FRONTISPIECE.—Carrie Bow Cay reef complex, aerial photo mosaic, March 1976. 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES -NUMBER 12 

The Atlantic Barrier Reef 

Ecosystem at Carrie Bow Cay, Belize, I 

STRUCTURE AND COMMUNITIES 

Klaus Riitzler 

and Ian G. Macintyre 

EDITORS 

ISSUED 

JUN1019K 

MTHSONIAN PUBLICATIONS 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1982 
ABSTRAC T 

Riitzler, Klaus, and Ian G. Macintyre, editors. The Atlantic Barrier Reef 

Ecosystem at Carrie Bow Cay, Belize, I: Structure and Communities. Smithsonian Contributions to the Marine Sciences, number 12, 539 pages, frontispiece, 

232 figures, 5 plates, 47 tables, 1982.—The results of the first series of 

multidisciplinary investigations of the Caribbean barrier reef complex near 

Carrie Bow Cay, Belize, are reported in 34 papers in this volume, which 

begins with a summary of past work on the Belizean reefs and cays. The first 

section treats the structure of barrier reef habitats in the vicinity of Carrie 

Bow Cay, influential physical parameters such as tides and currents, geological 

and sedimentological history of lagoon, reef, and island substrates, and the 

island's environment, including its climate and the effects of hurricanes. 

Subsequent papers analyze the distribution of endolithic microorganisms in 

carbonate substrates, and the diversity, standing crop, and production in 

selected lagoon and back-reef habitats. Related contributions report on the 

benthos of an unusual submarine cave and on the surface zooplankton over 

reef and lagoon bottoms. One section is devoted to the systematics and local 

distribution of flora and fauna. Marine plants covered are plankton diatoms, 

benthic algae—including a detailed study of the red alga Polysiphonia—and 

sea grasses. Faunistic studies focus on hydroids, medusae, stony corals, octocorals, sipunculans, anthurid isopods, pycnogonids, a marine chironomid, 

ophiuroids, and crinoids. In the papers on Polysiphonia, hydroids, stony corals, 

and anthurids, all species are illustrated for identification by nonspecialists; 

figures of important or unusual examples are shown in the other systematic 

contributions. New species are described among anthurids, pycnogonids, and 

ophiuroids. A section on ecological responses discusses the reaction of algae to 

grazing pressure, the life history of an ichthyo-parasitic hydroid, the growth 

response of the reef coral Montastrea annularis to a light gradient, and associations between zoanthids and their sponge hosts. Included in this section are 

discussions of the ecology of the zoanthid Isaurus duchassaingi, settlement 

behavior and development of the bivalve Malleus candeanus, and behavioral 

ecology of two closely related reef fishes, genus Acanthemblemana. The volume 

concludes with two general surveys of the barrier reef and cays, which discuss 

the Carrie Bow reef section and cay in relation to the overall barrier reef 

complex. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution's annual report, Smithsonian Year. 

Library of Congress Cataloging in Publication Data 

The Atlantic barrier reef ecosystem at Carrie Bow Cay, Belize. 

(Smithsonian contributions to the marine sciences ; no. 12) 

Bibliography: p. 

Contents: 1. Structure and communities. 1. Coral reef ecology—Belize—Carrie Bow Cay. 2. 

Carrie Bow Cay (Belize) I. Riitzler, Klaus. II. Macintyre, Ian G. III. Series. 

QH108.B43A87 574.5'26367'097282 81-607039 AACR2 
Contents 

Page 

FOREWORD ix 

Porter M. Kier 

PREFACE x 

INTRODUCTION 1 

Klaus Riitzler and Ian G. Macintyr e 

Topography, Bio-Geological Structure, 

and Physical Environment 

T H E HABITAT DISTRIBUTION AND COMMUNITY STRUCTURE OF THE BARRIER 

REE F COMPLEX AT CARRIE Bow CAY , BELIZE 9 

Klaus Riitzler and Ian G. Macintyre 

TIDES AT CARRIE Bow CAY , BELIZE 47 

Bjorn Kjerfve, Klaus Riitzler, and George H. Kierspe 

WATER CURRENTS ADJACENT TO CARRIE Bow CAY, BELIZE 53 

Jeffrey E. Greer and Bjorn Kjerfve 

WATER EXCHANGE ACROSS THE REE F CREST AT CARRIE Bow CAY, 

BELIZE 59 

Bjorn Kjerfve 

GEOLOGY AND SEDIMENT ACCUMULATION RATES AT CARRIE Bow CAY, 

BELIZE 63 

Eugene A. Shinn, J. Harold Hudson, Robert B. Halley, Barbar a 

Lidz, Daniel M. Robbin, and Ian G. Macintyre 

TERRESTRIAL ENVIRONMENT AND CLIMATE, CARRIE Bow CAY , BELIZE . . 77 

Klaus Riitzler and Joa n D. Ferraris 

Benthic and Planktonic Communities 

DISTRIBUTION OF MICROBORERS WITHIN PLANTED SUBSTRATES ALONG A 

BARRIER REE F TRANSECT, CARRIE Bow CAY, BELIZE 93 

Jeffrey A. May, Ian G. Macintyre, and Ronald D. Perkins 

PRODUCTION OF SOME BENTHIC COMMUNITIES AT CARRIE Bow CAY, 

BELIZE 109 

Paul E. Hargraves 

MACROBENTHIC INVERTEBRATES IN BARE SAND AND SEAGRASS {Thalassia 

testudinum) AT CARRIE Bow CAY, BELIZE 115 

David K. Young and Marth a W. Young 

A SUBMARINE CAVE NEAR COLUMBUS CAY, BELIZE: A BIZARRE CRYPTIC 

HABITAT 127 

Ian G. Macintyre, Klaus Riitzler, Jame s N. Norris, and Kristian 

Fauchald 

v 
VI SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

SURFACE ZOOPLANKTON AT CARRIE Bow CAY, BELIZE 143 

Joa n D. Ferraris 

Diversity and Distribution of Flora and Fauna 

PLANKTON DIATOMS (BACILLARIOPHYCEAE) FROM CARRIE Bow CAY , 

BELIZE 153 

Paul E. Hargraves 

MARIN E ALGAE AND SEAGRASSES FROM CARRIE Bow CAY , BELIZE 167 

Jame s N. Norris and Katin a E. Bucher 

T H E RE D ALGA Polysiphonia GREVILLE (RHODOMELACEAE) FROM CARRIE 

Bow CA Y AND VICINITY, BELIZE 225 

Donald F. Kaprau n and Jame s N. Norris 

HYDROIDEA (CNIDARIA: HYDROZOA) FROM CARRIE Bow CAY , BELIZE .. . 239 

Barry W. Spracklin 

MEDUSAE (CNIDARIA) FROM CARRIE Bow CAY , BELIZE 253 

Ronald J. Larson 

STONY CORALS (CNIDARIA: HYDROZOA, SCLERACTINIA) OF CARRIE Bow 

CAY , BELIZE 271 

Stephen D. Cairns 

OCTOCORALLIA (CNIDARIA) FROM CARRIE Bow CAY , BELIZE 303 

Katherine Muzik 

DISTRIBUTION OF SIPUNCULA IN THE CORAL REE F COMMUNITY, CARRIE 

Bow CAY , BELIZE 311 

Mar y E. Rice and Ian G. Macintyr e 

ANTHURIDEA (CRUSTACEA: ISOPODA) OF CARRIE Bow CAY , BELIZE 321 

Brian Kensley 

PYCNOGONIDA FROM CARRIE Bow CAY , BELIZE 355 

C. Allan Child 

Pontomyia EDWARDS (DIPTERA: CHIRONOMIDAE), A MEMBER OF THE CORAL 

REE F COMMUNITY AT CARRIE Bow CAY , BELIZE 381 

Gemo t Bretschko 

OPHIUROIDEA (ECHINODERMATA) FROM CARRIE Bow CAY , BELIZE 387 

Frederick H.C. Hotchkiss 

SHALLOW-WATER CRINOIDEA (ECHINODERMATA) FROM CARRIE Bow CAY , 

BELIZE 413 

D. Bradford Macurda , Jr. 

Behavior, Species Interaction, and Responses 

to the Environment 

CHEMICAL DEFENSE IN TROPICAL MARINE ALGAE 417 

Jame s N. Norris and William Fenical 

LIF E HISTORY OF THE HYDROMEDUSA Stomotoca pterophylla HAECKEL AND 

ITS ICHTHYOPARASITIC HYDROID 433 

Ronald J. Larson 
NUMBER 12 vn 

VARIATION IN GROWT H FORMS OF THE REE F CORAL Montastrea annularis 

(ELLIS AND SOLANDER): A QUANTITATIVE EVALUATION OF GROWT H 

RESPONSE TO LIGHT DISTRIBUTION USING COMPUTER SIMULATION ... . 441 

Richard R. Graus and Ian G. Macintyr e 

SPONGE-ZOANTHID ASSOCIATIONS: FUNCTIONAL INTERACTIONS 465 

Sar a M. Lewis 

O N THE ECOLOGY OF Isaurus duchassaingi (ANDRES) (CNIDARIA: ZOANTHIDEA) FROM SOUTH WATER CAY , BELIZE 475 

Kathleen S. Larson and Ronald J. Larson 

LARVAL SETTLEMENT BEHAVIOR AND SHELL MORPHOLOGY OF Malleus 

candeanus (d'ORBiGNY)(MoLLUscA: BIVALVIA) 489 

Thoma s R. Waller and Ian G. Macintyr e 

HABITAT AND RESOURCE PARTITIONING BETWEEN Tw o SPECIES OF Acanthemblemaria (PISCES: CHAENOPSIDAE) , WITH COMMENTS ON THE CHAOS 

HYPOTHESIS 499 

David W. Greenfield and Theresa A. Greenfield 

Carrie Bow Cay as Part of the Belizean 

Barrier Reef Complex 

RECONNAISSANCE STUDY OF THE GEOMORPHOLOGY AND BENTHIC COM -

MUNITIES OF THE OUTER BARRIER REE F PLATFORM, BELIZE 509 

Randolp h B. Burke 

SPECIES-AREA RELATIONSHIPS ON SMALL ISLANDS: FLORISTIC DATA FROM 

BELIZEAN SAND CAYS 527 

David R. Stoddart and F. Raymon d Fosberg 

PLATES 259 
Dedication 

This volume is dedicated to the 50th anniversary 

of the Great Barrier Reef Expedition 1928-1929, 

which pioneered the integration of many science disciplines 

for the better understanding of the coral reef system 
Foreword 

Museum scientists tend to be specialists in a particular discipline and to 

work alone. Those of us, however, who study living organisms learn sooner or 

later that we cannot hope to understand our animals and plants fully without 

some knowledge of the environment in which they exist. We also learn at 

some point to appreciate the benefits of collaborating with fellow scientists 

whose disciplinary focus may differ from ours, but whose interests are related 

to our own through the "environmental" link. That is to say, a great natural 

ecosystem such as the tropical coral reef draws together researchers of many 

diverse disciplines. 

