dendritic excurrent water channels with a single oscule. The incurrent ostia
are scattered over the entire surface of the sponge. Below left - This sponge, from Fiji, infiltrates the skeleton of living coral, only
the oscules are visible. Above right - This barrel sponge, Xestospongia, is one of the largest sponges on coral reefs. It is found in a
variety of habitats and colors. Below right - This unidentified sponge grows in small clumps, exposed on the reef. Different inter-
nal and external colors and textures are common in many sponge species.
Three classes of sponges are recognized on the eral different origins and do not represent a natural tax-
basis of their skeletal components. The Class onomic grouping. As their name implies sclerosponges
Demospongiae is by far the largest and most diverse are very hard and stony. They typically inhabit caves
group of sponges, it includes the familiar tube, vase, and shaded ledges on coral reefs. These sponges were
barrel, and fan sponges. Demosponges are characterized believed to be extinct for millions of years until their
by their skeleton, which consists of spongin fibers and rediscovery in the 1960’s.
siliceous spicules. In some demosponges one or both of
these skeletal components may be absent. The Class Some sponges occur in freshwater habitats,
Calcarea has calcium carbonate spicules in the mineral however most sponges are marine. They are found at
form calcite. Calcareous sponges are small and delicate all latitudes in the marine environment, but reach their
with a crunchy texture, due to a lack of spongin and col- greatest diversity on coral reefs in tropical seas.
lagen, they occur in limited numbers in all marine envi- Sponges are also abundant in seagrass beds, mangroves,
ronments. In the tropics, calcareous sponges are most and other environments.
often found on vertical reef faces. The Class
Hexactinellida, commonly called “glass sponges”, has No one knows exactly how many species of
distinct siliceous spicules with six rays. They are sel- sponges there are, but estimates range from 5,000-9,000
dom found at depths less than several hundred feet and species. As a general rule, each time scientists closely
we have not encountered glass sponges within diving investigate the sponge fauna of a certain region they
depths in the tropical Pacific thus far. A former fourth find it to be more diverse than originally thought. It is
class, the Sclerospongiae (sclerosponges), has been certain that in the eastern and central Caroline Islands of
divided among the first two classes. The species con- Micronesia there are at least 300-400 species of
sidered to be sclerosponges are known to have had sev- sponges. Palau, in the western Carolines, probably has
600 or more species. In areas such as Papua New
Guinea and Indonesia, the total number of species is
1,000 or more. Eventually we will probably find those
figures to be conservative underestimates. In addition to
the increase in diversity as one moves west across the
Pacific, some differences in species numbers can be
attributed to available habitats, i.e. the greater the diver-
sity of the habitats, the greater the number of species.
Across Micronesia, the lowest diversity in sponges is
found in the clear water atolls of the eastern Marshall
Islands. These atolls generally lack seagrasses and
mangroves and do not have high islands which enhance
the diversity of habitats and nutrient enrichment in their
lagoons. Areas with high lagoon productivity and many
different habitats, such as Pohnpei, Chuuk and Palau,
have greater sponge diversity.
The Phylum Porifera provides one of the great This close up view shows the multiple excurrent oscules of
challenges to marine taxonomists. The highly diverse Spheciospongia vagabunda, a widely occuring sponge in the
western Pacific and Indian Ocean faunas remain the tropical Pacific. Water is expelled through these openings
object of study where the basic elucidation and descrip- after it has been filtered in the body of the sponge.
tion of species is still far from complete. Consequently, Sponges reproduce both sexually and asexually.
although we have employed the best taxonomic identifi- Many individuals are hermaphrodites, producing both
cations available at the time of publication, a number of eggs and sperm. In sexual reproduction, sperm are
the species names will certainly change. The higher tax- released into the water; eggs may be released
onomy of sponges (Families, Orders and Classes) is also (oviparous) to undergo fertilization and development in
in a fluid state and many of the assignments to family or the water or retained and fertilized inside (viviparous)
order will change in the future. New discoveries by the sponge. In many species there is synchronous 19
sponge biologists have resulted in the continual revision release of sperm and eggs triggered by daily and lunar
of groupings to accommodate new information. The rea- cycles. Fertilized eggs develop into larvae. The larvae
sons for the taxonomic uncertainties are many; the char- swim or creep along the bottom for periods of up to sev-
acters used to differentiate many sponge species are few eral days, this aids dispersal of the offspring. Asexual
and often variable due to environmental and other fac- reproduction in sponges happens through a variety of
tors. Indeed, the concept of what constitutes a species of methods, including budding, fragmentation and a resting
sponge remains a matter of considerable investigation. stage known as the gemmule. In many cases, sponges
Above - The external color of many sponges varies with exposure to light. Usually a species that changes color is darker when liv-
ing in areas exposed to light and pale or white when living in dark areas. One theory for this color change is that the dark color is
due to the colorof photophilous micro-organisms living on the spong;, in the absence of light the micro-organisms leave the sponge,
and the dark color fades. The photograph on the left shows variation in color with exposure to light in Aaptos sp. This sponge is
living in a partially lighted area, hence its mottled appearance. The photograph on the right shows a normally purple colored sponge
living in a cave. It is almost completely white.
Above - Sponges reach theirgreatest diversity and abundance in tropical seas. In these rich habitats, sponges must compete with
otherbottom-dwelling invertebrates forattachment sites and hard surfaces. In the picture there are at least eight different species
of sponges living on, and competing for, the same two square foot area of wall. Many sponges succeed in obtaining open space
through “chemical warfare”, producing complex chemical weapons in the course of daily metabolism.
20
can be cut into pieces and each will reorganize itself to
survive as a separate individual. This phenomenon
forms the basis of the culture of the commercially valu-
able “bath sponges”.
While the majority of sponges will probably The sponge in the picture can exhibit several different growth
not harm humans if handled, there are a number of forms, even within the same habitat. Three growth forms are
species which are definitely irritating to human skin. visible in the photograph; erect columns, encrusting and small
The irritation is the result of chemicals, spicules or fans. This is one of the reasons many sponges are so diffcult to 21
both, and individual susceptibility varies greatly. identify in the field.
Sponges of the genus Tedania have the well earned
common name of “fire sponges”. In many sponges
sharp, spiny spicules easily penetrate skin and cause
severe pain, irritation and swelling. We have tried to
indicate in the photo notes a few species which we
know to be particularly irritating, but this list is far from
complete. There are many sponges among those illus-
trated which will prove to be irritating, so caution is
important. In general, it is simply best to leave sponges
alone.
23
7- Cinachyra schulzei * Papua New Guinea * 8- Paratetilla bacca * Palau
9- Craniella abracadabra * Federated States of Micronesia 10- Paratetilla lipotriaenosa * Federated States of Micronesia
11- Cinachyrella sp. * Papua New Guinea 12- Ancorina acervus * Federated States of Micronesia
brown internally and is stiff with lots of spicules. A given individual can be
quite large and typically this sponge “cements” rocks together, filling in the
spaces between large boulder sized pieces of reef rubble. A stony coral of the
genus Pocillopora is below the sponge. Reddish coralline algae occurs
above it.
15- Rhabdastrella sp. * Papua New Guinea 21- Thrombus sp. * Thrombidae * Astrophorida * Papua New Guinea *
New Britain * Agu Reef * 93 ft (28 m). This strange sponge looks as if
someone stretched a piece of rubber sheet over a rock. The area of reef where
this sponge occurred was quite silty. The genus is not particularly well
known, but has been recorded previously from Vanuatu, eastern Australia, the
northeastern Atlantic Ocean and Caribbean Panama.
25
19- Melophlus sarasinorum * Guam 20- Geodia sp. * Papua New Guinea
21- Thrombus sp. * Papua New Guinea 22- Chondrilla australiensis. * Papua New Guinea
23- Cliona cf jullieni * Papua New Guinea 24- Cliona sp. * Palau
time excavating chambers and greatly weakening the structure of the reef.
This is particularly true for those coral colonies where clionids bore into the
base, reducing the strength of the colony to the point it is easily toppled by
storm waves or other physical force. A characteristic of many clionid sponges
is the presence of raised sucker-like sieve plates seen on the surface of this
sponge.
27
31- Desmapsamma sp. * Philippines 32- Higginsia anfractuosa * Palau
33- Acanthochaetetes wellsi * Federated States of Micronesia 34- Spheciospongia vagabunda * Papua New Guinea
29
43- Theonella sp. * Indonesia 44- Theonella sp. * Papua New Guinea
47- Theonella sp. * Philippines 48- Agelas sp. cf clathrodes * Federated States of Micronesia
species is sometimes called Agelas mauritania , which is incorrect, Agelas
clathrodes is an earlier name. The sponge is undescribed but very closely
resembles the Caribbean Agelas clathrodes, hence the comparison in the
species name above. Either way, this massive orange sponge is hard to con-
fuse with anything else. It can be as much six feet or more across. The mor-
phology varies greatly, from the large fan-like structures commonly known as
“elephant ear sponges” to small encrusting individuals with no particular
form. The color and the surface texture varies little with size. The pho-
tographed individual has a white didemnid ascidian growing on it, a common
occurrence.
52- Coelocarteria singaporense * Papua New Guinea 53- Zyzzya sp. * Federated States of Micronesia
54- Clathria sp. * Federated States of Micronesia 55- Monanchora ungiculata * Papua New Guinea
31
56- Crella sp. cf. calypta * Federated States of Micronesia 57- Neofibularia hartmani * Fiji
58- Iotrochota sp. * Federated States of Micronesia 59- Iotrochota sp. * Federated States of Micronesia 60- Autletta sp. * Papua New Guinea
33
67- Clathria sp. * Papua New Guinea 68- Clathria sp. * Federated States of Micronesia
69- Clathria basilana * Federated States of Micronesia 70- Clathria mima * Federated States of Micronesia
71- Clathria vulpina * Federated States of Micronesia 72- Echinochalina intermedia * Indonesia
known from northeast Australia, New Caledonia, Indonesia and Micronesia.
35
79- Echinodictyum asperum * Papua New Guinea 80- Echinodictyum mesenterinum* Papua New Guinea
81- Raspailia nuda * Indonesia 82- Raspailia wilkinsoni* Papua New Guinea
83- Rhabderemia sorokinae * Palau 84- Acanthella cavernosa * Federated States of Micronesia
85- Acanthella sp. * Axinellidae * Halichondrida * Papua New Guinea *
Kavieng * 70 ft (21 m). This is another axinellid sponge, but with a chara-
teristic shape. In this case the sponge has an almost tubular body with one or
two oscules at the end, as is shown in the photograph. It is not very common,
and is presently known only from Papua New Guinea.
89- Phakellia sp. * Papua New Guinea 88- Phakellia sp.* Federated States of Micronesia
90- Auletta sp. * Papua New Guinea 91- Axinosa sp. * Papua New Guinea
37
92- Axinella sp. * Papua New Guinea 93- Cymbastella sp.* Federated States of Micronesia
94- Pseudaxinella sp. * Papua New Guinea 95- Stylotella aurantium* Palau
96- Phakellia sp. * Indonesia 97- Reniochalina sp. * Papua New Guinea
96- Phakellia sp. * Axinellidae * order* Indonesia * Manado * 110 ft (33
m). This axinellid sponge is tubular, and thus is superficially similar to mem-
bers of the genus Auletta. However, it has a rough outer surface, while
Auletta sp. are mostly smooth. The sponge is found on deep reef dropoffs.
98- Stylissa flabelliformis * Federated States of Micronesia 100- Axinyssa aplysinoides * Halichondriidae * Halichondrida *
Federated States of Micronesia * Chuuk Atoll * 80 ft (25 m). This
lagoon sponge is black and exudes abundant mucous. The sponge is lumpy
and can form fingers. It has characteristic ostia as shown in the photograph.
In Chuuk Atoll it occurs in the lagoon from 50-120 feet deep. A sea cucum-
ber lies next to the sponge.
101- Stylinos sp. * Federated States of Micronesia 102- Katiba milnei * Federated States of Micronesia
103- Myrmekioderma sp. * Guam 104- Myrmekioderma sp. * Federated States of Micronesia
39
105- Reniera osiros * Federated States of Micronesia 106- undescribed genus * Indonesia
107- Stylotella aurantium * Federated States of Micronesia 109- Callyspongia sp. * Papua New Guinea
One of the most common and easily recognized sponges on reefs in the
region. Although S. aurantium is quite variable in shape and surface texture,
it is nearly always the same flourescent orange yellow color shown in the pho-
tograph. It is soft and easily compressed, like “soggy bread”. Known
throughout the region from reefs, seagrass beds and mangroves.
110- Callyspongia aerizusa * Marshall Islands 110- Callyspongia aerizusa * Callyspongiidae * Haplosclerida * Marshall
Islands * Kwajalein Atoll * 80 ft (25 m). This well known species is a dis-
tinctive bluish green color, but can occasionally be light brownish-pink. The
species is typically tubular but can also form huge fans. Specimens from
deeper algal flats have thinner, more elongate tubes.
113- Amphimedon sp. * Indonesia 114- Callyspongia sp. * Papua New Guinea
115- Callyspongia sp. * Federated States of Micronesia 116- Callyspongia sp. * Papua New Guinea
41
117- Dactylia sp. * Papua New Guinea 118- Euplacella sp. * Indonesia
119- Euplacella sp. * Indonesia 120- Siphonochalina sp. * Papua New Guinea
121- Auletta sp. * Papua New Guinea 122- Adocia sp. * Papua New Guinea
volcanic island. The barrier reef around Bagabag experiences strong currents
and sometimes high waves. Despite the surge, the barrier reef at Bagabag is
rich with invertebrates which are well adapted to hanging on in the ocean
swell.
126- Sigmadocia amboinensis * Indonesia 127- Nara nematifera * Federated States of Micronesia
128- Haliclona cf. coerulescens * Federated States of Micronesia 129- Haliclona cymaeformis * Palau
43
130- Haliclona koremella * Indonesia 131- Acervochalina velinea * Palau
45
140- Haliclona sp. * Federated States of Micronesia 141- Amphimedon sp. * Papua New Guinea
142- Amphimedon sp. * Papua New Guinea 143- Amphimedon sp. * Papua New Guinea
144- Cribochalina sp. * Federated States of Micronesia 145- Cribrochalina olemda * Palau
148- Gelliodes fibulata * Niphatidae * Haplosclerida * Papua New Guinea
* Port Moresby * Lion Island * 40 ft (12 m). The spicules in this well
known species of Gelloides fuse together to form what are probably the
sharpest, stiffest spines found among tropical Pacific sponges. We like to call
these the “cactus” sponge.
47
152- Petrosia capsa * Papua New Guinea
153- Petrosia sp. * Indonesia 154- Petrosia sp. * Papua New Guinea
155- Petrosia sp. * Papua New Guinea 156- Strongylophora sp. * Papua New Guinea
in lagoonal areas of Micronesia. It takes a wide variety of forms from thin
encrusting mats, to encrusting with gnarled projections or as ridges arising
from a basal mass. The dark brown color is generally constant, despite the
large variations in shape. The small branches are brittle, and when broken
reveal a cream colored interior.
160- Pellina sp. * Federated States of Micronesia 161- Xestospongia sp. * Papua New Guinea
162- Xestospongia sp. * Papua New Guinea
49
164- Xestospongia testudinaria * Papua New Guinea 163- Xestospongia pacifica * Federated States of Micronesia
165- Xestospongia? sp. * Papua New Guinea 166- Vagocia sp. * Palau
51
175- Carteriospongia contorta * Indonesia 176- Coscinoderma sp. * Papua New Guinea
177- Dactylospongia sp.* Federated States of Micronesia 178- Dactylospongia sp. * Indonesia
179-Coscinoderma mathewsi * Federated States of Micronesia 180- Coscinoderma mathewsi * Papua New Guinea
184- Spongia sp. * Spongiidae * Dictyoceratida * Philippines * Santa
Rosa 6 * 165 ft (50 m). This sponge, again nominally S. officinalis, appears
quite different, being a brilliant white, perhaps in response to the depth where
it was photographed. A number of white synaptid holothurians (sea cucum-
bers) are on the outer surface of the sponge.
53
188- Hyrtios mela? * Federated States of Micronesia 189- Ircinia sp. * Federated States of Micronesia
190- Iricinia sp. * Philippines 191- Iricinia sp. * Papua New Guinea
192- Luffariella metachromia * Federated States of Micronesia 193- Luffariella variabilis * Federated States of Micronesia
195- Thorectandra sp. * Thorectidae * Dictyoceratida * Federated States
of Micronesia * Chuuk Atoll * barrier reef * 100 ft (30 m). These more
or less spherical sponges are distinctive from the superficially similar to “golf
ball” sponges. In Thorectandra the surface is hard, formed into ridges, with
much sand incorporated into it and with apical oscules. The illustrated
species is relatively common in the Caroline Islands on reefs at moderate
depths.