The project "Investigations of Marine Shallow-Water Ecosystems" 

(IMSWE) off Carrie Bow Cay has done this very thing for many of us at the 

Smithsonian Institution and elsewhere, and thus we have learned far more 

about our organisms and their environment than we might have done 

otherwise. As a result, we are more than enthusiastic about IMSWE's progress. 

We started with only a few investigators, but as the years have passed we have 

grown into a multidisciplinary contingent. 

Needless to say, part of IMSWE's success stems from the efforts of the 

principal investigator, Klaus Riitzler. He is responsible for the organizing, 

scheduling, coordinating, and orchestrating. He has done all this with good 

humor and energy. 

Of course, such an effort would not have been possible without the support 

and encouragement of the Belizeans themselves. They love their barrier reef 

and have high regard for its economic, recreational, and aesthetic value. 

Recognizing the need for understanding and protecting this precious resource, 

Belizean officials have approved and assisted our various endeavors since 

IMSWE's modest beginnings. In particular, Winston Miller, Fisheries Officer, 

has helped us in every way, as have members of the staff at the Ministry of 

Trade and Industry (after 1979 the Ministry of Health, Housing, and Cooperatives), and the Ministry of Finance. 

We all know that research cannot go forward without adequate funding. 

The Exxon Corporation has provided part of our support, and has done it in 

a most generous fashion, allowing the scientists to go about their studies 

independently. 

This volume thus represents the culmination of research effort and support 

from many quarters. It is the first of an open report series and serves as the 

basis for future contributions. It not only "sets the scene" by determining 

terminology and summarizing our knowledge to date, but it also points out 

the gaps yet to be closed. We look forward to the next decade. 

September 1980 Porter M. Kier, Director (1970-1979) 

National Museum of Natural History 

Smithsonian Institution 

ix 
Preface 

On a windy, overcast morning of February 1972, we discovered Carrie Bow 

Cay. Arnfried Antonius and I (K.R.) were on our way from Stann Creek (now 

Dangriga) to Glover's Reef, looking for the Tobacco Cay passage through the 

barrier reef. We could already hear the waves pounding the coral crest but 

neither of two islands in front of us matched our memory of Tobacco Cay, 

and the crew of the charter boat from Belize City, unfamiliar with southern 

waters, was uncertain too. The smaller but nearer one of the cays had 

buildings and a solid dock so we decided to have a closer look. Minutes later 

we tied up to the concrete pier and walked toward the stately main building. 

No one was around, except for a few mildly disturbed pelicans, but a 

weathered sign above the gate to the main porch said "Welcome to Carrie 

Bow Island." On that memorable day when we first walked around Carrie 

Bow—a speck of sand covered by about 80 coconut trees overlooking three 

cottages, two outhouses, and three water tanks—we could not have predicted 

the developments of the eight years to come. Despite its small size, this island 

has been the site of a simple but functional laboratory to which more than 70 

scientists from 30 institutions have come to study the well-developed reef 

complex nearby. 

Records indicate that up to 1927 Carrie Bow Cay, then known as Ellen 

Caye or Bird Caye, was an uninhabited sanctuary for migrating birds and for 

sea turtles coming there to lay their eggs. The surrounding reef "abounded 

with conch, lobster, turtles, and parrot fish, all of which have been depleted 

by extensive trade in these commodities," according to the historical account 

of Henry T. A. Bowman, current owner of Carrie Bow Cay. Mr. Bowman 

bought the island in 1943 and decided to name it after his wife Carrie. "At 

that time the cay was about twice the size it is now, and was surrounded by 

mangrove, and had about 75 coconut trees that were planted in the early 

1900's. The mangrove, breeding spot for mosquitos, was cleared off in 1944 

when [he] decided to build a summer home. Of the original coconut trees 

only six remain today." Mr. Bowman's account also cites the stresses introduced by countless storms and hurricanes, which were particularly harsh 

during the past three decades but whose damage Carrie Bow Cay and the reef 

have managed to survive. 

Notwithstanding these changes, the area remains relatively undisturbed by 

scientific standards, and thus is an ideal location for an ecological study of 

coral reefs. More significant, it is a segment of an enormous reef complex that 

is reasonably accessible from major cities on the North American mainland. 

This volume presents the first extensive scientific account of the barrier reef 

around Carrie Bow Cay, Belize. It is an outgrowth of the Smithsonian 

Institution Program "Investigations of Marine Shallow-Water Ecosystems" 

(IMSWE), which began in 1970 under the guidance of W. H. Adey, A. L. 
NUMBER 12 

Dahl, and the editors of this volume. It was our goal to introduce long-term 

cooperation and a truly interdisciplinary approach to the ecological study of 

coral reefs and related tropical environments. Our search for a study site that 

would be satisfactory to the special requirements of various disciplines led us 

to the reef off Belize. 

IMSWE evolved alongside a larger proposed project for the study of a 

coral-reef ecosystem that was to be a multi-institutional and multi-national 

program sponsored by the International Decade of Ocean Exploration. 

Glover's Reef atoll was contemplated as a site for that program's comparative 

studies and Carrie Bow Cay—although not yet seen by any of the planning 

committee—was discussed as an additional possibility because of its logistical 

advantages, particularly its location half way between the mainland of Belize 

and the atoll. Although the large program was not funded, Smithsonian 

Institution scientists carried forward the idea for an interdisciplinary study of 

a coral reef ecosystem. 

Since that time, many enthusiasts have joined in IMSWE's collaborative 

investigations. Because of the growing body of information resulting from our 

work, we are now able to present this volume, whose purpose is to give a 

general introduction to the topography, oceanography, geology, and biology 

of the Belizean barrier reef complex near Carrie Bow Cay. 

Some new terms have been introduced in this volume to describe the 

physiographic characteristics and zonation of the reef in this area. It is hoped 
X ll SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

that even if these terms are revised at a later date, they will serve as an 

important reference for future research reports. This contribution also opens 

a publication series that makes it possible to add information on this reef 

complex as it becomes available and to keep the results in focus for subsequent 

additions and for a later synthesizing monograph. 

The use of brand names in this publication is for descriptive purposes only 

and does not constitute endorsement by the Smithsonian Institution. 

ACKNOWLEDGMENTS.—Our program and the production of this volume has 

received, over the years, help and encouragement from so many supporters in 

direct and indirect ways that it is impossible to give a just account of them all. 

Those who have contributed specifically to papers published in the following 

pages are acknowledged in the article introductions. Here, we wish to highlight 

the accomplishments of individuals and groups whose efforts have made our 

program successful and therefore made possible the production of this publication. 

The Exxon Corporation extended grant support to us over the past five 

years; we are grateful not only for the financial aid but also for the personal 

interest in our studies shown by R. E. Chandler and E. W. Markowski, Public 

Affairs Department. Other funds came from the Smithsonian Environmental 

Sciences Program, National Museum of Natural History, Research Award 

Program and Fluid Research Fund, for which we thank D. Challinor, P. M. 

Kier, J. F. Mello, and S. D. Ripley. Special donations from AMOCO, Cities 

Service, Getty, Gulf, and Union Oil companies, and from K. Sandved and L. 

Taylor made possible the printing of the color plates in this volume. 

The people of Belize greatly enriched our days in the field, and its 
NUMBER 12 Xlll 

government officials helped us in many ways. W. Miller, Fisheries Officer and 

our scientific sponsor, patiently helped with certifications and permits. The 

Ministry of Trade and Industry (after 1979 the Ministry of Health, Housing, 

and Cooperatives) sanctioned our work, the Ministry of Finance granted 

exemption of scientific equipment from duty. H.T.A. Bowman ("Sir Henry") 

and Mrs. Bowman were our generous hosts for the past eight years. At the 

same time, H.T.A. Bowman, Jr. ("Junior") worked hard as our agent and 

guide, assisted by his wife Alice, sister Norma, and by other members of his 

family. H.T.A. Bowman III ("Henry") has read the tide gauge installed by us 

at Pelican Beach Motel, Dangriga, for the past four years. 

For many years the single most important person for the program was M. 

R. Carpenter, who solved logistical problems at home and in the field. Mike 

was preceded, or at times relieved, as field manager by H. Adolfi, A. Antonius, 

R. J. Larson, A. B. Rath, R. H. Sims, and B. Spracklin. Others who 

contributed to our comfort on the cay include people known familiarly to us 

as Frank and Japs, staff members of Pelican Beach Motel, and Emily, Ernie, 

and Genevieve, our cooks. Invaluable logistical help was also provided by C. 

Moore of Sen & Co., Belize City. At the National Museum of Natural History 

our study benefited greatly from the administrative expertise of C. A. Ossola, 

M. Parrish, and M. R. Tanner and from the technical assistance of C. G. 

Ahearn, W. T. Boykins, A. C. Cohen, L. G. Cole, R. J. and K. S. Larson, and 

K. B. Sandved. J. Petroski, Travel Services Office, did miracles with airplane 

reservations. 

This volume is the product of our collaboration with a number of patient 

authors and several dedicated assistants and goodwilled advisors. Presentation 

was shaped at an early stage by the skillful editing of V. V. Macintyre and 

the proficient illustrating of I. F. Jewett. M. Parrish and A. Stonework typed 

or retyped most of the contributions. S. D. Cairns, M. R. Carpenter, M. 

Parrish, and D. S. Robertson helped as editorial assistants. C. Schbpfer-Sterrer 

created the accompanying ink sketches and G. J. Thomas took the aerial 

photographs for the frontispiece. The following colleagues helped us as 

reviewers or scientific advisors: W. H. Adey, F. M. Bayer, T. E. Bowman, M. 
x i v SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

A. Buzas, S. D. Cairns, D. R. Calder, F. A. Chace, B. B. Collette, M. E. 

Downey, G. L. Hendler, R. S. Houbrick, E. Kirsteuer, C. G. Messing, J. N. 

Norris, D. L. Pawson, H. Pulpan, J. Rosewater, and T. Waller. Finally, special 

thanks are due to A. L. Ruffin, managing editor, B. J. Spann, editor, and L. 

J. Long, production manager, of the Smithsonian Institution Press, for seeing 

this publication through the final stages of production. 
The Atlantic Barrier Reef 

Ecosystem at Carrie Bow Cay, Belize, I 

Introduction 

Klaus Riitzler 

and Ian G. Macintyre 

The barrier reef off Belize, Central America, 

has received the concentrated attention of scientists only within the past two decades. Interest in 

this area dates back to the late nineteenth century, however, the first scientific reference to the 

Belizean reefs having been made by Charles Darwin, himself, in an 1842 work, 77?^ Structure and 

Distribution of Coral Reefs (pages 201-202). Using 

information provided by Captain B. Allen, Darwin discussed the bathymetry associated with 

Belizean reefs and included them in his classification of principal reef types. 

Early studies of the natural history of Belize, 

then known as British Honduras, focused not on 

the reefs but on incidental collections taken along 

the shore or by dredge hauls by residents of the 

colony and by visiting individuals and expeditions, most notably the "Pawnee" (1925) and 

"Rosaura" (1937-1938) expeditions. One of the 

oldest such records is the description of a new 

sponge, collected near Belize (now Belize City) by 

Priest (1881). Other systematic work on early 

collections dealt with algae (Taylor, 1935), seagrass Thalassia (den Hartog, 1970), sponges (Burton, 1954), cnidarians (Boone, 1928a), crustaKlaus Riitzler, Department of Invertebrate Zoology, and Ian G. 

Macintyre, Department of Paleobiology, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C. 20560. 

ceans (Boone, 1927), mollusks (Boone, 1928b; 

Marcus and Marcus, 1962; Robertson, 1975), 

echinoderms (Boone, 1928c; John and Clark, 

1954), and fishes (Breder, 1927; Tucker, 1954; 

Robins and Starck, 1965; Birdsong and Emery, 

1968). In addition, studies relating to commercial 

fisheries were reported by Smith (1939, 1941) and 

Craig (1966). 

The first investigations focusing on the cays 

and reefs of Belize were undertaken by the "Cambridge Expedition to British Honduras" (1959-

1960) led by J. E. Thorpe (Thorpe and Stoddart, 

1962). Some members of the expedition mapped 

40 cays, including Carrie Bow Cay, and collected 

samples of their flora. They also discussed the 

origin, formation, and distribution of reef and 

mangrove cays, and showed the migration of 

some of these cays on the basis of exposed beach 

rock (Stoddart, 1960). Another expedition team 

mapped bottom characteristics and identified 

coral species from around Rendezvous Cay, and 

conducted experiments on coral behavior in response to external stimuli (Thorpe and Bregazzi, 

1960). Subsequently Stoddart, one of this expedition's participants, surveyed in detail the biology and geology of the Belizean atolls, that is, 

Turneffe Islands, Lighthouse Reef, and Glover's 

Reef (Stoddart, 1962a). At this time he also initiated an interesting series of studies concerning 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

the hurricane destruction and post-storm recovery of cays and coral reefs (Stoddart, 1962b, 1963, 

1969, 1974). 

A second stage of intensive research activity 

was spearheaded by E. G. Purdy of Rice University, under whom several major projects—for the 

most part doctoral disserations—treating various 

geological aspects of the barrier reef complex were 

completed between 1961 and 1967. Most of this 

work appeared in a volume edited by Wantland 

and Pusey (1975) that focused on sediments and 

processes of sedimentation and diagenesis (L. R. 

High, Jr., pages 53-96; M. R. Scott, pages 97-

130; W. C. Pusey III; pages 131-233; W. J. 

Ebanks, Jr., pages 234-296), paleoecology and 

diagenesis of Pleistocene limestone (G. E. Tebbutt, pages 297-331), and distribution of benthic 

Foraminifera (K. F. Wantland, pages 332-339) 

and Ostracoda (J. W. Teeter, pages 400-499). 

Another member of this group of doctoral 

scholars documented elsewhere the genesis of recent lime mud (Matthews, 1966). In addition, 

Purdy himself produced two thought provoking 

reports dealing with the origin of the topographic 

relief of the Belizean reef and its influence on 

sediment distribution (Purdy, 1974a, 1974b). 

Among other studies of the Belizean reefs some 

of the most notable have been conducted by the 

University of Miami under the direction of R. N. 

Ginsburg. This work, to date, has focused on 

shallow-water submarine cements (Ginsburg and 

James, 1976; James et al., 1976) and has included 

geological investigations (James and Ginsburg, 

1978) and a study of deep-reef fishes (Colin, 1974) 

of the deep seaward margins of the barrier reef 

and Glover's Reef atoll. 

Several recent studies have concentrated on the 

barrier reef and the atolls. Devaney (1974) has 

reported on shallow-water echinoderms collected 

off Turneffe Island and Lighthouse and Glover's 

reefs. Wallace and Schafersman (1977) have examined the ecology and sedimentology of patch 

reefs in the Glover's Reef lagoon. This atoll was 

also the point of origin for a series of ichthyological studies that were expanded to the barrier 

reef complex and are still under way (Greenfield, 

1972, 1975a-c, 1979; Greenfield and Greenfield, 

1973; Greenfield and Johnson, 1981). 

With the establishment of the Smithsonian 

Institution's field laboratory at Carrie Bow Cay 

in 1972, most of the marine research in Belize 

concentrated on this part of the barrier reef; only 

some early site surveys and comparative studies 

included other locations. Smithsonian-sponsored 

physiographic surveys (Dahl et al., 1974; Miller 

and Macintyre, 1977) were important early steps 

of our programs, while hurricane reports and 

damage and recovery surveys (Antonius, 1972; 

Stoddart, 1974; Highsmith et al., 1980) appear to 

have become a continuing necessity. Among the 

methods developed or modified during our reef 

research were mapping (Riitzler, 1978a), underwater coring (Macintyre, 1975), and ecological 

sampling techniques (Dahl, 1973; Kirsteuer, 

1978; Riitzler, 1978b; Riitzler et al., 1980). 

Early in our program, attempts were made to 

monitor the physical environment of Carrie Bow 

Cay and the surrounding waters. Cooperation 

with colleagues at the University of South Carolina who contributed specialized equipment produced information on the diurnal energy balance 

on the island (Kjerfve, 1978) and on tidal patterns 

relative to the Caribbean system (Kjerfve, 1981). 

Inventory of flora and fauna was a prerequisite 

for the new program and produced an increasing 

number of checklists, distributional tabulations, 

taxonomic revisions, and descriptions of new 

taxa: algae and seagrasses (Tsuda and Dawes, 

1974); nemerteans (Kirsteuer, 1974, 1977); polychaetes (Fauchald, 1980); sipunculans (Rice, 

1976); copepods (H. B. Cressey, 1981; R. Cressey, 

1981); ostracods (Kornicker, 1978, 1981; Kornicker and Cohen, 1978); decapods (Kensley, 

1981; Kensley and Gore, 1981), bivalves (Waller, 

1978), holothuroids (Pawson, 1976); echinoids 

(Kier, 1975); and fishes (Greenfield, 1981; Greenfield and Johnson, 1981; Johnson and Greenfield, 

1982). A comparable inventory of novel chemical 

compounds from algae, sponges, and gorgonians 

is the subject of a series of publications of W. 

Fenical's group at Scripps Institution of Oceanography (McEnroe and Fenical, 1978; Mc-
NUMBER 12 

Connell and Fenical, 1978; Kokke et al., 1979; 

Kokkeet al., 1981). 

Participants of several programs have examined organisms producing limestone and calcareous sand. Environmental influences on skeletal 

development of corals were studied by Barnard 

et al. (1974), Macintyre and Smith (1974), Graus 

and Macintyre (1976), and Highsmith (1979). 

Related work on mollusks has dealt with growth 

rates of gastropods {Cerithium: Houbrick, 1974) 

and shell calcification of bivalves (Arcoida: 

Waller, 1980). Calcium carbonate breakdown by 

biological processes, on the other hand, was the 

subject of studies by Riitzler (1975), Rice (1976), 

and Highsmith (1981), and the contribution of 

noncalcareous (siliceous) components to reef 

sands was treated by Riitzler and Macintyre 

(1978). 

Another topic of research at Carrie Bow Cay 

has been the availability and quality of food as a 

major factor determining growth and distribution 

of invertebrates and fishes. Primary production 

by benthic algae, including spatial and temporal 

variability, were discussed by Dahl (1974, 1976). 

Distribution of many algae is controlled by herbivores (Hay, 1981) but some plants can defend 

themselves from grazers by producing toxic compounds that act as feeding deterrents (Gerwick et 

al., 1979; Sun and Fenical, 1979; Paul and Fenical, 1980; Gerwick and Fenical, 1981). On the 

other hand, strong symbiotic ties exist between 

certain algae and invertebrates that were studied 

by Kokke et al. (1980) and Riitzler (1981). Another report on invertebrate feeding presents behavioral observations on coronate medusae captured in plankton tows near Carrie Bow Cay 

(Larson, 1979). 

The earlier work summarized here will give the 

reader an indication of research projects under 

way at the Carrie Bow Cay laboratory or sponsored by the Smithsonian reef study program 

elsewhere in Belize. The following papers in this 

volume will add further data and important new 

perspectives to our knowledge. Many more contributions can be expected to follow. 

Literature Cited 

Antonius. 

1972. 

Barnard, 

1974. 

Birdsong, 

1968. 

Boone, L 

1927. 

Hurricane Laura, Witnessed in British Honduras. 

Atoll Research Bulletin, 162:11-12. 

L. A., I. G. Macintyre, and J. W. Pierce 

Possible Environmental Index in Tropical Reef 

Corals. Nature, 252:219-220. 

R. S., and A. R. Emery 

New Records of Fishes for the Western Caribbean. 

Quarterly Journal of the Florida Academy of Sciences, 30: 

187-196. 

Scientific Results of the First Oceanographic Expedition of the "Pawnee," 1925: Crustacea from 

Tropical East American Seas. Bulletin of the 

Bingham Oceanographic Collection, 1(2): 1-147. 

1928a. Scientific Results of the First Oceanographic Expedition of the "Pawnee," 1925: Molusca from 

Tropical East American Seas. Bulletin of the 

Bingham Oceanographic Collection, 1(3): 1-20. 

1928b. Scientific Results of the First Oceanographic Expedition of the "Pawnee," 1925: Echinodermata 

from Tropical East American Seas. Bulletin of the 

Bingham Oceanographic Collection, 1(4): 1-22. 

1928c. Scientific Results of the First Oceanographic Expedition of the "Pawnee," 1925: Coelenterata 

from Tropical East American Seas. Bulletin of the 

Bingham Oceanographic Collection, 1(5): 1-8. 

Breder, C. M., Jr. 

1927. Scientific Results of the First Oceanographic Expedition of the "Pawnee," 1925: Fishes from Tropical East American Seas. Bulletin of the Bingham 

Oceanographic Collection, 1(1): 1-90. 

Burton, M. 