55
200- Dendrilla sp. * Papua New Guinea 201- Euryspongia sp. * Federated States of Micronesia
202- Dysidea avara * Federated States of Micronesia 203- Dysidea sp. * Federated States of Micronesia
204- Dysidea herbacea * Palau 205- Lendenfeldia complex * Federated States of Micronesia
* Kavieng * Dyaul Island * 66 ft (20 m). This species occurred along a
deep reef dropoff.
57
212- Hyrtios sp. * Federated States of Micronesia 213- Iotrochota sp. * Federated States of Micronesia
214- Aplysinella rhax * Papua New Guinea 215- Aplysinella sp.cf. strongylata * Papua New Guinea
216- Aplysinella sp. * Papua New Guinea 217- Pseudoceratina verongiformis * Federated States of Micronesia
Micronesia * Mortlock Islands * Ettal Atoll * 66 ft (20 m). This occurs
along overhangs of reefs in moderate depths in the Caroline and Marianas
Islands.
Class Calcarea
59
223- Dactylospongia sp. * Federated States of Micronesia
227- Clathrina sp. * Papua New Guinea 228- Clathrina sp. * Papua New Guinea
229- Leucetta primigenia * Leucettidae * Clathrinida * Indonesia * Biak
* 86 ft (26 m). This sponge is one of the largest calcareous sponges in the
Indo-west Pacific and can occur in both inshore and offshore reef areas. It
also occurs commonly on the wrecks in Chuuk lagoon. The brown exterior
and creamy interior is normal for the species. It is very spiculous and can
pierce skin with its large sharp spicules.
229- Leucetta primigenia * Indonesia 232- Leucetta avocado * Leucettidae * Clathrinida * Federated States of
Micronesia * Chuuk Atoll * Fourup Reef * 60 ft (18 m). This sponge is
common on reefs in much of Micronesia, but is not seen so often in other
areas such as Papua New Guinea. Small individuals, such as the two shown
in the photograph, are easily recognized; a lobate or conical shape with an
oscular opening and the outer surface mottled brown with green underneath.
Large individuals, as much as two feet in length, particuarly those in calm
water, can assume grotesque shapes with the mass of the sponge appearing
to “drip” down a vertical face. The outer surface can also become very dark
when exposed to abundant light. The internal color is always a lime green.
This sponge was also described as Leucetta avocado by deLaubenfels, but
since P. hererorhaphis is the earlier name (1884 vs. 1954), it has priority.
61
235- Leuconia palaoensis? * Papua New Guinea 236- Plakina sp. * Papua New Guinea
237- Sycon sp. * Papua New Guinea 238- Sycon sp. * Philippines
This photograph,
taken inside a cave at
a depth of 100 feet
nearCebu in the
central Philippines
reveals several species
of calcareous sponges
in theirnatural
habitat.
62
Phylum Cnidaria
Hydroids,
Jellyfishes, Corals,
Sea Anemones and
Black Corals
Cnidarians, also known as coelenterates, are arguably the most common
and conspicuous invertebrates found in shallow tropical Pacific waters because this
phylum includes the many species of corals which build coral reefs. With more than 10,000
species, the cnidarians are predominantly marine and reach a level of diversity and importance
in shallow tropical waters unequalled by any other phylum. They form the basis of many trop-
ical reefs ecosystems, but are also abundant in colder water. There are four classes: the
Hydrozoa (hydroids), Scyphozoa (jellyfishes), Cubozoa (sea wasps), and Anthozoa (corals,
63
corallimorpharians, sea fans, sea anemones, zoanthids and black corals), distinguished on the
basis of life history and morphology. They are united by certain characteristics: radial sym-
metry, a central mouth surrounded by tentacles, a single opening through which food is ingest-
ed and expelled (coelenteron), a jelly-like middle germ layer (the mesoglea), and intracellular
stinging structures called nematocysts.
Two body forms, the polyp and the medusa, are found in cnidarians; some species
having only one form while others have both. The polyp lives attached to the substrate, has a
fleshy body and an upward directed mouth surrounded by tentacles. The medusa is free swim-
ming; the body is a dome-shaped bell with the mouth underneath and tentacles arranged
around the margin. In the Hydrozoa, Scyphozoa and Cubozoa most species have alter-
Left- This vertical wall at Pescador Island near Moalboal in the central Philippines is covered with Cnidarians and
other invertebrates. The species diversity in this area is very high. A large soft coral is visible in the center, also vis-
ible are numerous orange colonies of ahermatypic coral.
Above left- This photo shows three polyps of the cubomedusae Carybdea alata in the process of metamorphosis. The individual at
the far right is almost ready to swim away as a small jellyfish. The polyp at the far left has not started to change and is still able
to produce asexual buds. Below left- This is a young cubomedusae, Carybdea alata, several days after it has been released. As
64 the jellyfish grows, the tentacles will increase in length and the bell will become transparent. Within a year the jellyfish will be
sexually mature. Above right- Life history of Aurelia. Sperm and eggs produced by mature medusae are released into the water
where fertilization takes place. The fertilized eggs develop into a planula larvae which attach to the substrate and develop into
polyps. The polyps grow, increase the numberof theirtentacles and begin to bud off secondary polyps. Many clone-like secondary
polyps are produced. After a period of about nine months some of the older polyps begin to produce medusae. Unlike cubome-
dusae where one polyp converts to one medusa, Aurelia polyps produce many juvenile jellyfish (ephyrae). There are numerous
variations of this life history within the Cnidaria, in many species, only medusae or polyps develop.
nate polyp and medusa stages. Members of the remain- ual coral colonies which coexist and form much of the
ing class, Anthozoa, exist only as polyps, either solitaryframework of most coral reefs. Those corals which con-
or forming colonies. Within the Anthozoa the sea tribute skeletal material to the overall framework of the
anemones corallimorpharians and cerianthids are soli- reef are called hermatypic corals. Hermatypic corals
tary, the stony corals and zoanthids have species which generally belong to the scleractinia (stony corals), but
are either colonial or solitary, while the sea fans, sea can also include a few species of octocorals and hydro-
pens, softcorals (otcocorals) and black corals zoan corals such as Heliopora and Millepora respective-
(antipatharians) are colonial. ly. The ahermatypic, or non-reef building corals, gener-
ally do not contain zooxanthellae; they include many
The most abundant Cnidarians on coral reefs, species of solitary hydrozoans and scleractinians and do
the stony corals (Scleractinian corals), are enormously not contribute a significant amount of calcium carbonate
important in the ecology of tropical seas. In warm, to the reef structure..
clear water they build massive reefs, which are highly
productive “oases” in otherwise relatively barren water Cnidarians capable of stinging humans include
masses. The process of calcification, in which stony free-swimming jellyfishes of the Hydrozoa and
coral polyps take carbon dioxide and water and with the Scyphozoa and benthic fire corals, hydroids, sea
aid of intracellular symbiotic algae (zooxanthellae) pro- anemones, corallimorpharians, zoanthids and the polyp
duce calcium carbonate, appears simple, but it is actual- stage of some jellyfish. The injury they can cause
ly quite complex and still incompletely understood. The humans ranges from no discernible effect to the poten-
end result, however, is the enhanced growth of individ- tially fatal stings produced by the powerful nematocyst
65
Above left- These Scyphozoan jellyfish, Aurelia, swim just beneath the surface of the wateron calm days. From a a divers position
below them looking up, they resemble flying saucers in outer space. These jellyfish feed on small planktonic animals which have
also gathered together near the surface. Below left- Many other animals form symbiotic associations with Cnidarians. These juve-
nile fish (jacks) seek protection within the bell of the jellyfish. They are not imune to the sting of the medusa and will quickly become
prey if they become too tangled in the tentacles. Above right- These mushroom corals (Fungia fungites) and related genera, are soli-
tary unattached coral polyps as adults. Mushroom corals are able to roll over if turned upside down and they can move about the
reef. They are often found in aggregations on patch reefs. The color of these corals is variable and due to the presence of symbiot-
ic algae. Below right- This photo shows a section of reef in the Philippines, covered with soft and hard corals and otherinvertebrates.
toxins of some sea wasps and the Portuguese man-of- unpleasant burning sensation. The best treatment for
war. Hydroids, such as Millepora, Aglaeophenia these stings is 95% alcohol.
cupressina and Lytocarpus philippinus and certain coral-
limorpharians can be very abundant in areas frequented Some of the zoanthids are also potentially dan-
by divers and swimmers. The sting and subsequent rash gerous to humans. Members of Palythoa and Zoanthus
that results from contact with these Cnidarians is painful have toxins, generally known as palytoxins, which are
and may last for several days. present in mucous and the gonads. Palytoxins are
among the most toxic substances found in nature; fortu-
The nematocysts of octocorals usually will not nately they are not highly concentrated in the surface tis-
sting humans, however many octocorals possess sues of the zoanthid. While casual contact with zoan-
spicules, small calcareous structures which can scratch thids generally will not cause harm to humans, contact
and penetrate human skin. Additionally, sea fans are with Palythoa or Zoanthus through an open wound can
home for certain species of small brittlestars, with abun- be very painful and dangerous. The toxicity of Palythoa
dant needle-like spines. Touching a sea fan with brit- was well known to the ancient Hawaiians who coated
tlestars, even with a gloved hand, may result in a very spear tips with it to make them more deadly.
Class Hydrozoa- Hydrozoans
Among commonly observed hydrozoans on coral reefs are the hydroids, which are colonial, polyp-like
animals. Large, bushy or plume-shaped, hydroids at first glance look more like algae than animals. Hydroids,
less than an inch in height, are quite common although usually overlooked due to their small size and often cryp-
tic habits. Some hydroids, such as the Stylaster and Millepora spp. (fire coral), have calcareous skeletons and
resemble scleractinian corals. Hydroids and hydromedusae are carnivorous and feed on small planktonic ani-
mals.
Hydrozoans occur as either polyps or medusae or both. Hydromedusae are small jellyfish, generally
less than an inch or so in diameter, and are usually transparent. They can often be observed near the surface on
calm days in clear water. Some of these small jellyfish, and larger jellyfish-like hydrozoans such as the
Portuguese man-of-war, are capable of producing painful stings when they come in contact with a swimmer.
Use caution whenever dealing with hydroids. There are other animals which put these powerful nematocysts to
their own uses. Hydroids are the prey of various nudibranchs who store unfired, ingested nematocysts in spe-
cialized pockets of their digestive tract . The nudibranch is then able to sting and use the nematocysts for its
own defense.
Most hydroids have male and female individuals, but their life cycle is highly variable, sometimes com-
plex and poorly understood. Generally, attached colonial hydroids develop male or female medusae which bud
off, then swim away, develop gonads and reproduce. The fertilized egg divides and develops into a free swim-
ming larvae that subsequently attaches to the bottom and grows into a new hydroid. Hydroids may live for as
little as a few weeks or, as in the case of Millepora, many years. This alternation between medusae and attached
polyp has led to much confusion in naming hydroids; often there are separate generic names for polyp and
medusae of the same species.
66
Above left- Photomicrograph of nematocysts from the Portugese man of war. One large exploded nematocyst is visible in the cen-
ter, two large unfired nematocysts are just behind it, many small round nematocysts are also visible. Below left- This photo shows
small polyps which always live with the sponge. Although similarto hydroids, these are actually the polyps of a coronate medusa,
possibly Nausithoe. Right- Alarge head of the fire coral Millepora sp. with a crinoid on top is in the center of the photo.
239- Solanderia sp. * Solanderiidae * Hydroida Papua New
Guinea * Madang * barrier reef * 20 ft (6 m). The family and
genus is found worldwide in the tropics. While a hydroid, this genus
resembles a small sea fan or gorgonian with its branches usually in
one plane, perpendicular to the current or wave action. There are
several species, including some which live as deep as 300 feet or
more. S. misakinensis is known from Japan and Hawaii. S. minima
is known from Zanzibar and possibly Hawaii. S. secunda is known
from the central Pacific. The most common members are always
found in exposed areas on wave swept shallow outer reefs.
69
249- Unidentified hydroid * Indonesia 250- Antennellopsis integerrima * Papua New Guinea
251- Myronoema sp. * Papua New Guinea 252- Rhizogeton sp.?* Federated States of Micronesia
71
261- Stylaster sanguineus * Palau 262- Stylaster sp. * Indonesia
263- Stylaster sp. * Marshall Islands 264- Distichopora borealis * Federated States of Micronesia
265- Distichopora irregularis * Indonesia 266- Distichopora violacea * Federated States of Micronesia
267- Distichopora sp. * Stylasteridae * Hydroida * Marshall
Islands * Enewetak Atoll * leeward barrier reef * 33 ft (10 m).
Some species of Distichopora, such as this one, are often abundant
on the open reef face on the outer dropoffs. There is a species of vio-
let ascidian growing at the base of the coral colony.
73
274- Olindias sp. * Indonesia 275- Aequorea australis * Federated States of 276- Timoidesagassizi * Marshall Islands
Micronesia
The Scyphozoans are the animals we normally think of when someone mentions the word jellyfish.
These relatively large and often brightly-colored medusae are some of the most beautiful animals in the sea. They
often occur seasonally in large aggregations and are easily observed as they swim slowly near the surface.
Scyphozoans, unlike their Hydrozoan relatives, not only tend to be larger, but also have different types of nema-
tocysts. Most species are several inches to a foot in diameter, however, some attain bell diameters of several
feet, which makes them the largest Cnidarians, except for some colonial scleractinan corals. Scyphozoans are
entirely marine with their general habitats near shore. They sometimes are blown or carried into brackish water
estuaries.
The life cycle of most Scyphozoans resembles the alternation between polyp and
medusa stage found in the Hydrozoa, except that the polyp stage is much reduced.
Based on the early development of the newly-settled larvae and a few morpholog-
ical characters, the Scyphozoa comprise four groups; the Stauromedusae, Coronate
medusae, Seameostomae, and Rhizostomae. The Cubomedusae (sea wasps), for-
merly included in the Scyphozoa, are now thought, by some taxonomists, to repre-
sent a separate class of Cnidaria, the Cubozoa, based on their early life history,
however; for convenience, we have included them with the Scyphozoa .
Stauromedusae are jellyfish that do not swim. Usually less than an inch long, they have a stalk arising from
the top of their bell which they use to adhere to blades of sea grass. Coronate medusae live primarily in very deep
water but are represented in shallow tropical waters by several species. The Coronate medusae have larger polyps
than most jellyfish and some of these polyps have been classified in genera different from the medusae due to pre-
vious lack of knowledge concerning their life history. The Seameostomae and Rhizostomae include the medusae
we usually think of as jellyfish; Aurelia (moon jelly) and Cassiopea (upside-down jelly) respectively. They are
common near shore, often colorful and at times can be observed with commensal fishes, nudibranchs or shrimp.
The Cubomedusae are nearly transparent and have four tentacles or bundles of tentacles that originate at
the four corners of the squarish bell. They are represented by at least five species in the tropical Pacific, all of which
can inflict a painful sting.
Little is known about the longevity of jellyfish; however, it appears most mature within a year’s time,
reproduce and then die. The following year new medusae bud off the surviving polyp stage and enter the water col-
umn to repeat the life cycle.
74 Jellyfish appear to have few predators. Several species of butterfly fish have been observed eating pieces
of large jellyfish at Chuuk in Micronesia and there are reports of large turtles eating jellyfish. Certain large species
are harvested commercially for food, however the bell must be prepared properly in order to remove all nemato-
cysts before being consumed.
277- Carybdea rastoni * Federated States of 278- Carybdea marsupialis * Palau 279- Chiropsalmus sp. * Philippines
Micronesia
277- Carybdea rastoni * Carybdeidae * Cubomedusae * Cubozoa
* Federated States of Micronesia * Chuuk Atoll * open water.
This is the juvenile medusae of Carybdea alata several days after it
has metamorphosed from polyp to medusa. There are two species of
cubomedusae (sea wasps) known from Hawaii, C. alata and C. ras -
toni. Both occur in the tropical western Pacific as well. In other
areas of the tropical Pacific the species occurring are not well known.