1954. The "Rosaura" Expedition, 1937-1938: Sponges. 

Bulletin of the British Museum (Natural History), Zoology, 2:215-239. 

Colin, P. L. 

Observations and Collection of Deep-Reef Fishes 

off the Coasts of Jamaica and British Honduras 

(Belize). Marine Biology, 24:29-38. 

K. 

Geography of Fishing in British Honduras and Adjacent 

1974. 

Craig, A 

1966. 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Coastal Waters. 143 pages. Baton Rouge, Louisiana: 

Louisiana State University Press. 

Cressey, H. B. 

1981. Ceratocolax mykternastes, New Species (Copepoda, 

Bomolochidae) Parasitic in the Nasal Sinus of 

Haemulon sciurus (Pisces, Pomadasyidae) from Belize. Proceedings of the Biological Society of Washington, 

94(2) :514-524. 

Cressey, R. 

1981. Parasitic Copepods from the Gulf of Mexico and 

Caribbean Sea, I: Holobomolochus and Neobomolochus. Smithsonian Contributions to Zoology, 339: 24 

pages. 

Dahl, A. L. 

1973. Surface Area in Ecological Analysis: Quantification of Benthic Coral-Reef Algae. Marine Biology, 

23:239-249. 

1974. The Structure and Dynamics of Benthic Algae in 

the Coral Reef Ecosystem. In A. M. Cameron, B. 

M. Campbell, A. B. Cribb, R. Endean, J. S. Jell, 

O. A.Jones, P. Mather, and F. H. Talbot, editors, 

Proceedings of the Second International Coral Reef Symposium, 1:21-25. Brisbane, Australia: The Great 

Barrier Reef Committee. 

1976. Generation of Photosynthetic Surface Area by 

Coral Reef Algae. Micronesica, 12:43-47. 

Dahl, A. L., I. G. Macintyre, and A. Antonius 

1974. A Comparative Survey of Coral Reef Research 

Sites. In M.-H. Sachet and A. L. Dahl, editors, 

Comparative Investigations of Tropical Reef Ecosystems: Background for an Integrated Coral Reef 

Program. Atoll Research Bulletin, 172:37-120. 

Darwin, C. 

1842. The Structure and Distribution of Coral Reefs. 214 pages. 

London, 

den Hartog. See Hartog, C. den 

Devaney, D. M. 

1974. Shallow-Water Echinoderms from British Honduras, with a Description of a New Species of 

Ophiocoma (Ophiuroidea). Bulletin of Marine Science, 

24(1): 122-164. 

Fauchald, K. 

1980. Onuphidae (Polychaeta) from Belize, Central 

America, with Notes on Related Taxa. Proceedings 

of the Biological Society of Washington, 93(3): 797-829. 

Gerwick, W. H., and W. Fenical 

1981. Ichthyotoxic and Cytotoxic Metabolites of the 

Tropical Brown Alga, Slypopodium zonate. Journal of 

Organic Chemistry, 46:22-27. 

Gerwick, W. H., W. Fenical, N. Fritsch, and J. Clardy 

1979. Stypotriol and Stypoldione: Ichthyotoxins of 

Mixed Biogenesis from the Mrine Alga Slypopodium 

zonale. Tetrahedron Letters, 1979(2): 145-148. 

Ginsburg, R. N., and N. P. James 

1976. Submarine Botryoidal Aragonite in Holocene 

Reef Limestones, Belize. Geology, 4:431-436. 

Graus, R. R., and I. G. Macintyre 

1976. Light Control of Growth Form in Colonial Reef 

Corals: Computer Simulation. Science, 193:895-

897. 

Greenfield, D. W. 

1972. Notes on the Biology of the Arrow Blenny, Lucayablennius zingaro (Bdhlke) from British Honduras. 

Copeia, 1972:590-592. 

1975a. Emblemariopsis pricei, a New Species of Chaenopsid 

Blenny from Belize. Copeia, 1975:713-715. 

1975b. Cleplicus parrae, an Additional Sponge-Dwelling 

Fish. Copeia, 1975:381-382. 

1975c. Centropomus poeyi from Belize, with a Key to the 

Western Atlantic Species of Centropomus. Copeia, 

1975:582-583. 

1979. A Review of the Western Atlantic Starksia ocellataComplex (Pisces: Clinidae) with the Description 

of Two New Species and Proposal of Superspecies 

Status. Fieldiana: Zoology, 73:9-48. 

1981. Vancus imswe, a New Gobiid Fish from Belize. 

Copeia, 1981:269-272. 

Greenfield, D. W., and T. Greenfield 

1973. Triathalassothia gloverensis, a New Species of Toadfish from Belize (= British Honduras), with Remarks on the Genus. Copeia, 1973:560-565. 

Greenfield, D. W., and R. K. Johnson 

1981. The Blennioid Fishes from Belize and Honduras, 

Central America, with Comments on their Systematics, Ecology, and Distribution (Blenniidae, 

Chaenopsidae, Labrisomidae, Tripterygiidae). 

Fieldiana: Zoology, new series, 8:1-106. 

Hartog, C. den 

1970. Sea-Grasses of the World. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde, 59:1-275. 

Hay, M. E. 

1981. Spatial Patterns of Grazing Intensity on a Caribbean Barrier Reef: Herbivory and Algal Distribution. Aquatic Botany, 11:97-109. 

Highsmith, R. C. 

1979. Coral Growth Rates and Environmental Control 

of Density Banding. Journal of Experimental Marine 

Biology and Ecology, 37:105-125. 

1981. Coral Bioerosion: Damage Relative to Skeletal 

Density. American Naturalist, 117:193-198. 

Highsmith, R. C , A. C. Riggs, and C. M. D'Antonio 

1980. Survival of Hurricane-Generated Coral Fragments 

and a Disturbance Model of Reef Calcification/ 

Growth Rate. Oecologia (Berlin), 46:322-329. 

Houbrick, \J] R-

1974. Growth Studies on the Genus Cerithium (Gastropoda: Prosobranchia) with Notes on Ecology and 

Microhabitats. The Nautilus, 88:14-27. 
NUMBER 12 

James, N. P., and R. N. Ginsburg 

1978. The Deep Seaward Margin of Belize Barrier and Atoll 

Reefs. 201 pages. Miami Beach, Florida: University 

of Miami. 

James, N. P., R. N. Ginsburg, D. S. Marszalek, and P. W. 

Choquette 

1976. Facies and Fabric Specificity of Early Subsea Cements in Shallow Belize (British Honduras) Reefs. 

Journal of Sedimentary Petrology, 46(3):523-544. 

John, D. D., and A. M. Clark 

1954. The Echinodermata. In The "Rosaura" Expedition, 1937-1938. Bulletin of the British Museum [Natural History), Zoology, 2:139-162. 

Johnson, R. K., and D. W. Greenfield 

In press. Cling Fishes from Belize and Honduras, Central 

America, with a Redescription of Gobiesox barbalulus Starck. Northeast Gulf Science. 

Kensley, B. F. 

1981. Notes on Axiopsis (Axiopsis) serratifrons (A. Milne 

Edwards) (Crustacea: Decapoda: Thalassinidea). 

Proceedings of the Biological Society of Washington, 

93(4): 1253-1263. 

Kensley, B. F., and R. H. Gore 

1981. Coralaxius abelei, New Genus and New Species 

(Crustacea: Decapoda: Thalassinidea: Axiidae): 

A Coral-inhabiting Shrimp from the Florida Keys 

and the Western Caribbean Sea. Proceedings of the 

Biological Society of Washington, 93(4): 1277-1294. 

Kier, P. M. 

1975. The Echinoids of Carrie Bow Cay, Belize. Smithsonian Contributions to Zoology, 206: 45 pages. 

Kirsteuer, E. 

1974. Description of Poseidonemertes caribensis sp. n., and 

Discussion of Other Taxa of Hoplonemertini 

Monostilifera with Divided Longitudinal Musculature in the Body Wall. Zoologica Scripta, 3:153-

166. 

1977. Remarks on Taxonomy and Geographical Distribution of the Genus Ototyphlonemertes Diesing 

(Nemertina, Monostilifera). Mikrofauna Meeresboden, 61:167-181. 

1978. Coral-Associated Nemertina. In D. R. Stoddart 

and R. E. Johannes, editors, Coral Reefs: Research Methods. Monographs on Oceanographic Methodology, 5:315-327. Paris: Unesco. 

Kjerfve, B. 

1978. Diurnal Energy Balance of a Caribbean Barrier 

Reef Environment. Bulletin of Marine Science, 28: 

137-145. 

1981. Tides of the Caribbean Sea. Journal of Geophysical 

Research, 86:4243-4247. 

Kokke, W.C.M.C, L. Bohlin, W. Fenical, and C. Djerassi 

In press. Minor and Trace Sterols of Marine Invertebrates, XX: New Methyl Sterols from the Caribbean Gorgonians Briareum asbestinum and Muriceopsis flavida. Journal of Steroids. 

Kokke, W.C.M.C, W.Fenical, L. Bohlin, and C. Djerassi 

1980. Sterol Synthesis by Cultured Zooxanthellae: Implications Concerning Sterol Metabolism in HostSymbiont Association in Caribbean Gorgonians. 

Comparative Biochemistry and Physiology, 68B:281-

287. 

Kokke, W.C.M.C, C. S. Pak, W. Fenical, and C Djerassi 

1979. Minor and Trace Sterols in Marine Invertebrates, 

XII: Occurrence of 24(#+.S)-Isopropenylcholesterol, 24(/c-hS)-MethylchoIesta-5,25-dien3/?-ol, and 24(/c-hS)-Methylcholesta-7,25-dien3/?-ol in the Caribbean Sponge, Verongia cauliformis. Helvetica Chimica Ada, 62(4): 1310-1318. 

Kornicker, L. S. 

1978. Harbansus, a New Genus of Marine Ostracoda, and 

a Revision of the Philomedidae (Myodocopina). 

Smithsonian Contributions to Zoology, 260: 75 pages. 

1981. Revision, Distribution, Ecology and Ontogeny of 

the Ostracode Subfamily Cyclasteropinae (Myodocopina: Cylindroleberididae). Smithsonian Contributions to Zoology, 319: 548 pages. 

Kornicker, L. S., and A. C Cohen 

1978. Dantyinae, a New Subfamily of Ostracoda (Myodocopina: Sarsiellidae). Proceedings of the Biological 

Society of Washington, 91(2):490-508. 

Larson, R. J. 

1979. Feeding in Coronate Medusae (Class Scyphoza, 

Order Coronatae). Marine Behavior and Physiology, 

6:123-129. 

Macintyre, I. G. 

1975. A Diver-operated Hydraulic Drill for Coring Submerged Substrates. Atoll Research Bulletin, 165:21-

26. 

Macintyre, I. C , and S. V. Smith 

1974. X-Radiographic Studies of Skeletal Development 

in Coral Colonies. In A. M. Cameron, B. M. 

Campbell, A. B. Cribb, J. Endean, J. S. Jell, O. 

A. Jones, P. Mather and F. H. Talbot, editors, 

Proceedings of the Second International Coral Reef Symposium, 2:277-287. Brisbane, Australia: The Great 

Barrier Reef Committee. 

Marcus, E., and E. Marcus 

1962. Opisthobranchs from Florida and the Virgin Islands. Bulletin of Marine Science of the Gulf and Caribbean, 12:450-488. 

Matthews, R. K. 

1966. Genesis of Recent Lime Mud in Southern British 

Honduras. Journal of Sedimentary Petrology, 36:428-

454. 

McConnell, O. J., and W. Fenical 

1978. Ochtodene and Ochtodiol: Novel Polyhalogenated Cyclic Monoterpenes from the Red Seaweed 

Ochlodes secundiramea. Journal of Organic Chemistry, 

43:4238-4241. 
6 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

McEnroe, F. J., and W. Fenical 

1978. Structures and Synthesis of Some New Antibacterial Sesquiterpenoids from the Gorgonian Coral 

Pseudopterogorgia rigida. Tetrahedron Letters, 1978(34): 

1661-1664. 

Miller, J. A., and I. G. Macintyre 

1977. Third International Symposium on Coral Reefs, Field 

Guidebook to the Reefs of Belize. 36 pages. Miami 

Beach, Florida: The Atlantic Reef Committee. 

Paul, V., and W. Fenical 

1980. Toxic Acetylene Containing Lipids from the Red 

Alga Liagura farinosa Lamaroux. Tetrahedon Letters, 

1980(21):3327-3330. 

Pawson, D. L. 

1976. Shallow-Water Sea Cucumbers (Echinodermata: 

Holothuroidea) from Carrie Bow Cay, Belize. Proceedings of the Biological Society of Washington, 89(31): 

369-382. 

Priest, B. W. 

1881. On an Undescribed Species of Sponge of the Genus Polymastia, from Honduras. Journal of the Quekett 

Microscopical Club, 6:302-304. 

Purdy, E. G. 

1974a. Reef Configurations: Cause and Effect. In L. F. 

Laporte, editor, Reefs in Time and Space. Society 

of Economic Paleontolgists and Mineralogists, Special 

Publication, 18:9-76. Tulsa, Oklahoma. 

1974b. Karst-Determined Facies Patterns in British Honduras: Holocene Carbonate Sedimentation 

Model. The American Association of Petroleum Geologists Bulletin, 58:825-855. 

Rice, M. E. 

1976. Sipunculans Associated with Coral Communities. 

Micronesica, 12:119-132. 

Robertson, R. R. 

1975. Systematic List of Commonly Occurring Marine 

Mollusks of Belize. In K. F. Wantland and W. C. 

Pusey III, editors, Belize Shelf-Carbonate Sediments, Clastic Sediments, and Ecology. The American Association of Petroleum Geologists, Studies in Geology, 2:40-52. Tulsa, Oklahoma. 

Robins, C R., and W. A. Starck II 

1965. Opsanus astrifer, a New Toadfish from British Honduras. Proceedings of the Biological Society of Washington, 78:247-250. 

Riitzler, K. 

1975. The Role of Burrowing Sponges in Bioerosion. 

Oecologia (Berlin), 19:203-216. 

1978a. Photogrammetry of Reef Environments by Helium Balloon. In D. R. Stoddart and R. E. Johannes, editors, Coral Reefs: Research Methods. Monographs on Oceanographic Metholodogy, 5:45-52. Paris: 

Unesco. 

1978b. Sponges in Coral Reefs. In D. R. Stoddart and R. 

E.Johannes, editors, Coral Reefs: Research Methods. Monographs on Oceanographic Methodology, 5:299-

313. Paris: Unesco. 

1981. An Unusual Blue-Green Alga Symbiotic with Two 

New Species of Ulosa (Porifera: Hymeniacidonidae). Marine Ecology, 2:35-50. 

Riitzler, K , J. D. Ferraris, and R. J. Larson 

1980. A New Plankton Sampler for Coral Reefs. Marine 

Ecology, 1:65-71. 

Riitzler, K., and Macintyre, I. G. 

1978. Siliceous Sponge Spicules in Coral Reef Sediments. Marine Biology, 49:147-159. 

Smith, F.G.W. 

1939. Sponge Mortality at British Honduras. Nature, 

144:785. 

1941. Sponge Disease in British Honduras, and its 

Transmission by Water Currents. Ecology, 22:416-

421. 

Stoddart, D. R. 

1960. The Reefs and Sand Cays of British Honduras. In 

Cambridge Expedition to British Honduras, 1959-1960, 

General Report, pages 16-22. Cambridge, England. 

1962a. Three Caribbean Atolls: Turneffe Islands, Lighthouse Reef and Glover's Reef, British Honduras. 

Atoll Research Bulletin, 87:1-151. 

1962b. Catastrophic Storm Events on the British Honduras Reefs and Cays. Nature, 196:512-515. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays, October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 

1969. Post-Hurricane Changes on the British Honduras 

Reefs and Cays: Re-survey, 1965. Atoll Research 

Bulletin, 131:1-31. 

1974. Post-Hurricane Changes on the British Honduras 

Reefs: Re-survey, 1972. In A. M. Cameron, B. M. 

Campbell, A. B. Cribb, R. Endean, J. S. Jell, O. 

A. Jones, P. Mather and F. H. Talbot, editors, 

Proceedings of the Second International Coral Reef Symposium, 2:473-483. Brisbane, Australia: The Great 

Barrier Reef Committee. 

Sun, H. H., and W. Fenical 

1979. Rhipocephalin and Rhipocephenal: Toxic Feeding Deterrents from the Tropical Marine Alga 

Rhipocephalus phoenix. Tetrahedron Letters, 5:685-688. 

Taylor, W. R. 

1935. Botany of the Maya Area; Miscellaneous Papers, 

VII: Marine Algae from the Yucatan Peninsula. 

Carnegie Institution of Washington Publication, 461: 

115-124. 

Thorpe, J. E., and P. K. Bregazzi 

1960. Experiments and Observations on the Corals at 

Rendezvous Cay. In Cambridge Expedition to British 

Honduras, 1959-1960, General Report, pages 22-28. 

Cambridge, England. 

Thorpe, J. E., and D. R. Stoddart 

1962. Cambridge Expedition to British Honduras. Geography Journal, 128:158-171. 
NUMBER 12 

Tsuda, R. T., and C. J. Dawes 

1974. Preliminary Checklist of the Marine Benthic 

Plants from Glover's Reef, British Honduras. Atoll 

Research Bulletin, 173:1-13. 

Tucker, D. W. 

1954. Fishes, Part I: Families Carcharhiniidae, Torpedinidae, Rosauridae (nov.), Salmonidae, Alepocephalidae, Searsidae, Clupeidae. In The "Rosaura" Expedition, 1937-1938. Bulletin of the British 

Museum (Natural History), Zoology, 2:163-214. 

Wallace, R. J., and S. D. Schafersman 

1977. Patch-Reef Ecology and Sedimentology of Glovers 

Reef Atoll, Belize. In S. H. Frost, M. P. Weiss, and 

J. B. Saunders, editors, Reefs and Related Carbonates—Ecology and Sedimentology. The American Association of Petroleum Geologists, Studies in Geology, 4:37-52. Tulsa, Oklahoma. 

Waller, T. R. 

1978. Morphology, Morphoclines and a New Classification of the Pteriomorphia (Mollusca: Bivalvia). 

Philosophical Transactions of the Royal Society of London, 

B, 284:345-365. 

1980. Scanning Electron Microscopy of Shell and Mantle in the Order Arcoida (Mollusca: Bivalvia). 

Smithsonian Contributions to Zoology, 313: 58 pages. 

Wantland, K. K , and W. C Pusey III, editors 

1975. Belize Shelf-Carbonate Sediments, Clastic Sediments, and Ecology. The American Association of 

Petroleum Geologists, Studies in Geology, 2: 599 pages. 

Tulsa, Oklahoma. 
The Habitat Distribution and Community 

Structure of the Barrier Reef Complex 

at Carrie Bow Cay, Belize 

Klaus Riitzler 

and Ian G Macintyre 

ABSTRACT 

The reef complex near Carrie Bow Cay, which 

is representative of the entire Belizean barrier 

reef, is separated from the mainland by a deep 

and wide lagoon, which grades into shallow seagrass bottoms, patch reefs, and mangrove cays on 

the outer barrier platform. A study transect west 

(lagoon) to east (open ocean) shows a distinct 

zonation of substrates and organisms that reflects 

primarily water depth and the prevailing wave 

and current regime. The shallow back reef shows 

massive coral growth, extensive pavement areas, 

and large rubble accumulations caused by hurricane surge; it is separated from the inner fore 

reef by a narrow intertidal reef crest pounded by 

waves. The inner fore reef (to 14 m depth) shows 

a characteristic spur and groove structure, with 

high buttresses in the shallow depth zone (to 10 

m) and low-relief formations on the deeper terrace. The outer fore reef includes a steep innerreef slope, a sand trough and an outer coral ridge. 

The steep fore-reef slope drops off at the top of 

the outer ridge. Many topographic features are 

comparable to those present on north Jamaican 

reefs. Corals of the genus Acropora suffer heavy 

damage but also gain wide distribution from 

storm swells. Halimeda plates dominate the coarse 

fraction of the sand substrates across the entire 

transect even far below the occurrence of the alga. 

Submarine pavement lithification is most pronounced in areas of low sediment accumulation. 

The outer ridge, although now dominated by 

Klaus Riitzler, Department of Invertebrate Zoology, and Ian G. 

Macintyre, Department of Paleobiology, National Museum of Natural 

History, Smithsonian Institution, Washington, D. C. 20560. 

Acropora cervicornis on the study transect, appears 

to be built up by more solid framework. The reef 

at Carrie Bow Cay has a community composition 

representative of the central barrier reef province 

to which it belongs, but structurally it is a composite, including features characteristic of the 

discontinuous northern and southern province 

reefs. 