76
285- Pelagia noctiluca * Philippines 286- Phyllorhiza punctata * Hawaii 287- Mastigias papua * Papua New Guinea
284- Sanderia malayensis * Pelagiidae * Semaeostomeae * 287- Mastigias papua * Mastigiidae * Rhizostomae * Scyphozoa
Scyphozoa * Bahrain * open water. Although sometimes found at * Papua New Guinea * Madang * mangrove * open water. This
sea, this species more commonly occurs near shore in bays and estu- species appears to be quite variable in color and morphology. It
aries. The body attains lengths of nearly ten feet and the tentacles occurs over the entire western Pacific and is easily confused with P.
can extend twenty feet or more. Sanderia is a strong swimmer, but punctata above. It is possible there are several other species, close-
most often is carried with currents. Like most jellyfish, part of the life ly related to Phyllorhiza and Mastigias, which occur in this region.
of Sanderia is spent as an attached (benthic) form. Seasonally, the
polyps metamorphose to produce free-swimming jellyfish. 288- Mastigias sp. * Mastigiidae * Rhizostomae * Scyphozoa *
Palau * Jellyfish Lake * midwater.
285- Pelagia noctiluca * Pelagiidae * Semaeostomeae * This species inhabits marine lakes in Palau. The most interesting dif-
Scyphozoa * Philippines * Cebu * open water. The eight reddish, ference between this species and M. papua above is its near inability
stinging tentacles, shown here contracted, can extend several feet and to inflict a sting on bare skin. Both species have nematocysts and it
are used to stun and capture prey. The frilly lower portions of the appears those in the lake have lost almost all of their potency and
animal are the food-gathering oral arms which draw the food into the ability to sting. This species is filled with zooxanthellae and swims
mouth located on the underside of the bell. Most often encountered to those parts of the lake with the best sun exposure.
in warm offshore waters, these small (4 inches in length) jellyfish are
sometimes carried inshore by currents. 289- Cassiopea medusae * Cassiopeidae * Rhizostomae *
Scyphozoa * Philippines * Cebu * seagrass bed * 6 ft (2 m).
286- Phyllorhiza punctata * Mastigiidae * Rhizostomae * Cassiopea is usually found in shallow bays and estuaries where the
Scyphozoa* Hawaii * coastal open water. This jellyfish occurs water is still. They are most often observed with the top of the bell
around the world in warm water. The polyps of this species seem able laid against the substratum and the mouth and arms directed
to survive in the ballast tanks of large ships or as fouling organisms. upwards. The margin of the bell pulses occasionally as if to help the
It has been reported from Pearl Harbor, San Diego Bay, San Juan, jellyfish swim, however this effort is directed towards moving water
Puerto Rico, and Rio de Janeiro, all major ship harbors. This species and food across the oral region. The jellyfish is perfectly happy to be
can grow to almost three feet in diameter. upside down.
288- Mastigias sp. * Palau 289-Cassiopea medusae * Philippines
77
290- Cassiopea andromeda * Palau 291- Cephea cephea * Federated States of Micronesia
292- Crambione mastigophora * Federated States of Micronesia 293- Thystanostoma flagellatum * Palau
290- Cassiopea andromeda * Cassiopeidae * Rhizostomae * 292- Crambione mastigophora * Catostylidae * Rhizostomae *
Scyphozoa * Palau * Lighthouse Reef * sea grass bed * 6 ft (2m). Scyphozoa * Federated States of Micronesia * Chuuk * lagoon *
This jellyfish is similar in habit to C. medusae above, the principal open water. This jellyfish is common in the lagoon waters of Chuuk
difference being that this species prefers to inhabit seagrass areas. during late summer. The bell can be up to about one foot across. It is
not a strong swimmer and large aggregations of up to several hun-
291- Cephea cephea * Cepheidae * Rhizostomae * Scyphozoa * dred individuals may be observed gently drifting with the tidal cur-
Federated States of Micronesia * Chuuk * lagoon * open water. rents. Crambione can deliver a very painful sting if contact is made
This jellyfish varies a great deal in external appearance primarily as with exposed skin.
a function of size. There are several species described from the west-
ern Pacific, however we have only encountered this one in Chuuk 293- Thystanostoma flagellatum * Thysanostomatidae *
and Palau. Large individuals are about one foot in diameter. Small Rhizostomae * Scyphozoa * Palau * open water. This is a small
individuals have numerous frilly tentacle-like extensions off their jellyfish for a rhizostome with a bell about four inches in diameter.
oral arms. The medusae is generally accompanied by a small fish (jack) which
swims completely inside the bell. It seems to prefer open ocean, as
we have only rarely seen it near shore.
Class Anthozoa - Anthozoans
Anthozoa is derived from a Greek word which means flower animal. Anthozoans are Cnidarians in which
body form is based on the polyp. The medusa stage is absent. The Anthozoans are the most diverse class within
the Cnidaria and that diversity is reflected in the wide variation in polyp morphology. They may be convenient -
ly divided into octocorals and hexacorals, the former (as their name implies) have 8 branched tentacles while the
latter usually have 6 unbranched tentacles or multiples thereof. Within the Anthozoa the octocorals (sea pens,
soft corals, sea fans) are a fairly well-defined group, although taxonomy at the species level can be difficult.
Within the hexacorals (hard corals, corallimorpharians, anemones, zoanthids, black corals and tube anemones)
there is enough variation in nematocysts, skeletal structure and early life history to easily distinguish orders. In
spite of the prominence of Anthozoans in shallow water tropical habitats, very little is positively known about
the distribution (zoogeography) of species across the tropical Pacific due to inadequately and often times improp-
erly identified specimens.
In this book the Anthozoa are divided up into three picture and note sec-
tions; the octocorals, the stony hexacorals and the other hexacorals. A brief
introduction of each group precedes the picture section for that group.
78
Above left- This close up photo of an octocoral shows typical polyps with eight pinately branched tentacles, also visble just beneath
the red pigment are the spicules which make up the axial skeleton. Below left- This photo shows an ahermatypic coral,
Dendrophyllia with its unbranched tentacles expanded at night. In general , only the octocorals have branched tentacles. The small
orange spots on the transparent tentacles are clumps of nematocysts. Above right- This sea anemone Heteractis magnifica has com-
mensal fish, crabs and shrimp. Many Anthozoans serve as partners in symbiotic relationships with other reef organisms. Below
right- The coral, Porites has little defense against parrotfish which feed on the algae contained within the coral polyp. This photo
is a good example of one way new substrate is created on the reef.
Subclass Octocorallia - Octocorals
Octocorals have calcareous spicules within their body tissue which aid the support and maintenance
of form in large colonies. The shape and size of these spicules, which differ from sponge spicules in shape,
often determine classification of a species. The spicules are often times just under the outer surface of the
octocoral and may cut or scratch a diver’s hand if the animal is disturbed.
Most octocorals are filter feeders and inhabit areas where currents flow. Many shallow water species, 79
especially the brown-colored ones, contain symbiotic zooxanthellae. Photosynthetic products from the zoox-
anthellae certainly augment the nutrient intake of the octocoral and enhance calcification in the stony corals.
Dendronephthya are some of the most spectacular organisms found on Pacific reefs, with brilliant
colors, bizarre shapes and large sizes. They lack zooxanthellae so their sclerites are normally visible through
the translucent body wall. Normally photographed when they are inflated with water, they often deflate to a
small spiny lump on the bottom which is hard to identify as the same creature as the expanded individual.
Above- Soft corals of Dendronephthya are admired for their delicate beauty. Because they have no zooxanthellae, the calcareous
spicules are visible through the body wall. Their taxonomy is poorly known so they cannot be identified as a particular species.
294 - Heliopora coerulea * Helioporidae * Helioporacea *
Octocorallia * Philippines * Pamalican Island * 13 ft (4 m). This
abundant non-scleractinian coral, known as “blue coral”, has the
internal skeleton blue in color from iron salts. In the living colony
this is visible only if a branch or plate has been broken off. It varies
from delicate branches and vertical blades in shallow depths to hori-
zontal plates in deeper water. It superficially resembles Millepora
and some scleractinian corals, but once its characteristic appearance
is learned, it is hard to confuse with anything else. In some areas,
such as Ishigaki Island near Okinawa, it can be the dominant reef
coral. Blue coral only occurs as far east as the Marshall and Gilbert
Islands, and Samoa. It is not known from Fiji, French Polynesia or
Hawaii.
81
299 - Carijoa sp. * Federated States of Micronesia 300 - Carijoa sp. * Marshall Islands
303 - Tubipora musica * Papua New Guinea 302 - Tubipora musica * Federated States of Micronesia
307 - Minabea aldersladei * Alcyoniidae * Alcyoniina *
Octocorallia * Papua New Guinea * Madang * reef * 66 ft (20 m).
307 - Minabea aldersladei * Papua New Guinea 324 - Dendronephthya sp. * Nephtheidae * Octocorallia *
308 - Sinularia dura * Papua New Guinea 309 - Sinularia sp. * Papua New Guinea
83
310 -Sinularia sp. * Papua New Guinea 311 - Sarcophyton crassocaule * Papua New Guinea
312 - Sarcophyton sp. * Federated States of Micronesia 313 - Sarcophyton * Papua New Guinea
319 - Sarcophyton sp. * Palau 331- Nidalia simpsoni * Nidaliidae * Alcyoniina * Octocorallia *
320 - Sarcophyton sp. * Federated States of Micronesia 321 - Dendronephthya sp. * Palau
322 - Dendronephytha sp. * Federated States of Micronesia 323 - Dendronephthya sp. * Federated States of Micronesia
85
324 - Dendronephthya sp. * Federated States of 325 - Dendronephthya sp. * Federated States of 326 - Dendronephthya sp. * Papua New Guinea
Micronesia Micronesia
327 - Dendronephthya sp. * Papua New Guinea 328 - Dendronephthya sp. * Papua New Guinea 329 - Studeroites sp. * Palau.
Papua New Guinea * Madang * harbor entrance * 100 ft (30 m).
This photo was taken at night when Nidalia simpsoni is most active.
At dusk the round head (capitulum) of the colony inflates with water
and the polyps expand, exposing their tentacles. These are small
octocorals, about four to five inches in length. This species is usual-
ly found on vertical faces near shaded ledges. It is often overlooked
during the day.
87
336 - Cespitularia sp * Papua New Guinea 337 - Cespitularia sp. * Papua New Guinea
338 - Xenia sp. * Federated States of Micronesia 339 - Xenia sp. * Papua New Guinea
340 - Briareum sp * Papua New Guinea 341 - Briareum sp. * Papua New Guinea
342 - Ctenocella sp. * Papua New Guinea 343 - Ctenocella sp. * Federated States of 344 - Junceella sp. * Papua New Guinea
Micronesia
colony, which then falls to the bottom and starts a new individual.
89
350- Semperina sp. * Papua New Guinea 351 - Solencaulon sp. * Indonesia
352 - Solenocaulon sp. * Papua New Guinea 353 -Subergorgia sp. * Papua New Guinea
354 - Subergorgia suberosa * Federated States of Micronesia
91
361 - Melithaea sp. * Palau 362 - Melithaea sp. * Papua New Guinea
363 - Melithaea sp. * Papua New Guinea 364 - Acalycigorgia sp. * Papua New Guinea
365 - Acalycigorgia sp. * Federated States of Micronesia 366 - Acanthogorgia sp. * Federated States of Micronesia
Octocorallia * Federated States of Micronesia * Chuuk * Anaw
Reef wall * 165 ft (50 m).
370 - Anthogorgia sp. * Papua New Guinea 385 - Rumphella sp. * Gorgoniidae * Holaxonia * Octocorallia *
371 - Acanthogorgia sp. * Marshall Islands 372 - Anthogorgia sp. * Papua New Guinea
93
373 - Muricella sp. * Indonesia 374 - Astrogorgia sp. * Federated States of Micronesia
375 - Astrogorgia sp. * Papua New Guinea 376 - Astrogorgia sp. * Federated States of Micronesia
377 - Bebryce sp. * Federated States of Micronesia 378 - Bebryce sp. * Papua New Guinea
379 - Lophogorgia sp. * Federated States of Micronesia 380 - Menella praelonga * Federated States of Micronesia Pseudothesea sp.
Octocorallia * Papua New Guinea * Eastern Fields * 100 ft (30
m).
382 - Menella sp. * Palau 383 - Villogorgia sp? * Indonesia 384 - Isis hippuris * Papua New Guinea
385 - Rumphella sp. * Philippines 386 - Plumigorgia hydroides * Federated States of Micronesia
95
387 - Stephanogorgia sp. * Papua New Guinea 388 - Asterospicularia sp? * Papua New Guinea
389 - Rumphella sp * Federated States of 390 - Veretillum sp. * Indonesia 391 - Cavernulina sp. * Papua New Guinea
Micronesia
392 - Cavernularia cf. chuni * Philippines 393 - Pteroeides sp. * Indonesia 394 - Pteroeides sp. * Philippines
396 - Pteroeides sp. * Papua New Guinea 397 - Virgularia sp. * Philippines 398 - Virgularia sp. * Palau
Stony Corals-Scleractinia
As indicated previously, the stony corals fall into two general groups, the
reef-building or hermatypic and the non-reef building or ahermatypic corals.
Pacific Ocean stony corals are the most diverse coral fauna in the world, with
some sites having as many as 300 to 400 species. The highest generic richness,
about 90 genera, is believed to occur in the area formed by a triangle including
the Philippines, northern Indonesia and New Guinea, and extends to the northern
Great Barrier Reef. The taxonomy of Pacific Ocean reef corals is imperfectly
known, although much progress has been made in the last few decades to sort out the variable and similar
species. Identifications from dead skeletons may be difficult with many variations in growth form and skele-
tal characters depending on environmental conditions. Living colonies also pose problems since the colors of
many corals can vary greatly. Additionally the living tissue of polyps masks the skeletal characteristics which
are often necessary for accurate identification.
The stony corals fall into about 15 families. Assigning a particular coral to a specific family is the first
step in identifying it. Some families contain genera which are easy to recognize. Others are much less dis-
tinctive and cannot practically be identified from photographs unless the colony photographed is also available
as a specimen and primary literature is used.
The coral identifications presented in this book have been based on actual specimens in some cases, but
more often have been made using available literature from the photographs alone. This is why many of the
identifications are indefinite, often only to genus. It is hoped the species of stony corals included will assist in
generally placing a particular coral amongst the many families and genera.
Tissues of most hermatypic stony corals contain symbiotic algae, called zooxanthellae, which give the
coral polyps most of their color. In the presence of light these algal cells use nitrogen-containing waste prod- 1
ucts and carbon dioxide from the polyp to produce sugars and amino acids through the process of photosyn-
thesis. Enzymes in the coral tissue cause some of these nutrients to leak out of the zooxanthellae; the nutrients
are in turn used by the coral for its own growth. Obviously hermatypic corals grow best in shallow, clear, sun-
lit waters.
During the day most species of stony corals have the polyps retracted into cup-like calices, (fluted
depressions that make up the upper part of the skeleton). However, at night the polyps expand, and the coral
looks entirely different. To augment the nutrients produced by the zooxanthellae corals use their tentacles to
capture food at night. Corals defend their space on the reef at night as well; some species have special sweep-
er tentacles which can reach distances of several inches to attack neighboring organisms to keep them from
overgrowing the stony coral.
Some stony corals grow unattached to the bottom. Several genera of the family Fungiidae, the mush-
room corals, are free living to the extent they can even move themselves along the bottom. Although these
fungiid corals start out life as an attached polyp, early on the upper disc of the polyp breaks free. The disc set-
tles on the bottom and grows. If the coral finds itself in an unsuitable spot, it is able to inflate itself with water
and roll over along the bottom. Other families have fewer numbers of free-living stony corals; Goniopora
stokesi, grows in small hemispherical colonies on sandy bottoms near reefs.
Corals reproduce both asexually and sexually. Colonies grow by two types of asexual division of the
polyps, intratentacular (within the oral disc) and extratentacular (outside the oral disc) budding. Another form
of asexual reproduction can occur by a method known as “polyp bailout” in which stressed coral polyps leave
the skeleton and float away short distances and redevelop into new colonies. Some corals also form “satellite”
colonies, such as Goniopora stokesii, in which small buds form off the original colony and eventually break off
to form separate colonies.
Sexual reproduction also has several variations. Some corals have gonads of both sexes in each polyp
(hermaphroditic). Other corals have colonies with separate sexes. Fertilization is either internal, in which case
sperm released into the water swim to female polyps with eggs, or external, in which both eggs and sperm are
released into the water. Self-fertilization, most likely, does not occur. Whether eggs are fertilized externally
in the water or internally, they eventually produce larvae (planulae). The planulae are ciliated and able to
swim, their oblong bodies are at most about 1/4 of an inch long. Planulae may remain in the water column
for up to several months time and are the main means of medium to long distance dispersal for most corals.
The species of Pocillopora have planulae which can be extremely long lived in the plankton and this is
believed to be a major reason why this genus is found from the Red Sea to the western coast of North
America.