Introduction 

The barrier reef complex—10-32 km wide 

(James et al., 1976) and approximately 250 km 

long—off Belize, Central America, is said to be 

the largest continuous reef in the Caribbean Sea 

(Smith, 1948; Adey, 1977). Beginning as a fringing reef off the Pleistocene peninsula of Ambergris 

Cay, it extends southward into the Gulf of Honduras. This reef complex consists of an almost 

unbroken barrier reef and numerous patch reefs 

and mangrove cays in its shoreward lagoon. The 

shelf lagoon is 20-25 km wide in the northern 

section of the complex, but where the lower third 

of the reef bends eastward towards Gladden Spit, 

the lagoon becomes more than 40 km wide before 

it opens into the Gulf of Honduras. The point at 

which the reef complex bends eastward is marked 

by two islets, one of which—Carrie Bow Cay—is 

the subject of this volume (Figure 1; Plate 1). 

Carrie Bow Cay (16°48'N, 88°05'W), known 

as Ellen Cay up to 1944, is situated on top of the 

barrier reef proper, 22 km southeast of Dangriga 

(Stann Creek) (Figure 2) and 18 km east of Sittee 
10 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 1.—Tobacco Reef looking south toward South Water and Carrie Bow cays; note cuts 

isolating Carrie Bow Cay from barrier reef trend. 
NUMBER 12 11 

88°W 

:':' Blue Ground 

Range 

1 

Tobacco Range •'.'.'.•• 

Twin Cays 

Lagoon 

Pafch Reefs •/'.•'-•'•• 

BiRS 

-.5 

•':'.'p 

South Water C 

a

 S Carrie 

M Ca) 

Carrie Bow Cut 

jM\Cur\e\ 

^ ^ Bank 

Curlew Cut 

:;: ;:::^^ui«---:v : 

•;Wee Wee Cay 

•:vU:.5fc": 

: # & 

*'**• ^ 2 

FIGURE 2.—Index map of barrier reef complex in the vicinity of Carrie Bow Cay; area of larger 

map located on inset by rectangle. 
12 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Point, which is the nearest land. Shoreward from 

Carrie Bow Cay, the eastern portion of the lagoon 

(on the barrier platform) is less than 5 m deep 

whereas the western part is as much as 20 m deep. 

The reef at Carrie Bow Cay (Figure 3; Plate 1: 

center right) is separated from the main barrier 

trend by two channels, South Water Cut to the 

north (0.4 km wide and 4 m deep) and Carrie 

Bow Cut to the south (0.7 km wide and 5 m 

deep). These and other channels through the 

barrier reef allow oceanic waters from the Caribbean Sea to flow onto the shallow barrier platform and to transport platform sediments to deep 

off-reef areas. 

South Water Cut also separates Carrie Bow 

Cay (120 m long and 40 m wide) from the larger 

South Water Cay (440 m long and 100 m wide), 

a low island less than 1 km to the north that, like 

Carrie Bow Cay, is composed of reef rubble and 

sand. Twin Cays, locally known as Water Range, 

is a swampy mangrove island 2 km to the northwest of Carrie Bow and is about 1 km in diameter 

and is divided by a meandering canal (Figures 3, 

28 a). Approximately 0.5-1.5 km west and southwest of Carrie Bow Cay and within the range of 

strong tidal currents passing through breaks in 

the barrier reef, numerous patch reefs having low 

relief occur among Thalassia seagrass in the shallow water (3-6 m) of the lagoon. Similar coral 

build-ups known as "sand bores" that reach the 

water surface and that are topped by intertidal 

sand are clustered about 3 km to the southwest. 

Although many aspects of the Belize Shelf have 

already been studied and described—most notably by Stoddart (1963), Wantland and Pusey 

(1971), Purdy (1974), Purdy et al. (1975), and 

James and Ginsburg (1978)—most earlier work 

has been geologically oriented. In contrast, our 

description of reef zonation is based on detailed 

information about composition of flora and fauna 

FIGURE 3.—Aerial view of Carrie Bow Cay looking northwest. South Water Cay (right) is a reef 

island on the barrier trend; Twin Cays (left) is a mangrove island in the shallow lagoon. 
NUMBER 12 13 

as well as topographic and substrate characteristics. Our study focuses on a 650 m research 

transect (just north of Carrie Bow Cay) that 

extends west to east from the lagoon to the deep 

fore reef and is representative of the entire barrier 

reef (Burke, herein: 509). 

ACKNOWLEDGMENTS.—We are indebted to I. 

Jewett for patiently preparing the maps and 

graphs, some of which are based on low-altitude 

helicopter aerial photographs taken by Captain 

G. Thomas. H. Adolfi, A. Antonius, G. Bretschko, 

M. Carpenter, R. Larson, B. Spracklin, and P. E. 

Videtich assisted in establishing and surveying 

the main transect and in field mapping other reef 

and lagoon habitats. A. Antonius, W. T. Boykins, 

R. B. Burke, M. R. Carpenter, and K. Sandved 

assisted with photography. 

Methods 

Vertical overlapping aerial photographs of the 

Carrie Bow Cay environs were made by a helicopter-mounted 70 mm camera with timed motorized film advance. Black and white or color 

prints (12 X 12 cm) were used to make photo 

composites (for instance, Frontispiece) from 

which topographic features could be traced on 

transparent overlays. The scales of the original 

maps, shown here in reduced figures, are 1: 15,000 

(Figure 2), 1:1500 (Figure 4), and 1:800 (Figure 

5). Low-altitude aerial photographs (like Figure 

8) were taken by balloon-suspended camera (Riitzler, 1978a). The bottom topography in depths 

greater than eight meters is not discernible on 

aerial photographs and therefore had to be ascertained from underwater wide-angle photographs, 

as well as compass, depth-gauge, and tape measurements. Sediment thickness in each zone was 

determined with the aid of a steel probe (1 cm 

diameter) that could be extended in 3 m sections. 

The study transect is approximately 200 m 

north of Carrie Bow Cay, midway across and 

perpendicular to the crescent-shaped reef crest 

that half encloses the island (Figure 4). Metal 

stakes driven into the coral and submerged and 

surface buoys anchored permanently mark the 

transect along its projected length (not bottom 

contour) of 650 m. The zero point was established 

arbitrarily in a seagrass community in the shallow 

lagoon. Zones were determined on the basis of 

bottom configuration, substrates, and relative 

abundance of predominant organisms. Surveys 

were carried out along both the transect line and 

a 50 m strip on either side of it. Relative abundance of organisms within zones was measured 

by pointcounting organisms inside a 0.25 m2 

frame randomly positioned (Riitzler, 1978b: 310, 

fig. 3). Objects overlayed by a 16-intersection grid 

(line stretched across every 10 cm) were recorded. 

Vertical projections of the points were used where 

the three-dimensional configuration of the substrate did not permit direct contact. In areas of 

high bottom relief, this method was more reliable 

and efficient than surface coverage estimates employed by Riitzler (1975:206) for it avoids distortions due to nonhorizontal substrate surfaces 

(Dahl, 1973). 

The Transect 

On the basis of dominant biological and geological characteristics, the barrier reef along the 

transect off Carrie Bow Cay can be divided into 

five major units: lagoon, back reef, reef crest, 

inner fore reef, and outer fore reef (Figure 5; 

Table 1). Each unit except for the reef crest can 

be subdivided into distinct zones. The movement 

and depth of water apparently are the main 

factors controlling the biological and geological 

zonations of this area. The lagoon unit (1.5-2.0 

m depth) is marked by weak currents and a 

significant accumulation of fine sand and silt; the 

back reef (0.1-1.0 m) is subjected to strong currents and the lagoonward transportation of sediments; the water over the intertidal reef crest is 

in an almost constant state of agitation; the inner 

fore reef (1-12 m deep) similarly is strongly affected by both storm waves and waves related to 

normal trade wind conditions; the outer fore reef 

(at least 12 m deep), on the other hand, is affected 

only by long-period storm waves. Following is a 

detailed description of each unit. 

LAGOON.—This environment consists of a sand 

and seagrass {Thalassia) zone (Figure 6) and a 
14 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

5and and Rubble 

Pavement Rock 

Scattered Coral 

Dense Coral 

Thalassia 

0 

FIGURE 4.—Map of sea-floor characteristics of Carrie Bow reef with location of permanent study 

transect. 
NUMBER 12 15 

sand and rubble zone (Figure 7) that together 

extend only 50 m along the transect line. The 

first section, which extends for about 20 m from 

the zero point northwest of Carrie Bow Cay, 

covers a patch of seagrass {Thalassia testudinum 

Banks ex Kbnig, along with a few Syringodium 

filiforme Kiitzing) on a silty sand bottom (Figure 

6a; Plate 2: top left). This sand is fair sorted, 

dominantly very fine to medium, and 1.0-1.2 m 

thick. It consists mainly of coral, mollusks, 

benthic Foraminifera (including Homotrema), and 

Halimeda plates. During one of our surveys (May 

1975) the lower 10-15 cm of the plants were 

buried in sediments, presumably as a result of 

hurricane Fifi in 1974. The alga Dictyota sp., 

which was sparse in 1975, was very common 

during May and June of 1976, 1977, and 1978. 

Other algae regularly interspersed with Thalassia 

belong to the genera Halimeda, Udotea, and Penicillus. Empty conch shells and other rubble are 

covered by algal felts and patches of crustose 

coralline algae. Uncommon but conspicuous invertebrates include the anemone Bartholomea annulata (Lesueur) inside conch shells, the corals 

Acropora cervicornis (Lamarck) and Siderastrea radians (Pallas), the gastropods Strombus gigas Linnaeus and Turbinella angulata Lightfoot, the echinoid Tripneustes ventricosus (Lamarck), and the holothuroid Holothuria mexicana Ludwig. Benthic macroinvertebrates associated with this Thalassia 

community were studied by Young and Young 

(herein: 115). 

Between 20 and 50 m along the transect, sand 

appears along with rubble (sand and rubble zone) 

largely covered by algal felts (13%) and by Dictyota 

spp. (6%) (Figure 7). The poorly sorted sediment 

in this zone ranges in size from silt to gravel and 

its composition is similar to that of sediments in 

the Thalassia zone. The metal probe recorded 

sediment thickness of 1.0-1.4 m over a hard bottom. Acropora cervicornis was absent from the sand 

and rubble zone in 1972, appeared in only two 

percent of samples by 1975 (but was very common in patches nearby, particularly just north of 

the transect), and by 1977 had taken over extensive areas on both sides of the transect (Figure 

la). A subsequent survey in spring 1979 after 

hurricane Greta (September 1978) showed that 

thickets of A. cervicornis had been broken up and 

carried deep into the lagoon by storm surge. New 

growth, however, was evident on some coral 

branches that had not been completely buried in 

sediment. During this survey, thick growths of the 

red alga Champia parvula (C. Agardh) Harvey 

(Norris and Bucher, herein: 201) appeared in 

patches throughout both the Thalassia and the 

sand and rubble zones (Figures 6b, lb). 

BACK REEF.—This unit, in which the bottom 

rises steadily from an average of 1 m to the 

intertidal reef crest, consists of a patch-reef zone 

(Figures 8, 9) and a rubble and pavement zone 

(Figure 11) that together extend from the 50 m 

mark to the 245 m mark along the transect. The 

substrate in the patch-reef zone (50-150 m marks) 

consists of gravel scattered in a poorly sorted silt 

to very coarse sand. The coarse fraction is composed mainly of Halimeda, coral, Homotrema and 

other benthic foraminiferans, mollusks, crustose 

coralline algae, and echinoids. Maximum sediment thickness recorded with the metal probe 

was 0.3 m. This zone is characterized by local 

build-ups of dead coral framework forming isolated patch reefs that consist primarily of Montastrea annularis (Ellis and Solander) and some Diploma labyrinthiformis (Linnaeus) and Agaricia agaricites 

(Linnaeus) (Figure 9). Approximately 50 percent 

of the surface area of the coral is dead and 

overgrown by crustose coralline algae and algal 

turfs (Figure 8). Characteristically, the vertical 

sides of many coral heads and boulders are alive 

but the top surfaces are dead, evidently because 

exposure at low tide has limited the upward 

growth of the corals. At the same time, considerable damage to this coral population is being 

caused directly or indirectly by the blue-green 

alga Oscillatoria submembranacea Ardissone and 

Strafforello (Antonius, 1973, 1977). Overhangs 

and crevices are commonly populated by the sea 

urchin Diadema antillarum Philippi, which, along 

with several species of parrot fishes, is responsible 

for extensive bioerosion of dead coral surfaces 

typical of this zone. Clusters of the corals Acropora 
16 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Transect Across Barrier Reef at Carrie Bow Cay, Belize 

Lagoon 

Sand a 

Sea Grass 

Zone 

Sand a 

Rubble 

Zone 

Back Reef 

Patch Reef Zone 

100 

Rubble a Pavement Zone 

I2OO 

Reef 

Crest 

Inner Fore Reef 

High Spur a Groove Zone 

I3OO L00 

- 10 

- 2 0 

- 3 0 

.'.' •: Sand and rubble 

///// / Coral rock 

* ///<*' Tf-

•••••••///••••- III" •• •• ••••••///••••• 

\t^^\\ Mounds or spurs 

8

 ® Corol heads 

•••••••:•.•.•••.•/// "»5 " '" ^ ""•£&!-

? Plaly corol 

^ M / Th" 1 0 "

1 0

 testudinum 

ft P\ Ponfera \ P i \

 Mil,e P ° r a

 complanata 

%i Millepora- Agaricia- Pontes £=-^- Acropora palmato 

FIGURE 5.—Map and bottom profile of the permanent study transect with zonation 

terminology (© — permanent markers). 

cervicornis and Pontes astreoides Lamarck, and the 

octocoral Plexaura spp. are dispersed throughout 

the sandy flats (Figure 8), with a variety of biota 

including Dictyota sp., Cliona caribbaea Carter, Thalassia testudinum, Strombus gigas, Udotea flabellum (Ellis and Solander) Lamouroux, Amphiroa sp., and 

Penicillus capitatus Lamarck—in order of abundance—growing in between. In the spring of 

1979, as a result of hurricane Greta, most of the 

A. cervicornis showed extensive damage, and drifts 

of A. cervicornis debris were piled up against the 

patch reefs (Figure 10; Plate 2: center left). The 

corals Acropora palmata (Lamarck), Pontes pontes 

(Pallas), Diploria strigosa (Dana), Siderastrea siderea 

(Ellis and Solander), and Agaricia fragilis Dana 

were observed close to this zone but were not 

included in the counts. 

The rubble and pavement zone appearing between 150 and 245 m along the transect consists 

of gravel in a silt to very coarse sand matrix that 

grades into a relatively smooth and undulating 

rock pavement adjacent to the reef crest (Figure 

11). As can be seen on the map (Figure 4) the 

pavement is generally 40 m wide and forms the 

shoreward border of the entire reef crest trend. 

This pavement (described in detail by James et 

al., 1976) consists of a conglomerate of coral 

{Millepora), mollusk, and crustose coralline algal 

fragments lithified by a magnesium calcite submarine cement. We noted a maximum pavement 

thickness of only 4 cm although James et al. 

(1976) reported lithification down to 0.5-1.0 m 

in other areas of the barrier reef off Belize. Rock 

surfaces in this zone are overgrown by isolated 
NUMBER 12 17 

;y. *,'"} 

Low Spur a Groove Zone 

Uoo 

Outer Fore Reef 

Inner 

Reef 

Slope 

Sand Trough 

1600 

Outer 

Ridge Fore Reef Slope 

meters 

_ V Acropora cervicornis 

Gorgonocea E 

coral heads {Siderastrea siderea and Pontes astreoides), scattered Dictyota spp., coralline crusts, 

and the boring sponge Cliona caribbaea. Less abundant organisms are algae of the genera Halimeda, 

Caulerpa, and Penicillus and the corals Agaricia 

agaricites, Diploria clivosa (Ellis and Solander), and 

Acropora palmata. The depth of water in this zone 

averages 0.6 m. Acropora cervicornis was abundant 

in this zone in spring 1978 (Figure 11a) but had 

nearly disappeared by the 1979 survey (Figure 

1 lb), apparently owing to storm surges associated 

with hurricane Greta, which transported almost 

all living A. cervicornis lagoonward. Due south 

from the 150-200 m marks, between the transect 

and the island, the same zone changes into a 

Thalassia-dom'mated rubble flat, with Pontes porites, Siderastrea radians, crustose corallines (on rubble), and Diadema (Plates 1: bottom right, 2: top 

right). 

REEF CREST.—A transition zone (0.2 m deep) 

occurs between the back reef and fore reef at the 

245-265 m marks along the transect. This zone is 

distinguished by a framework of dead (60%) and 

living Acropora palmata along with Agaricia agaricites 

and Pontes astreoides (Figure 12; Plate 2: bottom left, 

bottom right). The bottom consists mainly of rubble 

and rock pavement covered by crustose coralline 

algae along with patches of fine to medium, wellsorted sand, the coarser fraction of which is 

mainly Halimeda, coral, Homotrema, and crustose 

coralline algae. This sand supports scattered 

growths of Dictyota sp. whereas the dead coral 

rock is commonly covered by algal turfs and 

dense stands of Caulerpa racemosa (Forsskal) J. 

Agardh. The section of the reef crest unit between 

265 and 270 m along the transect (depth of 0.1 

m) is dominated by the coral Acropora palmata, the 

hydrocoral Millepora complanata Lamarck, and the 
18 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 1.—Relative abundance of dominant organisms on the Carrie Bow Cay main transect 

(column heads designate region, structural and biological zone, transect position in meters 

from 0, depth in meters as average or range, and substrate in order of abundance; ++ + = 

Biota 

Fleshy macro-algae 

Calcareous macro-algae 

Crustose Corallinacea 

Algal felts and turfs 

Thalassia testudinum 

Massive Demospongea 

Cliona spp. 

Millepora alcicornis 

M. complanata 

Palythoa caribaeorum 

Stephanocoenia michelinii 

Madracis mirabilis 

Acropora cervicornis 

A. palmata 

Agaricia agaricites 

A. fragilis 

A. lamarcki 

A. tenuifolia 

Leptoseris cucullata 

Siderastrea siderea 

Porites astreoides 

P. porites furcata 

Diploria clivosa 

D. labyrinthiformis 

D. strigosa 

Colpophyllia nalans 

Montastrea annularis 

M. cavernosa 

Meandrina meandrites 

Dichocoenia stokesi 

Dendrogyra cylindrus 

Mycetophyllia danaana 

Eusmilia fasligiata 

Erythropodium caribaeorum 

Erect Gorgonacea spp. 

Strombus gigas 

Diadema antillarum 

Didemnidae 

Lagoon 

Sand-mud 

Thalassia 

0-

2 

M 

20 

uddy sand 

++ 

++ 

+ ++ 

P 

Sandrubble 

Algal 

felts 

20-50 

1.8 

Sand 

rubble 

++ 

+++ 

+ 

Back reef 

Patch reefsand 

MontastreaDiploria 

50-150 

1 

Sand 

rock 

rubble 

+ 

+ 

++ 

+ 

+ 

+ 

++ 

P 

++ 

P 

P 

++ 

P 

+++ 

P 

+++ 

++ 

+ 

++ 

Rubblepavement 

SiderastreaPorites 

150-245 

0.6 

Rubble 

rock 

sand 

+++ 

++ 

++ 

+ 

++ 

P 

+ 

++ 

+ ++ 

+++ 

+ 

+ 

P 

P 

+ 

+ 

Coral rocksand 

AcroporaAgaricia 

245-265 

0.2 

Rock 

sand 

rubble 

++ 

++ 

++ 

+ 

+ 

-r-r-r-

-r-r-t-

+ 

++ 

+ 

Reef crest 

Coral rockrubble 

AcroporaMillepora 

265-270 

0.1 

Rock 

rubble 

+ 

++ 

++ 

+ ++ 

+ ++ 

+ ++ 

+++ 

++ 

Rubblecoral rock 

CorallinesMillepora 

270-275 

0.1 

Rubble 

rock 

+ ++ 

+ 

+ ++ 

++ 

++ 

+ 

+ 
NUMBER 12 19 

very common (>10% presence); + + = common (5%-10% presence); + = less common (1%-

5% presence); P = present and obvious but not encountered in the statistical samples) 

Coral pinnacles 

MilleporaAcropora 

275-330 

1.8 

Rock 

sand 

rubble 

+ + 

Inner fore reef 

Spur 

High relief 

AgariciaAcropora 

330-410 

5 

Sand 

rock 

rubble 

+ + 

and groove 

Low relief 

GorgonaceaMontastrea 

410-550 

10 

Sand 

rubble 

rock 

+ + 

Inner reef 

slope 

AcroporaMontrastrea 

550-575 

15-22 

Rubble 

+ 

+ 

Sand trough 

GorgonaceaMontastrea 

575-615 

23 

Sand 

rubble 

Outer fore reef 

Outer ridge 

Inner slope 

AcroporaDiploria 

615-625 

12-22 

Rubble 

+ 

+ + 

Top 

AcroporaGorgonacea 

625-645 

14 

Rubble 

sand 

+ + 

Fore reef 

slope 

MontaslreaGorgonacea 

645-655 

15-30 

Rock 

rubble 

sand 

+ 

+ 

+ 

+++ 

+++ ++ 

+ + + 

+ 

+ + + 

+ + + 

+ + 

+ + 

p 

p 

+ + 

+ 

+ 

+ 

+ + 

+ 

+ + 

+ + + 

+ + 

+ + + 

+ 

+ 

+ + 

+ + 

p 

p 

+ 

+ 

+ 

++-f-

+ 

+ 4-

+ + 

+ 

+ 

+ 

+ + 

+•+ 

+ + + 

+ + 

+ + + 

+ + 

+ + 

+ + 

+ 

+ + + +++ 

++ 

p 

p 

++ ++ 

++ 

+-1-

+++ 

p 

p 

+ 

+ + 

+ + + 

+ + 

+ 

+ + + 

+ + 

+ 

+ + 

++-1-

+ + 

+ + 

+ + 

+ 

+ + 

++ 

++ 

++ 

+++ 

p 

p 

++ 

++ 

p 

+++ +++ ++ —I—I—I — 

++ 

+++ 
20 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 6.—Sand and seagrass zone before and after hurricane Greta: a, March 1978; b, April 

1979. (Note extensive development of epiphytic red alga Champia parvula on Thalassia testudinum 

blades; scale = 40 cm.) 