In the past decade scientists have discovered that in some areas many species of stony corals spawn
nearly simultaneously, resulting in a spectacular release of gametes over large areas of reef almost on cue. On
the Great Barrier Reef the mass coral spawning occurs just after the full moon of November. In other areas
timing is different; for example, off western Australia coral spawning occurs in April. In Micronesia coral
spawning is less well known, but a number of species are believed to spawn during July.
When the planula larva settles to the bottom, it produces a single polyp which starts building the cal-
cium carbonate skeleton. From that single polyp and its skeleton, the coral colony grows by budding of indi-
vidual polyps, initially forming an area of firm attachment to the substrate. Once a sizeable base is estab-
lished, the colony can begin to grow upward. Acropora species (one of the most common corals) have two
types of polyps, axial (at the tips of the branches) and radial (along the sides of the branches). Growth is rapid
at the tips of the branches, where the axial corallites occur, and members of this genus are among the fastest
growing of all corals.
The genus Acropora, and other genera of corals occurs in many different growth forms and these
forms are often times the best character for identifing species in the field. Color is not a good character to
use, as it is often quite variable within a single species. Growth forms include: plate or table-like, tree-like
98 (arboresecent ), encrusting, corymbose (short branches arising from horizontal mass), pillow-like, digitate
(short non-dividing, unconnected), bushy, and massive.
In just about any area of the tropical Pacific numerous species of Acropora are to be found. For
example, in Australia, 73 species are reported from the eastern coast and 54 from western Australia. In
Micronesia, 36 species are reported from Guam, and quite a few more occur in other Micronesian waters.
Above - This diagram of coral polyps is cut away to show the relationship of soft tissue to the coral skeletonn In the cen-
ter of the cut away portion is a large gastrovascular cavity divided by mesentaries. The septa of the coral skeleton form
against these folds.
399- Stylocoeniella guentheri * Federated States of Micronesia 400- Pocillopora damicornis * Palau
403- Pocillopora eydouxi * Pocilloporidae * Scleractinia * 402- Pocillopora danae * Federated States of Micronesia
Federated States of Micronesia* Fujikawa Maru * 50 ft (15 m).
This species of Pocillopora is quite distinctive with upright flattened
branches with pale ends. A number of crabs and fishes are associat-
ed with P. eydouxi, the crabs nestled deep in the crooks of the branch-
es. This species occurs from the Red Sea and east Africa to Hawaii.
404- Pocillopora verrucosa * Federated States of Micronesia 408- Seriatopora hystrix * Pocilloporidae * Scleractinia * Papua
New Guinea * Madang * barrier reef * 10 ft (3 m). This species
is often called “needle coral” because of the fine sharp points on the
branches. It is found from east Africa across the Indian Ocean and the
Pacific to as far east as Micronesia and Samoa.
101
410- Stylophora mordax * Federated States of Micronesia 411- Acropora granulosa * Federated States of Micronesia
412- Acropora palifera * Federated States of Micronesia 413- Acropora pruinosa * Hong Kong
414- Acropora tenella * Federated States of Micronesia 415- Acropora robusta * Philippines
414- Acropora tenella * Acroporidae * Scleractinia * Federated
States of Micronesia * Chuuk Atoll * Ozen Island * 133 ft (40 m).
This remarkable species of Acropora is branched like many other
species, but is greatly flattened with calices only on its upper surface
to maximize exposure to light. Not surprisingly, it is found in deep-
er reef waters, usually below 80 feet, although small specimens are
occasionally encountered shallower. It is quite a distinctive species
that is easy to recognize.
103
422- Acropora valenciennes* Marshall Islands 423- Acropora sp. * Indonesia
424- Acropora sp. * Federated States of Micronesia 425- Acropora sp. * Federated States of Micronesia
426- Acropora sp. * Federated States of Micronesia 427- Anacropora sp. * Palau
428- Astreopora gracilis * Acroporidae * Scleractinia * Federated
States of Micronesia * Chuuk Atoll * lagoon * 30 ft (9 m).
105
434- Alveopora sp. * Federated States of Micronesia 435- Alveopora sp. * Federated States of Micronesia
436- Goniopora stokesii * Papua New Guinea 437- Goniopora sp. * Papua New Guinea
438- Goniopora cf. tenuidens * Federated States of Micronesia 439- Goniopora sp. * Federated States of Micronesia
bled to new areas, where they survived and started growing as
separate reef patches. Often, in Micronesia, P. cylindrica is yellow.
It can be confused with Palauastrea ramosa which is superficially
similar, but has distinct polyps and calices. P. ramosa is limited in its
distribution, being found only in Palau among Micronesian islands.
107
446- Porites sp. * Federated States of Micronesia 447- Porites sp. * Federated States of Micronesia
448- Porites rus * Federated States of Micronesia 449- Porites australiensis * Federated States of Micronesia
450- Porites sp. * Philippines 451- Psammocora contigua * Federated States of Micronesia
454- Leptoseris papyracea * Agariciidae * Scleractinia *
Federated States of Micronesia * Chuuk * lagoon bottom * 112 ft
(34 m). This small coral can occur in dense masses in moderate
lagoon depths. The photograph was taken in an area of the northwest
Chuuk lagoon where this coral dominated the bottom, the dead skele-
ton building small hills on the bottom. It is found from the western
Indian Ocean to Hawaii, and is much more delicate than the similar
Leptoseris gardineri.
452- Psammocora digitata * Federated States of Micronesia 457- Pavona cactus * Agariciidae * Scleractinia * Papua New
Guinea * Laing Island * 40 ft (12 m). At the opposite extreme
from Pavona minuta is P. cactus, a delicate form which is most com-
mon in turbid water. Stands of P. cactus can be quite extensive, mea-
suring many meters across. The species is bifacial, with polyps on
both sides of the fronds. It occurs throughout the region as far east as
the Marshall Islands.
109
458- Pavona clavus * Papua New Guinea 459- Pavona decussata * Hong Kong
460- Pavona decussata * Federated States of Micronesia 461- Pavona minuta * Federated States of Micronesia
462- Fungia fragilis * Federated States of Micronesia 463- Fungia fungites * Federated States of Micronesia
the tissue of the polyp with water in order to push the coral right side
up. In this manner they are also able to move from one area to anoth-
er. The ridges on the surface of the coral are very sharp and handel-
ing these corals is not advised.
467- Halomitra pileus * Federated States of Micronesia 468- Heliofungia actiniformis * Papua New Guinea
469- Herpolitha limax * Marshall Islands 470- Podobacia? sp.* Hong Kong
111
471- Polyphyllia talpina * Federated States of Micronesia 472- Podabacia crustacea * Philippines
473- Sandalitha robusta * Palau 474- Acrhelia horrescens * Papua New Guinea
475- Galaxea sp. * Papua New Guinea 476- Galaxea paucisepta * Palau
common in a wide range of habitats. It occurs in a wide variety of
colors.
478- Mycedium elephantotus * Pectiniidae * Scleractinia *
Philippines * Pescador Island * 20 ft (6 m).
480- Oxypora glabra * Palau 481- Pectinia lactuca * Papua New Guinea
482- Pectinia paonia * Marshall Islands 483- Acanthastrea echinata * Papua New Guinea
113
484- Cynarina lacrymalis * Papua New Guinea 485- Lobophyllia cf. corymbosa * Papua New Guinea
486- Lobophyllia corymbosa * Federated States of Micronesia 487- Lobophyllia corymbosa * Federated States of Micronesia
488- Lobophyllia hemprichii * Philippines 489- Lobophyllia hemprichii * Papua New Guinea
490- Lobophyllia sp. * Federated States of Micronesia 491- Lobophyllia sp. * Federated States of Micronesia
115
498- Hydnophora exesa * Papua New Guinea 499- Paraclavarina triangulata * Palau
500- Merulina amplicata * Federated States of Micronesia 501- Scapophyllia cylindricus * Federated States of Micronesia
502- Barabattoia amicorum * Federated States of Micronesia 503- Caulastrea curvata * Papua New Guinea
504- Caulastrea furcata * Palau 505- Diploastrea heliopora * Papua New Guinea
508- Favia stelligera * Federated States of Micronesia 509- Favites flexuosa * Federated States of Micronesia
510- Favites cf. halichora * Papua New Guinea 511- Goniastrea actinata * Federated States of Micronesia
117
512- Goniastrea pectinata * Federated States of Micronesia 513- Platygyra ?lamellina * Federated States of Micronesia
514- Montastrea curta * Federated States of Micronesia 515- Moseleya latistellata * Papua New Guinea
516- Oulophyllia crispa * Federated States of Micronesia 517- Platygyra daedalea * Federated States of Micronesia
514- Montastrea curta * Faviidae * Scleractinia * Federated
States of Micronesia * Nama Island * 40 ft (12 m).
515- Moseleya latistellata * Faviidae * Scleractinia * Papua New
Guinea * Eastern Fields *40 ft (12 m).
521- Platygyra sp. * Federated States of Micronesia 522- Euphyllia ancora * Papua New Guinea
523- Euphyllia divisa possibly * Papua New Guinea 524- Euphyllia glabrescens * Indonesia
119
525- Euphyllia parancora * Marshall Islands 526- Euphyllia parancora * Papua New Guinea
527- Euphyllia paradivisa * Papua New Guinea 528- Physogyra lichtensteini * Palau
531- Plerogyra sinuosa * Marshall Islands 535- Tubastraea micrantha * Dendrophyllidae * Scleractinia *
Papua New Guinea * Madang * barrier reef * 60 ft (18 m).
534- Tubastraea diaphana * Hong Kong 535- Tubastraea micrantha * Papua New Guinea
536- Tubastraea micrantha * Federated States of Micronesia 537- Tubastraea sp * Philippines
121
538- Tubastraea sp. * Philippines 539- Tubastraea sp. * Federated States of Micronesia
540- Turbinaria bifrons * Papua New Guinea 541- Turbinaria peltata * Papua New Guinea
542- Turbinaria reniformis * Federated States of Micronesia 543- Turbinaria reniformis * Federated States of Micronesia
Corallimorpharians, Sea Anemones, Cerianthids, Zoanthids and Antipatharians
For all practical purposes corallimorpharians are corals without a skeleton. Their internal anatomy, nema-
tocysts and tentacles are identical to the scleractinia. Many species are brightly colored. One species of
Pseudocorynactis, which stays closed all day, spreads its disk open at night to reveal a beautiful red column, trans-
parent tentacles and bright orange capitate spheres at the tentacle ends. Often times species of the genus Rhodactis
cover large areas of bottom. We saw one entire patch reef north of New Britain in Papua New Guinea that was
covered densely with nothing but Rhodactis over an area of thousands of square yards. Interestingly, other patch
reefs nearby had very few. Corallimorpharians reproduce asexually through a process called transverse fission
where the animal literally pulls itself apart and reforms into two individuals. Sexual reproduction is probably sim-
ilar to that observed in corals.
Sea Anemones are always solitary polyps. As in other Cnidarian polyps, the mouth is situated on an oral
disc surrounded by tentacles. The body is columnar in shape with a flattened “foot” at the base for attachment.
The main difference between anemones and the Cnidarians covered previously is the former have a well devel-
oped structure called the siphonoglyph located along one or both sides of the pharynx (octocorals have a reduced
siphonglyph). The siphonoglyph is lined with cilia which beat in rhythmic fashion to bring water into the gas-
trovascular cavity or reverse direction to help expel waste material. By aiding circulation of oxygenated water,
the siphonoglyph, enables anemones to attain larger sizes than most cnidarian polyps. Actinians resemble coral-
limorpharians, however they are generally firmer, larger and more variable in morphology.
There is great diversity in the body plan of different sea anemone species in the tropical Pacific. Sea
122 anemones are common in shallow water where they attach to rocks and other hard substrate. Many species live
on sandy and mud bottoms and several species live in association with other animals. The genus Calliactis has
several species which are only found on the shells of hermit crabs. A number of species of large shallow water
anemones harbor anemone fishes which only live in association with sea anemones. Much has been written about
this association and it is generally accepted that the anemone fish develop a mucous coating that renders them
unrecognizable as prey to the anemone. The association is well developed and there is certainly behavior on the
part of the fish and probably on the part of the anemone which is not fully understood.
Cerianthids are solitary, tube-dwelling anemones. They differ morphologically from the sea anemones by
having tentacles around the mouth (oral tentacles) as well as around the margin of the oral disc. In cerianthids,
the bottom of the column is rounded and not modified into an attachment structure (there are a few anemones like
this as well). They live mostly buried in the sand in tubes which they secrete. The tubes are made up of fired
nematocysts and encrusted sand particles. The tubes of large cerianthids may be up to several feet in length. When
disturbed the cerianthid quickly retracts into the safety of its tube. There are several species of shallow water ceri-
anthids in the tropical Pacific. Cerianthids have planktonic larvae that may live for six months in the water col-
umn. It is quite possible that many species of cerianthids are widely distributed. Unfortunately most of the species
are poorly known.
Zoanthids are generally colonial although a few are solitary. Colonial zoanthids may superficially
resemble coral heads. Colonial polyps are connected by stolons or, in species forming large mats, the polyps are
embedded in a tissue-like body (coenenchyme). Certain species of Palythoa and Zoanthus are capable of cover-
ing many square yards of reef flat or rocky bottom. When large areas of reef become overgrown with zoanthids
it is usually a good indication that the water quality of the area has changed for the worse. Many species of zoan-
thids live in association with other Cnidarians. The genus Parazoanthus lives almost exclusively with other
invertebrates such as sponges, hydroids and black corals. As previously mentioned, most zoanthids contain paly-
toxin and care should be taken to avoid contact with open cuts.
Order Antipatharia - Black Corals
In the black corals only the horny material of the axial skeleton is black. Polyps are colonial and-
cover the surface of the black skeleton; they may be white, yellow, orange or brown. The skeleton often has
small hooks or thorns on it which enable the polyps to grip the surface. The polyps are non-retractile and
there are no “cups” into which they can retract as in stony corals or gorgonians. Members of the genus
Cirrhipathes have a single whip-like skeleton. Other species, such as those of Antipathes, can be delicately
branched on one or more planes or can be extremely bushy. Some black corals in deep water can reach sev-
eral feet in height and more than four inches in diameter at the base of the skeleton.
The name antipatharian (Greek for “against disease”) was given at a time when it was believed black
corals possessed medicinal properties. To date, however, no pharmaceutical products have been obtained from
black corals. Black coral, certain octocorals and even stony corals are used to make jewelry, but their true
commercial potential is probably best realized by leaving them on the reef for tourist divers to observe.
Right- This photo shows a large black coral colony
(antipatharian) on a reef outcrop near Madang in
Papua New Guinea. Black corals grow slowly and a
large individual such as this may be fifty years old or
more. These corals are attached firmly to the reef by
a modification of the central axis. Superficially,
black coral seafans like this one resemble octocoral
sea fans. Both types of seafans are filterfeeders; they
can be distinguished by the number and type of
polyp tentacles. Black coral polyps have only six
unbranched tentacles. Numerous other inverte-
brates, such as, shrimp, fish, oysters, worms, and
other cnidarians are often found in association with
black corals.
125
550- Palythoa sp. * Marshall Islands 551- Sphenopus * Papua New Guinea
552- Parazoanthus sp. * Papua New Guinea 553- Parazoanthus sp. * Papua New Guinea
554- Parazoanthus sp. * Papua New Guinea 555- Parazoanthus sp. * Indonesia
isms to overgrow other organisms. Several species of Parazoanthus
live on sponges and hydroids. Studies have indicated that the zoan-
thid makes the host unpalatable, while the host provides substrate for
the zoanthid. The color of the zoanthid usually contrasts with that of
the host which makes it easier for predator species to recognize and
avoid (aposymatic coloration). This species of Parazoanthus is com-
monly found growing on the surface of the orange sponge Stylissa
flabelliformis. This is a typical growth form for Parazoanthus, simi-
lar to that of P. axinella from the Caribbean and Mediterranean.
127
563- Zonathella larve * open water 564- Anthopleura nigrescens? * Hong Kong
567- Dofleinia sp. * Indonesia 568- Entacmea quadricolor * Federated States of Micronesia
habit of asexual division. The crevices on a coral head inhabited by
this anemone will practically be filled with the tentacles of this
anemone. E. quadricolor occurs in many areas of the Indo-West
Pacific. This anemone harbors commensal fish (anemone fish).
129
576- Aiptasia pulchella * Palau 577- Alicia pretiosa * Federated States of Micronesia
131
588- Actineria villosa * Papua New Guinea 589- Cryptodendrum adhesivum * Papua New Guinea
590- Heterodactyla hemprichii * Federated States of Micronesia 591- Heteractis aurora * Marshall Islands
592- Heteractis malu * Papua New Guinea 593- Stichodactylus gigantea * Papua New Guinea
594- Heteractis magnifica * Papua New Guinea 595- Heteractis magnifica * Papua New Guinea
133
602- Amplexidiscus fenestrafer * Philippines 603- Discosoma nummiformis * Indonesia
604- Discosoma nummiformis * Federated States of Micronesia 605- Discosoma sp. * Papua New Guinea
606- Discosoma sp. * Federated States of Micronesia 607- Discosoma sp. * Marshall Islands
608- Discosoma sp. * Federated States of Micronesia 609- “Rhodactis” * Papua New Guinea
135
615- Arachnanthus oligopodus * Marshall Islands 616- Arachnanthus sp. * Marshall Islands
617- Arachnanthus sp. * Papua New Guinea 618- Cerianthus sp. * Indonesia
619- Cerianthus sp. * Federated States of Micronesia 620- Cerianthus sp. * Philippines
614- Ricordea sp. * Corallimorphidae * Corallimorpharia *
Indonesia * Biak Island * 60 ft (18 m).
137
627- Antipathes cf. recticulata * Palau 628- Antipathes cf. reticulata * Federated States of Micronesia
631- Antipathes sp. * Federated States of Micronesia 632- Antipathes sp. * Philippines
632- Antipathes sp. * Antipathidae * Antipatharia * Philippines *
PescadorIsland * 40 ft (12 m). In some areas black corals can occur
in clear water openly exposed to light, such as is seen here from a reef
in the Philippines.
139
639- Cirrhipathes sp. * Federated States of Micronesia 640- Cirrhipathes sp. * Federated States of Micronesia
The Ctenophores are a relatively small taxonomic group with 100 or so species. They are marine ani-
mals that superficially resemble jellyfish. Most ctenophores are transparent and pelagic, and they are found
drifting in open water. Unlike most cnidarians, they have only a single body form during their life history
and they are never colonial. The name Ctenophore comes from the eight rows of ciliary combs (ctenes)
which traverse the body. The common name for the group, “comb jellies”, refers to the appearance of these
specialized structures. The fused cilia making up an individual comb diffract light and account for the irrides-
cence seen in color photographs of ctenophores. Ctenophores have tentacles armed with adhesive cells called
colloblasts which aid in capturing prey from the surrounding water. Other species of ctenophores, which lack
large tentacles, capture their prey by enveloping them, much as we would capture small fish underwater with
a plastic bag.
Some ctenophores (order Platyctenida) are creeping forms which live primarily in association with
other invertebrates such as soft corals and echinoderms. They are usually pigmented rather than clear and
have two long tentacles which they extend to capture prey in the water column.
646- Beroe forskali * Beroida * open Pacific. Beroe is a member urchin spines, in this case Diadema sp. There is also a similar
of the Class Nuda which lacks tentacles, even as larvae. This species species which lives on the Crown-of-thorns starfish, Acanthaster
is cosmopolitan in warm waters. planci. The members of this order are sessile, living attached to
another creature.
647- Pleurobrachia sp. * Cydippida * open Pacific. This species
has short tentacles, one of which is easily visible in the photo. 652- Coeloplana astricola * Platyctenea * Philippines* Batangas
Members of this order have globular forms and can retract the tenta- * 40 ft (12 m). The mottled blotches on the skin of this starfish are
cles quickly. actually a sessile ctenophore. These creatures are poorly known, but
interestingly these ctenophores are believed to multiply by a process
648- Pleurobrachia sp. * Cydippida * open Pacific. In this view called fragmentation that produces new individuals. Generally any
the tentacles are fully extended fishing for prey. starfish which has this ctenophore has several. It is unknown for cer-
tain whether these ctenophores cause any damage to the starfish. This
649- Bolinopsis sp. * Lobata * open Pacific species, as well as the one that follows, is most active at night.
650- Leucathea sp. * Lobata * open Pacific. This ctenophore can 653- unidentified benthic ctenophore. * Platyctenea * Marshall
become seasonally abundant in Micronesian lagoons such as Chuuk Islands * Kwajalein Atoll * reef * 20 ft (6 m). This ctenophore is
lagoon during January through April. During this time, Leucathea is attached to a soft coral. While it may appear that it is damaging the
quite abundant in the upper several feet of water, but few individuals soft coral, it is actually fishing using its tentacles which are clearly
occur below about forty feet. visible. When prey are captured by the tentacles, they are drawn in
and the prey enveloped by the animal.
651- Coeloplana bannworthi * Coeloplanidae * Platyctenea *
Philippines * Pamalican Island * 9 m. This species lives on sea
646- Beroe forskali * open Pacific. 647- Pleurobrachia sp. * open Pacific
141
648- Pleurobrachia sp. * open Pacific. 649 - Bolinopsis sp. * open Pacific
652- Coeloplana astricola * Philippines 653- Unidentified benthic ctenophore * Marshall Islands
142
Phyla Platyhelminthes,
Nemertea, Annelida, Sipuncula,
Echiurida and Hemichordata
Marine Worms
The “worms”, as grouped here, do not constitute a naturally related
group, but because of their general form and the relatively few species that
would be observed by non-specialists, they are considered together here for con-
venience.
Phylum Platyhelminthes
The flatworms are most commonly known for their parasitic members, which include the
flukes (Class Trematoda) and tapeworms (Class Cestoda). The free-living flatworms (Class
Turbellaria) found in shallow tropical waters however, are the most spectacular members of
the group. They are among the most brightly colored of marine invertebrates, rivaled only by,
and often confused with, the nudi-
branchs. They can swim with 143
undulations of their thin, flattened
bodies, but normally crawl along
the bottom.
Opposite- The tube worm, Protula sp., is common nearsheltered areas on outside reefs of Indonesia and other
areas of the western Pacific. Protula uses the feathery cilliated structure, called the brachial plume, for filter
feeding and as an aid in respiration. If disturbed they pull back into their tube in an instant.
Phylum Nemertea - Ribbon worms
feeders are equipped with jaws for catching small, with the previous three groups, and they are more close-
mobile prey. Active crawlers tend to be scavengers. ly related to the echiurids, below. They are unsegment-
Many tube dwelling polychaetes have evolved special- ed and live in sand, rocks and coral. Sipunculids are
ized feeding structures, such as branchial or tentacular one of the major borers of coral skeletons, causing
crowns, which remove suspended particles from the weakening of the skeleton and its eventual destruction.
water and also double as respiratory organs. Their boring is believed to be a combination of mechan-
ical and chemical action. They are eaten by some fish-
Terebellid polychaetes live in tubes and spread es and molluscs, especially Mitra spp. Some of the
their long tentacles over the surface around their tubes. sand burrowers are almost a foot long, with unlined 145
The tentacles resemble white, red or green spaghetti and non-permanent burrows.
are drawn back to the tube when touched. The main
body of the worm is lodged in fissures in the reef and is Diversity is moderate in the sipunculids, but a
rarely seen. The worm feeds on algal and bacterial films few species are usually present in most habitats. There
with the food particles being brought back to the mouth are 23 species known from Great Barrier Reef while
via a ciliated groove on each tentacle. seven species have been collected from Enewetak Atoll.
Most polychaetes have separate sexes, although Phylum Echiura- Echiurid worms
some are hermaphrodites and a few change sex.
Fertilization of eggs takes place in the water column for Echiurids are unsegmented worm-like animals,
those species which release gametes into the water. with a highly extendible proboscis. They live in bur-
Other species mate and lay encapsulated eggs in the rows in mud, sand and rock. On reefs, echiurids are
tube of the female (a few species retain fertilized eggs most often found in coral rock formed by overhangs of
in the body of the female). Planktonic larvae develop coral heads. The feeding proboscis, with a bifurcate tip,
from the fertilized eggs and eventually settle to the bot- is occasionally seen extended out along the bottom for
tom to become juveniles. Among the most spectacular a distance of several feet or more from the body of the
examples of polychaete spawning are the palolo worms. echiurid. If touched the proboscis is quickly drawn back
In Samoa, after sunset during the first lunar cycle of to the worm.
October or November, the reproductive portion of the
body, the epitoke, breaks free and swims to the surface Phylum Hemichordata - Acorn worms
The epitokes are light sensitive and large numbers of
them swarm on the surface, eventually breaking apart to Acorn worms live in sediment and the main
release the eggs and sperm. evidence of their presence is a mound of coiled “cast-
ings” on the surface of the sand. The castings are sand,
Phylum Sipuncula- peanut worms sheathed in a thin layer of mucous that has passed
through the worm’s gut. They live in sandy bottom
Sipundulids do not belong phylogenetically habitats. Some polychaetes make similar castings.
654- Acanthozoon sp. * Pseudocerotidae * Polycladida *
Platyhelminthes * Papua New Guinea * Madang * barrier reef *
30 ft (9 m). The most diverse and colorful group of flatworms in the
western Pacific are members of the family Pseudocerotidae. The
family includes several genera, four of which are included in this sec-
tion.
657- Pseudoceros dimidiatus * Philippines 658- Pseudoceros ferrugineus * Papua New Guinea
659- Pseudoceros tritriastus * Philippines 660- Pseudoceros dimidiatus * Marshall Islands
147
661- Pseudoceros sp. * Papua New Guinea 662- Pseudoceros sp. * Papua New Guinea
663- Pseudoceros sp. * Marshall Islands 664- Pseudoceros sp. * Marshall Islands
665- Pseudoceros sp * Marshall Islands 666- Pseudobiceros affinis * Papua New Guinea
668- Pseudobiceros damawan * Pseudocerotidae * Polycladida *
Platyhelminthes * Marshall Islands * Enewetak Atoll * channel
* 32 ft (10 m).
669- Pseudobiceros cf. fulgor * Federated States of Micronesia 670- Pseudobiceros paralaticlavus * Philippines
671- Pseudobiceros gloriosus * Federated States of Micronesia 672- Pseudobiceros gratus * Papua New Guinea
673- Pseudobiceros sp. * Papua New Guinea 674- Pseudoceros sp. * Hawaii
149
675- Pseudobiceros sp. * Palau 676- Pseudobiceros sp. * Philippines
151
688- Cerebratulus sp. * Palau 689-Paralepidonotus ampulliferus * Marshall Islands
692- Notopygos albiseta * Marshall Islands 693- Leocrates sp. * Marshall Islands
694- Unidentified Sabellarid * Sabellaridae * Polychaeta * Palau
* Malakal Harbor entrance * 100 ft (35 m). We are uncertain
exactly what this tube dwelling polychaete is, but it is unusual in
appearance. Twin “antennae” protrude from its tube and the worm
pulls back deep into the tube at the slightest disturbance.
697- Bispira sp.* Philippines 698- Sabellastarte sp.* Papua New Guinea
699- Sabellidae * Papua New Guinea 700- Sabellidae * Papua New Guinea
153
701- Sabellidae * Papua New Guinea 702- Sabellidae * Papua New Guinea
705- Spirobranchus giganteus* Papua New Guinea 706- Protula magnifica* Philippines
707- Protula sp. * Serpulidae * Polychaeta * Papua New Guinea
* West New Britain * reef wall * 50 ft (15 m).
155
714- Unidentified Sipunculid * Marshall Islands 715- Unidentified Sipunculid * Philippines
716- Bonellia fuliginosa * Marshall Islands 717- Achaetobonellia maculata * Marshall Islands
718- Ptychodera flava * Marshall islands 719- Hemichordata mound * Marshall Islands
156
Phylum Mollusca
Molluscs
The historical popularity of shell collecting and the durable
nature of the hard shells of specimens, with a wealth of characters avail-
able, are the reasons for molluscs having perhaps the best known taxonomy of
all marine invertebrate groups. After the arthropods, the phylum Mollusca contains the greatest
number of described living species (100,000), plus an additional 60,000 known fossil species.
About half of the species are marine, the rest being freshwater or terrestrial.
Although the molluscs appear to be a heterogeneous assemblage, they are derived from 157
the same fundamental body plan. The body is typically divided into a head, with well-developed
sensory organs, a large, soft visceral mass, from which the phylum gets its name, and a muscular
foot. As possession of a muscular foot is believed to be the primitive condition in molluscs, loco-
motion by crawling is common in the group. Octopods and squids can take in seawater, expel it
from their mantle cavity, and spurt forward by means of jet propulsion, in addition to their ability
to crawl over the substrate with their well-developed tentacles. These animals are collectively
called cephalopods (“head-footed”) because the tentacles are derived from the muscular foot and
emanate from the head. In sessile forms, such as oysters, the muscular foot is greatly reduced.
Most molluscs possess an external calcareous shell which they secrete. Evolutionary
change in some groups, like the nudibranchs, squids, and octopus, has resulted in reduction, inter-
nalization, or complete loss of the shell. These species have developed other means of protection
and evasion, for example, the octopus confuses predators by emitting a cloud of ink.
Opposite- The giant clam, Tridacna gigas, on a coral reef at Bagabag Island, offshore of
Madang, Papua New Guinea. The giant clams were a regularcomponent of most western Pacific
coral reefs, but have been exploited in many areas, causing a great decrease in theirabundance
throughout the region.
Class Polyplacophora- Chitons
The chitons are strictly marine, but occur in all seas. They are flat-
tened, with eight overlapping plates comprising the shell, surrounded by a
girdle. Most tropical Pacific species areintertidal or shallow subtidal, and have
a large foot by which they can clamp down to rock so tightly that they cannot
be moved. Most feed on algae, grazing the surface of rocks, while a few also
feed on assorted encrusting invertebrates. They are generally slow-moving,
inconspicuous animals. There are approximately 500 living species of chitons
world-wide.
Most gastropods have a large, fleshy foot which is used for locomotion
over a variety of substrates, propelled by either ciliary action or waves of fine
muscular contraction along the surface of the foot. Mucous secreted at the foot
helps the animals glide over the substrate. Some, such as members of the genus
Strombus, use their claw-like operculum to “pole” themselves along the bottom
with what appears to be a loping motion. Some species have an escape reac-
Above- The molluscs have theirbasic tion when a potential predator is detected, which consists of a rapid rolling or
morphology modified in (from top to leaping action, usually with the aid of the operculum.
bottom) cephalopods, snails, chitons,
clams and nudibranchs. In these Shell shape can indicate a lot about the habitat of a gastropod. The
drawings the foot is green, the shell is
blue, the gills are yellow and the spiny species of Murex are inhabitants of soft, muddy bottoms, their spines
mouth is orange. helping to protect them from predators or possibly to aid in prey capture (usu-
ally other molluscs). Limpets and abalones have low, broad
shells which offer less resistance in high wave habitats.
163
726 - Turbo petholatus * Papua New Guinea 727 - Serpulorbis grandis * Papua New Guinea
728 - Dendroderma maxima * Papua New Guinea 729 - Cerithium aluco * Indonesia
730 - Thyca crystallina * Philippines 731 - Lambis scorpius * Federated States of Micronesia
733 - Strombus dentatus * Strombidae * Mesogastropoda *
Marshall Islands * Kwajalein Atoll * lagoon pinnacle * 33 ft (10
m). This is a small species of Strombus which has a glossy shell and
reaches about 2.6 inches (60 mm) in length. Species of Strombus
have their eyes on stalks.
165
738 - Coriocella sp. * Palau 739 - Cypraea annulus * Federated States of Micronesia
740 - Cypraea annulus * Federated States of Micronesia 741 - Cypraea arabica * Hong Kong
167
750 - Cypraea tigris * Marshall Islands 751 - Cypraea vitellus * Indonesia
752 - Cypraea vitellus * Federated States of Micronesia 753 - Cypraea chinensis * Palau
169
762 - Casmaria erinaceus * Philippines 763 - Malea pomum * Papua New Guinea
766 - Charonia tritonis * Marshall Islands 767 - Cymatium aquatile * Federated States of Micronesia
species is often seen with a thin orange periostracum covering the
outside of the shell, which is not the shell’s true color.
171
774 - Pleuroploca trapezium * Indonesia 775 - Oliva reticulata * Indonesia
173
786 - Conus circumcisus * Papua New Guinea 787 - Conus crocatus * Marshall Islands
788 - Conus eburneus * Federated States of Micronesia 789 - Conus floccatus * Marshall Islands
790 - Conus geographus * Papua New Guinea 791 - Conusimperialis * Federated States of Micronesia
795 - Conus textile * Conidae * Neogastropoda * Philippines *
Cebu * Mactan Island * 20 ft (6 m). Most cones with a tented pat-
tern on the shell, such as C. textile, feed on other molluscs. This
species hunts at night. It buries in the sand during the day.
175
799 - Janthina janthina * Central Pacific 800 - Melanella sp. * Papua New Guinea
801 - Chelidonura electra * Papua New Guinea 802 - Chelidonura hirundinina * Philippines
803 - Chelidonura inornata * Federated States of Micronesia 804 - Chelidonura varians * Indonesia
804 - Chelidonura varians * Aglajidae * Cephalaspidea *
Opisthobranchia * Indonesia * Manado * 30 ft (9 m).
808 - Bulla ampulla * Philippines 809 - Elysia ornata * Federated States of Micronesia
810 - Elysia sp. * Federated States of Micronesia 811 - Plakobranchus ocellata * Hawaii
177
812 - Plakobranchus sp. * Philippines 813 - Cyerce sp. * Palau
816 - Dolabella auricularia * Papua New Guinea 817 - Dolabella auricularia * Papua New Guinea
species of Aplysia can be relatively large, up to 4-8 inches (10-20
cm). They release a purple secretion when disturbed which is an irri-
tant and may deter predators.
179
824 - Pleurobranchus grandis * Palau 825 - Pleurobranchus peroni * Indonesia
826 - Pleurobranchus sp. * Philippines 827 - Ardeadoris egrettae * Papua New Guinea
833 - Halgerda sp. * Papua New Guinea 834 - Jorunna funebris * Palau
835 - Platydoris cruenta * Palau 836 - Platydoris scabra * Papua New Guinea
181
837 - Platydoris sp. * Papua New Guinea 838 - Ceratosoma moloch * Solomon Islands
839 - Ceratosoma trilobata * Papua New Guinea 840 - Chromodoris albopunctata * Marshall Islands
841 - Chromodoris annae * Papua New Guinea 842 - Chromodoris annulata * Papua New Guinea
841 - Chromodoris annae * Chromodorididae * Nudibranchia *
Papua New Guinea * Madang * lagoon patch reef * 40 ft (12 m).
845 - Chromodoris kunei * Papua New Guinea 847 - Chromodoris lochi * Indonesia
846 - Chromodoris lineolata * Papua New Guinea 848 - Chromodoris magnifica * Philippines
849 - Chromodoris reticulata * Papua New Guinea 850 - Chromodoris willani * Philippines
183
851 - Chromodoris cf. tinctoria * Marshall Islands 852 - Chromodoris sp. * Indonesia
853 - Chromodoris sp. * Indonesia 854 - Chromodoris sp. * Papua New Guinea
860 - Hypselodoris kanga * Federated States of Micronesia 861 - Hypselodoris mardadilus * Marshall Islands
862 - Miamira sinuata * Palau 863 - Risbecia imperialis * Papua New Guinea
185
864 - Reticulidia fungia * Federated States of Micronesia 865 - Hexabranchus sanguineus * Philippines
866 - Hexabranchus sanguineus * Papua New Guinea 867 - Hexabranchus sanguineus eggs * Papua New Guinea
868 - Fryaria ruppeli * Indonesia 869 - Phyllidia babai * Papua New Guinea
previous, demonstrates this. It is possible that much of the color vari-
ation can be attributed to changes during growth, rather than specif-
ic differences.
872 - Phyllidia madangensis * Papua New Guinea 874 - Phyllidia cf. ocellata * Papua New Guinea
873 - Phyllidia ocellata * Palau 875 - Phyllidia cf. ocellata * Federated States of Micronesia
876 - Phyllidia pustulosa * Philippines 877 - Phyllidia tula * Indonesia
187
878 - Phyllidia varicosa * Philippines 879 - Phyllidia sp. * Papua New Guinea
880 - Phyllidia sp. * Papua New Guinea 881 - Phyllidia sp. * Papua New Guinea
882 - Phyllidia sp. * Philippines 883 - Phyllidiella sp.* Papua New Guinea
875 - Phyllidia cf. ocellata * Phyllidiidae * Nudibranchia *
Opisthobranchia * Federated States of Micronesia * Chuuk Atoll
* lagoon reef * 100 ft (30 m).
189
890 - Nembrotha sp. * Philippines 891 - Nembrothapurpureolineata * Indonesia
892 - Tambja morosa * Federated States of Micronesia 893 - Notodoris minor * Papua New Guinea
894 - Notodoris minor eggs * Papua New Guinea 895 - Notodoris minor * Philippines
892 - Tambja morosa * Polyceridae * Nudibranchia *
Opisthobranchia * Federated States of Micronesia * Chuuk *
lagoon * 85 ft (25 m).
191
902 - Cuthona cf. sibogae * Indonesia 903 - Phyllodesmium briareus * Papua New Guinea
904 - Phyllodesmium longicirra * Papua New Guinea 905 - Pteraeolidia ianthina * Marshall Islands
906 - Pteraeolidia ianthina * Philippines 907 - Arca ventricosa * Papua New Guinea
(12 m). This species is the largest aeolid in Hawaii, but it is still rel-
atively small, only a few inches long and very slender. It is wide-
spread in the Indo-Pacific. It eats hydroids, including the widespread
Halichordyle disticha, and stores the nematocysts for future use.
This species also contains zooxanthellae, which may account for
many of the color differences found among individuals.
193
914 - Pinctada maxima * Papua New Guinea 915 - Pteria penguin * Federated States of Micronesia
916 - Pteria sp. * Federated States of Micronesia 917 - Pedum spondyloideum * Palau
918 - Spondylus sp. * Federated States of Micronesia 919 - Spondylus sp. * Indonesia
919 -Spondylus sp. * Spondylidae * Bivalvia * Indonesia * Biak
Island * reef * 40 ft (12 m).
927 - Lopha frons* Ostreidae * Bivalvia * Hong Kong * Shek Ngau Chau
* 33 ft (10 m). This species has a velvety black mantle with a striking thin
white edge. It was found on the under surfaces of huge blocks of rock on an
offshore island in Mirs Bay, Hong Kong. We have not yet been able to iden-
tify it, but it is almost certainly a member of the oysters (Ostreidae).
195
925 - Hyotissa sp. * Papua New Guinea 926 - Lopha cristagalli * Federated States of Micronesia
929 - Chama lazarus * Federated States of Micronesia 930 - Unidentified bivalve * Palau
cea and T. squamosa. It has numerous close, set scutes on the shell
valves, and most often bores into the reef, though not as deeply as T.
crocea, so that the scutes are still visible. It reaches about fifteen
inches in length. The mantle is variable in pattern and often bright-
ly colored, though not usually iridescent. It has the widest distribu-
tion of all the giant clams, from east Africa to Polynesia.
197
937 - Tridacna derasa * Fiji
938 - Tridacna gigas * Marshall Islands 939 - Periglypta “clathrata” * Papua New Guinea
940 - Nautilus pompilius * Papua New Guinea 941 - Metasepia pfefferi * Indonesia
945 - Sepia latimanus * Sepiidae * Sepioidea * Cephalopoda *
Indonesia * Ruang Island * 33 ft (10 m).
199
948 - Sepioteuthis eggs * Indonesia 949 - Hapalochlaena lunulata * Indonesia
950 - Octopus macropus * Bahrain 951 - Octopus sp.* Federated States of Micronesia
Crustaceans
Most of the larger crustaceans found in the shallow Pacific tropics have
planktonic larvae, which is one reason for the wide geographic distribution of most
species. In crustaceans such as spiny lobsters, the planktonic larval stage can last as
long as 6 months, ample time for currents to carry larvae thousands of miles. In other
crustaceans planktonic larvae are short-lived or or absent, and these species have
more restricted distributions.
rocky shores. Those found on intertidal rocks are gener- with the distasteful sponge.
ally fast and agile, scurrying over the rocks both above Finally, there are some other orders of non-deca-
and below the water. The mud dwellers, typified by the pod, generally small crustacea which are abundant on
fiddler crabs Uca, live in burrows they dig in the mud. reefs, but not very apparent to human visitors. These
Some slow species have developed various means of include the mysid shrimps, amphipods, isopods and
camouflage or deception. Some are cryptic, closely copepods. Mysids occur in dense schools in and around 205
resembling a mixed algal bottom, where they hide. crevices and caves, looking more like baby fishes than
Other actually employ pieces of algae and invertebrates, crustaceans. Some copepods have similar habits, form-
attaching these materials to their exoskeleton (decorator ing dense swarms of individuals even smaller than the
crabs), to aid in their cryptic endeavors. A few species mysids in the water above and near crevices of reefs.
attach or hold, with their modified last legs, pieces of Amphipods are occasionally found on reefs, living on
sponge on the top of the carapace to discourage predators the bottom in groups. Isopods are most evident on reefs
as parasites of fishes.
Above- This photograph shows an advanced larval stage of a slip- Above- This is the megalopa or last larval stage of a crab. The
per lobster in the process of settling on a reef after its life of sev- larva swims in the plankton, it is about one half inch in length.
eral months in the plankton. The transparent nature of this and After it settles to the bottom, it may walk fora short time until
most crustacean larvae is a benefit for living in the open water of it molts and completes the transformation into a juvenile crab.
the planktonic environment. Such transparent plankton stages quickly produce pigment
after taking up residence on the bottom.
954 - Nymphon sp. * Pycnogonida * Chelicerata * Hawaii * Puako
* on sponge * 20 ft (6 m). Many pycnogonids are small, like these
in the photograph, and are tricky to spot. This individual was found
living on a sponge.
207
960- Caprella sp. * Philippines 961- Lepas sp. * Federated States of Micronesia
962- Unidentified Barnacle * Federated States of Micronesia 963- Megabalanus sp. * Federated States of Micronesia
964- Barnacle Pyrgomatidae * Philippines 965- Barnacle Pyrgomatidae* Federated States of Micronesia
968 - Euchaeta sp. * Copepoda * Crustacea * Hawaii * Open
Pacific. Copepods can be found, with careful examination, in the
water column. Most species are less than a quarter of a inch long.
Copepods are primary consumers of phytoplankton and are in turn an
important food source for other carniverous zooplankton.
209
972- Thalassina anomala * Philippines 973- Ranina ranina * Indonesia
974- Birgus latro * Fiji 975- Aniculus maximus * Papua New Guinea
976- Dardanus guttatus * Federated States of Micronesia 977- Dardanus megistos * Fiji
These unusual hermit crabs do not occur in mollusc shells, but live in
tubes in Millepora sp. fire coral. They are tiny and filter food from
the water using their feathery antennae. Paguritta harmsi occurs on
the Great Barrier Reef where it lives in dead serpulid worms tubes in
corals.
211
984- Porcellanella picta * Indonesia 985- Unidentified porcellanid
213
996- Parthenope validus * Hong Kong 997- Portunus sp. * Hong Kong
998- Lissocarcinus orbicularis * Hawaii 999- Charybdis sp. * Federated States of Micronesia
215
1008- Trapezia rufopunctata * Federated States of Micronesia 1009- Trapezia sp. * Palau
1010- Zosimus aeneus * Indonesia 1011- Unidentified crab * Federated States of Micronesia
1012- Etisus sp. * Federated States of Micronesia 1013- Unidentified crab * Palau
claws useful in picking up algae to convey it to the mouth.
1013 - Unidentified crab * Xanthidae * Decapoda * Palau *
Lighthouse Reef * night * 10 ft (3 m). In many xanthids the fingers
of the claw are black. This species has the opposite situation with the
fingers being lighter than the rest of the claw.
217
1020- Matuta lunaris * Philipines 1021- Metapenaeus sp.* Philippines
1024- Stenopus hispidus * Federated States of Micronesia 1025- Stenopus pyrsonotus * Papua New Guinea
Papua New Guinea * West New Britain * Farmers Reef * night *
33 ft (10 m). Despite being a different species, this coral shrimp has
the “look” which immediately identifies it as a member of Stenopus.
There are several species in the genus not included here.
219
1032- Thor amboinensis * Philippines 1033- Rhynchocinetes conocolor * Federated States of Micronesia
1034- Rhynchocinetes conocolor * Federated States of Micronesia 1035- Rhynchocinetes conspiciocellus * Hong Kong
1040- Periclimenes amboinensis * Marshall Islands 1044 - Periclimenes imperator female * Palaemonidae * Caridea *
Papua New Guinea * Kalihi Harbor * 20 ft (6 m). This is one of
the best known of the commensal Periclimenes, being found on
opisthobranchs, including Pleurobranchus forskali, and the spanish
dancer nudibranch, Hexabranchus sanguiensis. The male and female
differ in color and generally only a single pair are found per mollusc.
221
1044- Periclimenes imperator * Papua New Guinea 1045- Periclimenes imperator * Papua New Guinea
1046- Periclimenes kororensis * Palau 1047- Periclimenes soror * Papua New Guinea
223
1056- Alpheus djiboutiensis * Marshall Islands 1057- Unidentified alpheid * Marshall Islands
1058- Synalpheus carinatus * Marshall Islands 1059- Unidentified alpheid * Federated States of Micronesia
1064- Panulirus versicolor * Palau 1069 - Parribacus antarcticus * Scyllaridae * Palinura * Federated
States of Micronesia * Chuuk * lagoon reef * 50 ft (15 m). This
species grows to about 8 inches (20 cm) in length.
225
1068- Arctides regalis * Hawaii 1069- Parribacus antarcticus * Federated States of Micronesia
Lophophorates
The first three phyla in this section are superficially quite
different. However, because they share a common feeding structure,
the lophophore, they are grouped together. The lophophore is a ciliated tentacular
crown surrounding the mouth. They have other similarities: their general body
plan, a U-shaped gut, a transient reproductive system, and outer casings, including
tubes, with compartments or shells. The fourth phylum, the Kamptozoa, is a close-
ly related group.
Phoronids are found only in marine waters, living in chitinous tubes which
they secrete. There are only two genera (Phoronis and Phoronopsis) with about 15
species. The lophophore functions in feeding, respiration and protection.
Tentacular ciliary bands filter particulates from the water and deliver them to the
mouth. The gut is U-shaped, with the mouth at the base of the lophophore and anus
just outside the lophophore. Each tentacle of the lophophore contains a coelonic
extension. Phoronids have a free-swimming larvae, called an actinotroch, which
usually has a lengthy existence in the plankton. In our region, phoronids are most
often seen as associates of various other invertebrates such as sponges and tube
anemones.
Members of the Phylum Ectoprocta, usually called bryozoans from the out-
moded Phylum name Bryozoa, are sessile colonial animals which encrust on rocks
and various living organisms and resemble algae, hence their common name “moss
animals”. The colonies are composed of zooids, in essence replicated individuals.
Like hydroid polyps and individuals of other colonial animals the zooids of many
bryozoans are polymorphic (different in form and function). Autozooids occur in a
horny or calcified exoskeleton, sometimes with a small door (operculum) which can
cover the opening where the lophophore is extended. Zooids specialized for feed-
ing, the autozooids, have a U-shaped gut with the mouth inside the lophophore, sim-
ilar to that of the phoronids, and the anus opening outside the lophophore. Other
types of zooids include avicularia, which have the operculum modified into a jaw;
Left- This bright red bryozoan Tropidozoum cellariiforme, it is hard to distinguish from a cal-
careous red algae. Only when carefully examined is the zooid structure of the fine flexible
branches evident. The species is found on drop offs and sloping reefs in the Philippines.
and marine species. The marine
members have traditionally been
considered as comprising the Class
Gymnolaemata. Overall the bry-
ozoans are one of the most poorly
known groups of marine inverte-
brates on tropical Pacific reefs.
Inner reef flats on the Great Barrier
Reef, although not an ideal habitat
for bryozoans, are known to support
at least 80 species, while a similar
number of species is known from
Enewetak Atoll in the Marshall
Islands. Studies from Chuuk Atoll
indicate that perhaps as many as
300-400 species might occur in that
environmentally diverse area.
Many species, even from shallow
water, remain undescribed. Among
known species many appear to have
broad geographic ranges, often
from Hawaii to the western Indian
Above- This bryozoan, Caulibugula intermis seems to like areas with strong currents. In Ocean within the tropical and sub-
Palau it is found on the bottom and sides of deep channels between the lagoon and ocean. tropical belts.
The flower-like structures bear the clonial zooids.
Recently a potential anti-cancer
and vibracula, with the operculum modified into a bris- compound called bryostatin was isolated from the bry-
tle. ozoan Bugula neretina and is presently undergoing clin-
228 Sessile colonies of zooids are produced by asex- ical trials. There may well be other compounds of med-
ual budding. The structure and repetitive nature of the icinal value in the bryozoans.
zooids quickly allow determination of whether the
organism in question is a bryozoan or not. Some bry-
ozoans are encrusting on mangrove roots or rocks.
Some species can be heavily calcified, finely branched,
often white in color, and could easily be confused with
stylasterine corals (Stylasterina). Bryozoans are often
surprisingly common on reefs.
Above- The ciliated tentacles of the zooids of this bryozoan are Above- Bryozoans are colonial animals, consisting of many small
easily seen protruding out of the opening of the exoskeleton. The zooids, each encased in a chitinous cup, and then organized into
exoskeletons of bryozoans can be somewhat calcified, in some delicate colonies. Each zooid has a lophophore with tentacles
species to the point where the resemble small coral colonies. which protrude when the animal is feeding.
Phylum Brachiopoda- lamp shells or brachiopods.
The brachiopods, commonly called lamp shells,
consist of only about 300 living species, but over 12,000
extinct species dating back 600 million years are known
from the fossil record. During that time brachiopods
have changed little; they are truly living fossils.
Brachiopods superficially resemble bivalve molluscs in
having a bivalved calcareous shell, but they are actually
quite different. The mantle cavity has coiled arms, the
brachia, that bear the lophophore which is used in col-
lecting suspended food particles. The body is organized
similarly to the phoronids, hinting at a common ances-
try. It has been suggested competition from bivalve
molluscs after the Paleozoic led to the gradual decline of
the brachiopods.
Brachiopods are solitary and live in benthic
marine environments. There are two basic types. In the
articulate brachiopods, the body is enclosed in hinged,
dorsoventrally-oriented calcium carbonate valves
(shells), which contrast with the laterally oriented shells
of bivalve molluscs. They attach to the substratum by
the pedicle, a fleshy foot, (although some lack it) and
normally sit ventral side up. Most occur in caves or in
sheltered areas beneath boulders. In the second group,
the inarticulate brachiopods, the valves are unhinged
and composed of calcium phosphate, plus chitin and
protein. Most inarticulates occur in sand and mud and
the pedicle is adapted for burrowing and anchoring in
soft substrata. 229
One species, Lingula reevei, can be particularly
common. It lives in vertical sand burrows in shallow
water; its presence is evident by a three (two incurrent, Above- - The small brown polychaete-like animal on the lower
one excurrent) siphonal openings visible on the sand left side of the cerianthid tube is a phoronid, Phoronis australis.
surface. It is widespread in the Pacific, including Phoronids are one of the few organisms that live on the tube of
Hawaii, Indonesia, and the Philippines. the cerianthid anemones. This species is sometimes found liv-
ing in soft sediment as well.
231
1079 - Serripetraliella sp. * Palau 1080 - Iodictyum sp. * Federated States of Micronesia
1081 - Reteporella sp. * Papua New Guinea 1082 - Schizoporella serialis * Federated States of Micronesia
1083 - Stylopoma sp. * Papua New Guinea 1084 - Catenicella sp. * Papua New Guinea
1084- Catenicella sp. * Scrupocellariidae * Cheilostomata *
Papua New Guinea * Port Moresby * barrier reef * 60 ft (18 m).
This is an undescribed species we have found in Papua New Guinea
and Palau. It is a lovely bryozoan, a golden brown tuft of the softest
sort of branches. It is most abundant along deep reef dropoffs and
varies greatly in density along any given section of wall.
233
1091 - Zoobotryon sp. * Papua New Guinea 1092 - Bugula sp. * Federated States of Micronesia
1095 - Frenulina sanguinolenta * Federated States of Micronesia 1096 - Lingula reevi * Philippines
234
Phylulm Echinodermata
Echinoderms
All adult Echinoderms display a five-part radial
symmetry, which seems to serve their sedentary benthic lifestyle
well. They also possess an internal skeleton of calcium carbonate
plates, called ossicles, and a water vascular system unique to echino-
derms. The water vascular system consists of a series of canals which radiate
throughout the body and terminate in structures called tube feet. The tube feet
penetrate the body wall and often have a tiny suction or adhesive cup at their tip.
Body fluid, with an ionic composition close to seawater, circulates through the water vascular system for
hydraulic expansion or contraction of the tube feet. Tube feet serve echinoderms in a variety of ways, 235
including adhesion, locomotion, feeding, and respiration.
The crown-of-thorns starfish, Acanthaster planci, (above) is probably the single most important echinodermin influencing the
nature of Pacific reef communities. This large species reaches 20 inches (50 cm) in diameter. The long spines on the arms and disk
are sharp, stiff and venomous, easily penetrating the skin of a diver who accidently steps on or is thrown by a wave against this starfish. 239
It may be the only venomous starfish in the world. Long a part of the normal community on Pacific coral reefs, the crown-of-thorns is
a predator of stony corals. It feeds by everting its stomach out of its mouth in a thin sheet that covers all or part of a coral colony and
digesting the living coral tissue from the skeleton. After several hours, the starfish moves away leaving a portion of dead coral which
is starkly white. This feeding site soon becomes darkened with algae, but remains apparent for many days. In low numbers the crown-
of-thorns is a predator which helps to keep the balance between stony corals and the many other organisms competing for space on the
reef. In the 1960’s or early 1970’s, though, conditions changed on some Pacific reefs which allowed the population of starfish to
explode to “plague” levels. The starfish devastated the coral communities of many reefs and entire islands. The “plague” levels of
Acanthaster receded in later years, but the damage has remained on many reefs which have not regenerated to any significant degree.
Large populations, although not at previous “plague” levels, still remain on many reefs in Micronesia and elsewhere. Whether these are
equilibrium levels or not is a matter of conjecture.
How these waves of starfish abundance occur is still largely a mystery. A single individual can produce as many as 65 mil-
lion eggs in a spawning season. Small changes in the survival of larvae can have a magnifying effect on the number of juveniles sur-
viving. The most widely accepted theory suggests that increased phytoplankton production, perhaps from increased “fertilization” of
waters from runoff from agricultural land or other human activities, resulted in increased larval survival with resultant increases in juve-
nile and adult populations. It has also been suggested that waves of starfish abundance are natural fluctuations, not the result of human
activities, and have occurred many times in the past.
The effect of the crown-of-thorns on reefs goes far beyond the simple predation on coral. When large amounts of coral are
killed, algae colonize the bare coral skeletons. The many organisms which live in and around coral colonies have lost their proper habi-
tat and become rare or disappear from a reef. Herbivorous fishes and other invertebrates feed upon the benthic algae, their scraping
and biting eroding the reef surface. Some organisms increase in abundance, but many others are reduced, and the overall result is a
decrease in the diversity of organisms living on a given reef. If all the reefs in an area are affected, then entire species may disappear
where they previously occurred in abundance.
There are few predators of the crown-of-thorns. The trumpet triton shell is known to eat the starfish, but populations of this
gastropod are never very high and have potentially been decreased by shell collecting. The small paddle shrimp, Hymenocera picta,
will kill and eat Acanthaster if starving, but much prefers other starfish prey such as Linckia. Only a single fish, the humphead wrasse,
Cheilinus undulatus, is known to eat Acanthaster, but again populations of that large tasty fish are reduced in many areas, eliminating
predation as an effective biological control on the starfish.
Where juvenile crown-of-thorns occur was a mystery for a number of years. They were finally found to be sheltering under
rocks and rubble during daytime and were not normally visible. This illustrates how often we are ignorant of even the most basic facts
about the organisms of the sea.
The crown-of-thorns occurs from the east coast of Africa and the Red Sea across the tropical Indian and Pacific Oceans to
the west coast of the Americas, from the Sea of Cortez south to Panama. It is not found in the Galapagos. Some authorities consider
eastern Pacific populations to represent a separate species.
1097 - Comanthina schlegelii * Palau 1098 - Comanthus alternans * Indonesia 1099 - Comanthus parvicirrus * Papua New
Guinea
241
1103 - Comaster gracilis * Marshall Islands 1104 - Comaster multifidus * Marshall Islands
1105 - Comaster multibrachiatus * Indonesia 1106 - Comatella maculata * Papua New Guinea
1107 - Comatella stelligera * Papua New Guinea 1108 - Comissia pectinifer * Indonesia
1109 - Oxycomanthus bennetti * Papua New 1110 - Colobometra perspinosa * Papua New 1111 - Cenometra bella * Indonesia
Guinea Guinea
exact identity of the photographed individual cannot be determined. Caledonia, but is not known from the Marshall Islands. In Palau it is
It is resting among Anacropora coral branches. Known from not common, but can sometimes be found in large numbers else-
Indonesia, Australia, Papua New Guinea, Philippines and Palau, where in the western Pacific.
where it is not common.
1113 - Himerometra robustipinna * Himerometridae * Crinoidea
242 1108 - Comissia pectinifera* Comasteridae * Crinoidea * * Indonesia * Manado * fringing reef * 60 ft (18 m). This species
Indonesia * Biak Island * fringing reef * 50 ft (15 m). This sits in the open on corals near the reef crest, both day and night. The
crinoid has long arms, which it extends up from the bottom leaving deep red color is typical. It is known from Sri Lanka, Australia, the
the disk protected. Its occurs in the East Indies, Micronesia and Philippines and Indonesia.
Australia.
1114 - Liparometra regalis * Mariametridae * Crinoidea *
1109 - Oxycomanthus bennetti * Comasteridae * Crinoidea * Marshall Islands * Kwajalein Atoll * 60 ft (18 m). The photo-
Papua New Guinea * Madang * barrier reef * 33 ft (10 m). This graph shows clearly how crinoids hold on with their cirri. This
species is often placed in the genus Comanthus. It is a very common species was not previously known from the Marshall Islands, the
crinoid which is active and exposed both day and night. Hence their geographic distribution is poorly known.
appearance as shown in the photograph is the rule. There are sever-
al color varieties. This species likes currents and it is very common 1115 - Stephanometra indica * Mariametridae * Crinoidea *
in passes and other areas with good water flow. It hangs on with its Federated States of Micronesia * Chuuk Atoll * Yanagi Island *
cirri to coral, rocks or gorgonians. It can occur singly or in aggrega - night * 6 ft (2 m). This is a common crinoid on some inshore reefs
tions and is known from the eastern Indian Ocean to the Marshall in Chuuk, climbing up corals and gorgonians at night to feed. During
Islands. the day they hide among the coral branches. In the photograph one
individual can be clearly seen hanging on to the coral with its cirri.
1110 - Colobometra perspinosa * Colobometridae * Crinoidea * This species does not get very large, only about one foot across. It
Papua New Guinea * Madang * barrier reef * 66 ft (20 m). This occurs over a broad area of the region, but the exact limits are not
species lives on gorgonians and antipatharians, as is shown in the known.
photograph, usually in reef waters below 66 ft (20 m). It has ten arms
and a number of color varieties. It is known from Papua New 1116 - Genus species unknown * Mariametridae * Crinoidea *
Guinea, Australia, Lord Howe Island and Fiji. Federated States of Micronesia * Nama * reef face * night * 30 ft
(9 m). This unidentified species is clearly seen holding on to the
1111 - Cenometra bella * Colobometridae * Crinoidea * Indonesia reef with its cirri.
* Biak Island * fringing reef * 66 ft (20 m). This species is usu-
ally found on gorgonians and antipatharians. The photograph shows 1117 - Genus species unknown* Mariametridae * Crinoidea *
an individual holding on to a whip black coral of the genus Papua New Guinea * Madang * barrier reef * 60 ft (18 m).
Cirrhipathes with its cirri. Crinoids with such cirri are able to climb
out into open water where filter feeding is better than close to the 1118 - Genus species unknown* Mariametridae * Crinoidea *
reef. This species is also capable of active swimming. It has sever- Papua New Guinea * West New Britain * Kimbe Bay * night * 33
al color varieties and generally between 19 and 39 arms. It is known ft (10 m).
from the western Pacific, including Indonesia, Vietnam, the
Philippines and Marshall Islands. 1119 - Luidia cf. avicularia * Luidiidae * Asteroidea * Philippines
* Cebu * Mactan Island * 3 ft (1 m).
1112 - Oligometra serripinna * Colobometridae * Crinoidea *
Papua New Guinea * Madang * Cape Croiselles * 3 ft (1 m). 1120 - Archaster typicus * Archasteridae * Asteroidea * Indonesia
This small crinoid has only ten arms and it is almost always found on * Manado * sand bottom * 6 ft (2 m). This species is very com-
gorgonian fans or wire corals such as Cirrhipathes sp. It is known mon on sandy bottoms inshore. The starfish is often slightly buried
from the Red Sea and east Africa to the Philippines, Palau and New in sand and only the general impression in the surface of the sand
1112 - Oligometra sirripinna * Papua New Guinea 1113 - Himerometra robustipinna * Indonesia
243
1114 - Liparometra regalis * Marshall Islands 1115 - Stephanometra indica * Federated States of Micronesia
1116 - Unidentified crinoid * Federated States of Micronesia 1117 - Unidentified crinoid * Papua New Guinea
1118 - Genus species unknown* Papua New Guinea 1119 - Luidia cf. avicularia * Philippines
may be visible. It has numerous gastropod parasites. A. typicus is
known from southeast Asia, Melanesia, Australia and western
Polynesia, but not from eastern Polynesia. In the Indian Ocean it is
replaced by Archaster angulatus, a similar species.
245
1127 - Protoreaster nodosus * Indonesia 1128 - Culcita novaeguineae * Marshall Islands
1129 - Culcita novaeguineae * Marshall Islands 1130 - Celerina heffernani * Papua New Guinea
1131 - Celerina heffernani * Papua New Guinea 1132 - Fromia indica * Indonesia
1133 - Fromia cf. milleporella * Marshall Islands 1134 - Fromia monilis * Philippines
247
1140 - Linckia guildingi * Papua New Guinea 1141 - Linckia laevigata * Federated States of Micronesia
1144 - Linckia multifora * Federated States of Micronesia 1145 - Linckia sp. * Philippines
by small gastropod molluscs of the genera Stylifer and Thyca, which
can be found by examining the underside and arms of the starfish
(see Mollusc section for photograph). It is common throughout the
Indo-Pacific, particularly in areas with wave action.
1149 - Nardoa tuberculata * Indonesia 1150 - Nardoa tuberculata * Papua New Guinea
1151 - Neoferdina cumingi * Papua New Guinea 1152 - Mithrodia clavigera * Papua New Guinea
249
1153 - Thromidia catalai * Papua New Guinea 1154 - Acanthaster planci * Palau
1155 - Acanthaster planci * Federated States of Micronesia 1156 - Echinaster callosus * Federated States of Micronesia
large starfish, the arms a foot (30 cm) or more in length, and looks
more like an overstuffed toy than a living organism. It is known from
Guam and the Bonin Islands to New Caledonia and Hawaii.
251
1166 - Ophiothrix nereidina * Solomon Islands 1167 - Ophiothrix sp. * Marshall Islands
253
1179 - Ophiarthrum elegans * Marshall Islands 1180 - Ophiomyxa sp. * Marshall Islands
255
1192 - Astropyga radiata * Indonesia 1193 - Diadema savignyi * Papua New Guinea
1196 - Mespilia globulus * Papua New Guina 1197 - Temnopleurus toreumaticus * Bahrain
1193 - Diadema savignyi * Diadematidae * Echinoidea * Papua
New Guinea * Port Moresby * Bootless Bay * 50 ft (15 m).
257
1204 - Colobocentrotus mertensi * Mariana Islands 1205 - Echinometra mathaei * Indonesia
1206 - Echinostrephus sp. * Papua New Guinea 1207 - Anthocidaris crissipinina * Hong Kong
1208 - Heterocentrotus mammillatus * Marshall Islands 1209 - Heterocentrotus trigonarius * Marshall Islands
(3 m). During the day these urchins hide in cracks and crevices in
shallow water in wave swept areas on outer reefs. At night they leave
their shelter holes and graze over the reef. They are known for their
stout spines which remain intact after the urchin dies, often washing
up on beaches or being found in reef sediments. The species is
known from Hawaii through the tropical Pacific and Indian Oceans
to the coast of Africa. Their spines are often used in handicrafts.
259
1216 - Maretia planulata * Marshall Islands 1217 - Maretia planulata * Philippines
1222 - Actinopyga miliaris * Federated States of Micronesia 1223 - Actinopyga palauensis * Holothuridae * Aspidochirotida *
Federated States of Micronesia * Chuuk Atoll * Eten Island * 40
ft (12 m). This dark sea cucumber is common in lagoon areas of
Chuuk and Palau.
261
1228 - Bohadschia marmorata * Federated States of Micronesia 1229 - Bohadschia sp. * Federated States of Micronesia
1230 - Holothuria atra * Hawaii 1231 - Holothuria edulis * Federated States of Micronesia
1232 - Holothuria flavomaculata * Federated States of Micronesia 1233 - Holothuria fuscopunctata * Papua New Guinea
1235 - Holothuria leucospilota * Holothuridae * Aspidochirotida
* Hong Kong * Cape d’Aguilar * sand and rock bottom * 3 ft (1
m). This is the most common holothurian in Hong Kong and is
found throughout the tropical Indo-Pacific.
263
1240 - Stichopus “variegatus” * Palau 1241 - Stichopus variegatus * Federated States of Micronesia
1242 - Stichopus sp. * Federated States of Micronesia 1243 - Tyrone sp. * Hong Kong
1244 - Thelenota ananas * Papua New Guinea 1245 - Thelenota anax * Marshall Islands
Dendrochirotida * Papua New Guinea * Port Moresby * 66 ft (20
m). This certainly is one of the more unusual sea cucumbers as it
looks more like a sea anemone than anything else. The white body
is buried in sand and only the branchial tree protrudes anemone-like
above the bottom. The arms of the tree capture plankton and food
particles and are inserted, one at a time, into the mouth, and the food
items sucked off as the arm is pulled out of the closed mouth. This
behavior must be seen to be appreciated.
265
1252 - Euapta godeffroyi * Papua New Guinea 1253 - Opheodesoma sp. * Philippines
1254 - Opheodosoma spectabilis * Hawaii 1255 - Opheodesoma sp. * Papua New Guinea
1256 - Synaptula lamperti * Papua New Guinea 1257 - Synaptula lamperti * Federated States of Micronesia
266
Phylum Chordata
Ascidians
The Phylum Chordata includes not only vertebrates (which are
beyond the limits of this book) but also invertebrates. Of the invertebrate chor-
dates, the tunicates, comprising the subphylum Urochordata, are important
inhabitants of the shallow water tropical Pacific. Several classes of tunicates (e.g.
Pyrosoma, Salpa, Doliolida and Appendicularia) are entirely pelagic and will be
covered briefly. The remaining class, Ascidiacea will be treated in most detail here
since they are very common benthic animals throughout the tropical Pacific.
Ascidians take many different forms, but can be conveniently divided into solitary and colo-
nial species. The solitary species are easiest to recognize and they are often relatively large and usu-
ally have two readily apparent siphons, even when the body is covered with other organisms growing
on the surface of the ascidians. The solitary species range from the size of a grain of rice to the dimen-
sions of a soccer ball. Solitary ascidians generally live as isolated individuals, but sometimes occur in
such high densities that they resemble colonial species. The colonial species are made up of small
individual units, called zooids, which can number many thousands in a large colony. They grow as
sheets, stalked bouquets, large masses and lumps, and occur on many types of living and dead sub-
strata including live coral, dead coral, rock, sponges, gorgonians, and other ascidians. Some hang,
appearing to drip like candle wax, from whip coral or gorgonians. They occur in a bewildering vari-
ety of color and patterns, often with a great deal of intraspecific variation, producing some of the most
spectacular visual treats to be found in the ocean.
Opposite- The orange bodies of these colonial ascidians, Didemnum sp. stand out on this reef wall in the Philippines.
Ascidians are found in many reef environments, but their presence is often overlooked by divers in favor of the more
familiar hard and soft corals.
The siphons, particularly in the solitary species,
are equipped with sphincters which can close the open-
ing in an instant if the ascidian is disturbed or touched.
This is their only possible response to potential danger,
since they cannot flee and have no defensive structures
like pincers or teeth. Some also react to the approach of
a diver by siphon closure, either through detection of the
pressure wave produced by a large object moving
through the water or by changes in light. The sensory
perception abilities of ascidians are not understood,
although cells supposedly sensory in nature have been
described from the epithelium of the body wall. It is
uncertain what stimuli they can detect, however.
Above- Some colonial ascidians occurs as a Two genera of unicellular algae, the prochlorophyte Prochloron
single stalk with head containing many zooids and the procyanophyte Synechocystis, are symbiotic in ascidians.
on it (Phillipines). Members of Prochloron are found on and near the ascidian’s surface,
where they can be easily dislodged by rubbing the
colony, and within the cloacal chambers of some
species, such as the very common Didemnum molle.
Additionally, Prochloron and Synechocystis occur with-
in the tissues and tunic of some didemnids, the latter
alga tending to color colonies pink with their red pig-
ments. Interestingly, the ascidians do not harbor the
symbiotic dinoflagellate Symbiodinium microadriaticum
(zooxanthellae), found in many other tropical inverte-
brates, including stony corals, sponges and cnidarians.
Above top- Two individuals of the salp Tethys vagina were pho- Above- The salp Pyrosoma atlanticum can reach lengths of more
tographed in open water. Above- Doliola sp. is similarin appear- than 10 ft (3 m) and is found in open water.
ance to salps, but belongs in the closely related doliiolids.
1258 - Aplidiopsis sp. * Polyclinidae * Aplousobranchia * Indonesia *
Manado * 23 ft (7 m). This beautiful ascidian, probably an undescribed
species, has a circular system of zooids around a common cloaca. This is a
series of colonies, the zooids embedded in a firm tunic, and growing among
sponges, soft corals and crinoids. Crinoid arms lie above the ascidian and a
contracted pink Dendronephthya soft coral is visible in a lower corner.
273
1264 - Aplidium sp. * Indonesia 1265 - Aplidium sp. * Papua New Guinea
275
1276 - Eudistoma reginum * Indonesia 1277 - Eudistoma reginum * Indonesia
1280 - Exostoma sp. * Federated States of Micronesia 1281 - Exostoma sp. * Federated States of Micronesia
1282 - Sigillina sp. * Indonesia 1283 - Didemnum sp.* Papua New Guinea 1284 - Didemnum molle * Federated States of
Micronesia
with a translucent head with the zooids. This species has only been found
along deep reef dropoffs.
1287 - Sigillina sp. * Polycitoridae * Aplousobranchia * Papua New 1294 - Didemnum rubeum * Didemnidae * Aplousobranchia * Palau *
Guinea * Eastern Fields * 100 ft (30 m). This is another stalked species Airai Channel * 33 ft (10 m). This species occurs as encrusting sheets made
1286 - Oxycorynia fascicularis * Papua New Guinea 1287 - Sigillina sp. * Papua New Guinea
277
1288 - Sigillina signifera * Federated States of Micronesia 1289 - Didemnum gutatum * Papua New Guinea
1290 - Didemnum molle * Philippines 1291 - Didemnum mosleyi * Federated States of Micronesia
1292 - Didemnum sp. * Philippines 1293 - Didemnum psammathodes * Papua New Guinea
up of mammilate groups of zooids clustered around a single cloaca. It is usu-
ally inflated with water, resulting in the appearance shown in the photograph,
but it can also retract to form a thin featureless sheet which makes it hard to
recognize as the same ascidian. This species is commonly found on the sides
of clear water channels in Micronesia.
279
1300 - Didemnum sp. * Indonesia 1301 - Didemnum sp. * Federated States of Micronesia
1304 - Diplosoma sp. * Papua New Guinea 1305 - Leptoclinides reticulatus * Papua New Guinea
1308 - Leptoclinides sp. * Didemnidae * Aplousobranchia * Indonesia *
Manado * 125 ft (38 m). This is an encrusting didemnid which occurs on
dead corals and rocky substratum in deep reef environments. Like all didem-
nids it has tiny zooids. The cloacal opening of each zooid empties into a sys-
tem of channels, which combine to form the large cloacal openings seen here.
This colony is inflated. The genus Leptoclinides possesses spicules.
1306 - Leptoclinides sp. * Indonesia 1311 - Lissoclinum patella * Didemnidae * Aplousobranchia * Papua New
Guinea * Madang * Pig Island reef * 10 ft (3 m). This is an ascidian, that
forms large flattened masses with distinct valleys and depressions on its upper
surface. The species varies from near white to dark green depending on the
density of symbiotic algae, often with differences in color between valleys
and ridges on the surface. The photographed individual is living among live
coral, a common locality for this species which likes level reef bottoms with
abundant light. The species is well known and is widely distributed, includ-
ing the Indian Ocean to the central tropical Pacific.
281
1312 - Lissoclinum sp. * Federated States of Micronesia 1313 - Lissoclinum sp. * Papua New Guinea
1314 - Trididemnum cyclops * Papua New Guinea 1315 - Diazona sp. * Indonesia
1323 - Rhopalaea crassa * Diazonidae * Phlebobranchia * Papua New 1331 - Plurella sp. * Plurellidae * Phlebobranchia * Philippines * Cebu *
Guinea * Bagabag Island * fringing reef * 66 ft (20 m). The clear and col- Mactan Island * 66 ft (20 m). This is a second species of Plurella, with a
ored tunics of Rhopalaea are graced with delicate colored lines which differ different shape and color from the previous species. The taxonomy of the
between species. These are among the most lovely of ascidians. This species genus is poorly known, so it is uncertain whether these species are unde-
is found on reef dropoffs at 12 to 100 ft (30 m) depths. scribed and what their geographic ranges might be.
1324 - Rhopalaea sp. * Diazonidae * Phlebobranchia * Indonesia * 1332 - Perophora namei * Perophoridae * Phlebobranchia * Indonesia *
Manado * 33 ft (10 m). This small blue species is close to R. crassa, but Biak Island * 100 ft (30 m). This lovely ascidian is one of a group which
differs in several regards. The genus is known through the tropical Indo- have the tiny zooids on a stem that grows out from the reef face. Needless to
Pacific, with probably several undescribed species similar to R. crassa. say, such ascidians are not immediately obvious and only with careful search-
ing in the right area can they be found.
1325 - Ascidia sp. * Ascidiidae * Phlebobranchia * Papua New Guinea *
New Britain * Kimbe Bay * Restorf Island * 33 ft (10 m). It is common 1333 - Botryllus sp. * Styelidae * Stolidobranchia * Papua New Guinea *
for large solitary ascidians to grow in crevices with only their siphons pro- Port Moresby * Basilisk Passage * 100 ft (30 m). This undescribed species
truding, as is seen in this photograph. Different species have different color of Botryllus occurs in Papua New Guinea and perhaps Indonesia. It varies
markings around the siphons. somewhat in color, but is always encrusting as is shown in the photograph.
1326 - Ascidia sp. * Ascidiidae * Phlebobranchia * Philippines * Santa 1334 - Botryllus sp. * Styelidae * Stolidobranchia * Indonesia * Manado
Rosa * 66 ft (20 m). This solitary Ascidia shows another variation in siphon * 66 ft (20 m). Botryllus contains some of the most colorful of ascidians.
color. Again the ascidian is growing in a crevice with the siphons exposed. Color within the species of the genus is also quite variable. This colony from
1322 - Diazona sp. * Indonesia 1323 - Rhopalaea crassa * Papua New Guinea
283
1324 - Rhopalaea sp. * Indonesia 1325 - Ascidia sp. * Papua New Guinea
285
1336 - Botryllus sp. * Indonesia 1337 - Botryllus sp. * Philippines
1340 - Eusynstyela sp. * Palau 1341 - Polycarpa argentata * Papua New Guinea
90 foot depths. The white and gold colors of the siphons, contrasted with the
dark tunic, make them stand out along a reef wall.
1345 - Polycarpa cryptocarpa * Palau 1346 - Polycarpa papillata * Papua New Guinea
1347 - Polycarpa sp. * Federated States of Micronesia 1348 - Polycarpa sp. * Papua New Guinea
287
1349 - Polycarpa sp. * Papua New Guinea 1350 - Polycarpa sp. * Indonesia
1351 - Polycarpa sp. * Papua New Guinea 1352 - Symplegma viride * Federated States of Micronesia
1353 - Herdmania momus * Indonesia 1354 - Pyura sp. * Papua New Guinea
SUGGESTIONS FOR FURTHER READING
Introduction. Phylum Cnidaria and Ctenophora
Coral Reefs:
Allen, Gerald R. and Roger Steene. 1994. Indo-Pacific Coral Faulkner, D. and R. Chesher. 1979. Living Corals, Clarkson
Reef Field Guide, Tropical Reef Research, Singapore, 378 pp. N. Potter, New York, 311 pp.
Ryan, Paddy. 1994. Snorkleller’s Guide to the Coral Reef. Fautin, Daphne G. and Gerald R. Allen. 1992. Field Guide to
University of Hawaii Press, Honolulu, 184 pp. Anemonefishes and their Host Sea Anemones. Western
Australian Museum Press, 160 pp.
Steene, R. 1990. Coral Reefs Natures Richest Realm, Mallard Veron, J.E.N. 1986. Corals of Australia and the Indo-Pacific,
Press, New York, 336 pp. Angus and Robertson Publ., 644 pp. (reissued in 1992 by
Micronesia: University of Hawaii Press).
Devaney, D.M., E.S. Reese, B.L. Burch and P. Helfrich. 1987, Worms
The Natural History of Enewetak Atoll. U.S. Dept. of Energy Stephen, A.D. and S.J. Edmonds. 1972. The phyla Sipuncula
(DOE/EV/0073-T1), 2 vols. and Echiura. British Museum Natural History, London.
Faulkner, D. 1974, This Living Reef, Quadrangle/The New Fauchaud, K. 1977. The polychaete worms. Natural History
York Times Book Co., New York, 183 pp. Museum of Los Angles County, Science Series 28: 188 pp.
Myers, R.F. 1989. Micronesian Reef Fishes. Coral Graphics, Molluscs
Guam, 299 pp.
While dealing with the fishes of the Micronesia, this Bertsch, H. and S. Johnson. 1981. Hawaiian nudibranchs,
book has an excellent introduction to the area. Oriental Publishing Co, Hawaii, 112 pp.
New Caledonia: Burgess, C.M. 1970. The Living Cowries, A.S. Barnes and
Co., Cranbury, New Jersey, 389 pp.
Laboute, P., M. Fuega and R. Garndperrin. 1991. Le Plus Hinton, Alan. 1972. Shells of New Guinea and the Central
Deau Lagon du Monde, Editions Alizes, Noumea, New Indo-Pacific. The Jacaranda Press, Milton, Qld. Australia, 94
Caledonia, 272 pp. pp.
288 Laboute, P. and Y. Magnier. 1979. Underwater guide to New Pechar, Chris, Chris Prior and Brian Parkinson. no date. Mitre
Caledonia, les Editions du Pacifique, 160 pp. Shells from the Pacific and Indian Oceans. Robert Brown and
The Great Barrier Reef Associates, Australia, no pagination.
Bennett, Isobel. 1988. The Great Barrier Reef, Lansdowne Radwin, G.E. and A. D’Attilio. 1976. Murex Shells of the
Press, Sydney, 184 pp. World. Stanford University Press, 284 pp.
Mather, P. and I. Bennett. 1993. Coral Reef Handbook, A Willan, Richard C. and Neville Coleman. 1984. Nudibranchs
Guide to the Geology, Flora and Fauna of the Great Barrier of Australasia. Australasian Marine Photographic Index,
Reef. Surrey Beatty and Sons, Norton, NSW, Australia, 4th Sydney, Australia
ed. 264 pp. Arthropods- Crustaceans
Reader’s Digest. 1983. Reader’s Digest Book of the Great Debelius, Helmut, 1983. Armoured Knights of the Sea.
Barrier Reef, Reader’s Digest, Sydney, 384 pp. Kernen Verlag, Essen, 120 pp.
Hawaiian Islands Lophophorate Phyla.
Devaney, Dennis M. and Lucius G. Eldredge (eds). Reef and Emig, C.C. 1982. The biology of Phoronida. Adv. Mar. Biol.
Shore Fauna of Hawaii, Section 1. Protozoa through 19:1-89.
Ctenophora; Section 2. Platyhelminthes through Phoronida;
Section 3. Sipncula through Annelida; Section 4. Mollusca. Soule, D.F. and J.D. Soule. 1976. The status of faunistic infor-
Bishop Museum Special Publication 64 (1-4). mation on tropical reef bryozoans. Micronesica 12: 157-164.
Fielding, Ann and Ed Robinson. 1987. An Underwater Guide Echinoderms
to Hawai’i, University of Hawaii Press, Honolulu, 156 pp.
Hobson, Edmund and E.H. Chave. 1990. Hawaiian Reef Guille, Alain, Pierre LaBoute and Jean-Louis Menou. 1986.
Animals, University of Hawaii Press, Honolulu, 137 pp. Guide des etoiles de mer, oursins et autres echinodermes du
lagon de Nouvelle-Caledonie, Editions de l’ORSTOM, Paris,
Hong Kong and South China Sea: 238 pp.
Morton, Brian and John Morton. 1983. The Seashore Ecology Chordates- Ascidians
of Hong Kong, Hong Kong University Press, 350 pp. Monniot, C., F. Monniot and P. Laboute. 1991. Coral Reef
Phylum Porifera Ascidians of New Caledonia, Editions de l’ORSTOM, Paris,
248 pp.
Hooper, J.N.A. and F. Wiedenmayer. 1994. Zoological This book is an excellent source for information
Catalogue of Australia Volume 12 Porifera, CSIRO about the biology of coral reef ascidians.
Melbourne, 624 pp.
INDEX - General