FIGURE 7.—Sand and rubble zone before and after hurricane Greta: a, March 1978, stands of 

Acropora cervicornis on plain substrate; b, April 1979, heavy red algal growth (Champia parvula) on 

substrate devoid of living A. cervicornis. (Scale = 40 cm.) 

zoanthid Palythoa caribaeorum Duchassaing and 

Michelotti (Figure I3a,b; Plate 3: top left, top 

right). Coralline algal crusts, algal turfs, and Porites astreoides are attached to massive, extensively 

bored coral rock. The last windward section of 

the intertidal reef crest unit (270-275 m along the 

transect) is built up by rubble and coral rock 

(60%), covered by coralline crusts, Palythoa caribaeorum, Millepora complanata, some Porites porites, 

Agaricia agaricites, algal felts, and extensive crusts 

of the alcyonacean Erythropodium caribaeorum (Duchassaing and Michelotti). Just south of the transect line and at four or more other locations the 

reef crest is interrupted by perpendicular channels (Figure 13<r), which facilitate the water exchange between lagoon and fore reef (Kjerfve, 

herein: 59). 

At its southern limit, due southeast of Carrie 

Bow Cay, the reef crest zone grades into a coralrubble storm ridge, which is actively accreting 

over the rubble and pavement zone (Figures 4, 

14; Plates 2: center right, 5: top left). Our observations indicate that the shoreward transportation 

of this material occurs only during severe storms 
NUMBER 12 21 

FIGURE 8.—Low-altitude aerial view of patch reef zone at the 100 m transect mark (lower 

center). Most of the patch reef is built up by Montastrea annularis (light areas are dead), 

accompanied by Agaricia agaricites and Acropora cervicornis; the latter also forms an extensive 

thicket on the right flank of the patch reef. (Picture area: 6X 6 m.) 

or hurricanes. During lesser storms this rubble is 

quite stable because much of it is platy (derived 

from Acropora palmata) and offers protection to a 

diverse interstitial invertebrate fauna (Plate 5: top 

right). Another comparable area on the reef crest 

is a small incipient rubble field located 50 m 

north of the transect line (Figure 4). 

INNER FORE REEF.—Just east of the 275 m 

mark along the transect the bottom drops to a 

depth of 1.8 m and the inner fore reef begins in 

a transition zone (to 330 m along the transect) of 

tall but unorganized coral pinnacles on a sand 

and rubble bottom (Figure 15; Plate 3: center left). 

These pinnacles are the result of turbulent water 

movement related to oscillating wave energy that 

is partly reflected by the reef crest. This area is 

comparable to Chevalier's (1973) Type II Outer 

Reef at Mururos, in which he related "pillar 

structure" of corals to heavy wave swell. Approximately half of these massive pinnacles off Carrie 
22 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 9 (above).—Montastrea annularis patch reef on rubble 

substrate. Massive corals in this zone tend to form pedestalshaped bases. 

FIGURE 10 (below).—Acropora cervicornis thicket in patch reef 

zone badly damaged by hurricane Greta, September 1978. 

(Photograph taken April 1979.) 
NUMBER 12 23 

FIGURE 11.—Rubble and pavement zone before and after hurricane Greta: a, March 1978, 

extensive Acropora cervicornis and Dictyota sp. on sand and rubble bottom; b, April 1979, most of 

the dominant organisms and large quantities of sand have been transported lagoonwards. 

(Scale = 40 cm.) 

FIGURE 12.—Inner margin of reef crest; algal turf covered coral rock surfaces show evidence of 

rasping action by parrot fish; Caulerpa racemosa, Millepora complanata, and Porites astreoides in 

foreground. 
24 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

*%> r%. 
NUMBER 12 25 

FIGURE 14.—Coral rubble storm ridge, south of the transect, photographed after hurricane Fifi 

(1974); rubble prograding over back-reef pavement, partly burying Montastrea annularis colony. 

Bow Cay are formed by build-ups of dead Acropora 

palmata, the rest by live Millepora complanata, A. 

palmata, and Agaricia tenuifolia Dana. The dead 

framework is extensively covered by encrusting 

Cliona caribbaea, coralline algae, and some Palythoa 

caribaeorum. Diploria strigosa, Porites astreoides, P. 

porites, Montastrea annularis, and M. cavernosa (Linnaeus) are massive corals of secondary importance. 

The spur and groove zone of high relief (330-

410 m along the transect) has an average depth 

FIGURE 13.—Seaward margin of reef crest: a, Millepora complanata, with Porites porites (foreground) and Acropora palmata 

(background); b, A. palmata community; c, Reef channel 

through Millepora ridge looking toward shallow inner reef 

crest (note overturned live colony of A. palmata in foreground). (Scale frame = 50 X 50 cm.) 

of 5 m, but depth ranges from 3 to 10 m between 

the highest coral spurs and deepest sand grooves 

(Figure 16; Plate 3: center right). This well-defined 

zone has been discussed by Wantland and Pusey 

(1971) and has been described in detail by James 

et al. (1976). The high-energy oscillating movement of water in this zone has promoted coral 

growth on the spurs (Shinn et al., herein: 63) and 

has caused erosion in the grooves. The buttresses are characterized by foliate Agaricia tenuifolia and Millepora complanata enclosing clusters of 

Porites porites (Figure 17; Plate 3: bottom left). The 

tops of many buttresses are dominated by stands 

of Acropora palmata and A. cervicornis (Figure 18) 

and their flanks and the sand grooves in between 

by Agaricia agaricites, Diploria strigosa, various species of gorgonians (Muzik, herein: 303) and fleshy 
26 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 15.—Pinnacles of Millepora complanata and Agaricia tenuifolia in an area of transition 

between reef crest and high-relief spur and groove zone. 

green algae (Figure 16; Plate 3: center right, bottom 

right). Of lesser importance are Siderastrea siderea, 

Pontes astreoides, Dichocoenia stokesi Milne Edwards 

and Haime and Stephanocoenia michehnii Milne 

Edwards and Haime. A characteristic and abundant—but quantitatively unimportant—component of this fauna is the purple hydrozoan Stylaster 

roseus (Pallas) that occurs in niches and overhangs 

along the sides of the buttresses. Another hydrozoan, Millepora alcicornis Linnaeus, commonly 

grows over the dead skeletons of gorgonians, 

mainly of Gorgonia ventalina Linnaeus. The medium sand to gravel-sized sediment in the grooves 

has a maximum thickness of 0.3 m. It has fair 

sorting and is composed mainly of coral, Halimeda, 

Homotrema, and mollusk and echinoid debris. 

The spur and groove system of low relief extends between 410 and 550 m along the transect 

(average depth, 10 m). A diverse population of 

gorgonians dominates the rock and rubble substrates of the sand flats as well as the low (about 

1 m relief) coral spurs, which are formed of 

Montastrea annularis, M. cavernosa, Acropora cervicornis, and Diploria strigosa (Figure 19, 20; Plate 4: 

top left, top right). A few island-like coral pinnacles 

attain 3 m in height and diameter. Massive Demospongea become quantitatively important in 

this zone of reduced agitation of near-bottom 

water, for example, Neofibularia nolitangere (Duchassaing and Michelotti), Callyspongia spp., Aplysina spp., Geodia neptuni (Sollas), Ircinia spp., as 

well as the thickly encrusting Anthosigmella vahans 

(Duchassaing and Michelotti). Halimeda spp. and 

Dictyota spp. are common algae. Conspicuous corals having patchy distribution are Agaricia agaricites, A. tenuifolia, Diploria labyrinthiformis, Dendro-
NUMBER 12 27 

FIGURE 16.—High-relief spur and groove zone, having 5 m relief. (Water depth at sand bottom: 

6 m.) 

gyra cylindrus Ehrenberg, and Eusmilia fastigiata 

(Pallas). Probes of the sand-filled grooves indicated a maximum depth of 1.2 m at the shallow 

end of this zone. Seaward, these sand lenses thin 

out and eventually give way to sand pockets in a 

rock pavement. The sediment is similar to that 

found in the high-relief spur and groove zone. 

James et al. (1976:532) referred to this low-relief 

spur and groove zone as the "deep spur and 

groove," which was described as being separated 

from a "shallow spur and groove" system by a 

"rubble-covered terrace." There is no indication, 

however, that a terrace separates the high- and 

low-relief spur and groove zones off Carrie Bow 

Cay. In fact, the spurs having low relief are 

commonly a continuation of the shallower highrelief spurs, with one or more shallow grooves 

spilling into one of the deeper and wider grooves. 

Furthermore, the relief of the spurs is considerably 

less than that reported by James' group who 

described spurs having relief of 3-4 m that rise to 

within 2 m of the surface of the waters. 

OUTER FORE REEF.—This region begins with a 

25-degree slope—the inner reef slope—where the 

transect drops from 15 m depth at 550 m along 

the transect to 22 m at a position 575 m along 

the transect. Most of the bottom is covered by a 

thicket of living and dead Acropora cervicornis that 

offers substrate to some massive sponges—Verongula gigantea (Hyatt), Callyspongia vaginalis (Lamarck)—and various gorgonians. Columnar colonies of Montastrea annularis at the top of the slope 

give way to large platy colonies towards the base 

that are accompanied by Porites astreoides, Siderastrea siderea, and Agaricia tenuifolia (Figure 21; Plate 

4: center left). Very poorly sorted sediment com-
28 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 17.—Dominant framework builders of buttresses in high-relief spur and groove zone: a, 

Agaricia tenuifolia capped by Acropora palmata; b, close-up view of characteristic coral association 

consisting of A. agaricites, Millepora complanata, and Porites porites. (Scale = 40 cm.) 

FICURE 18.—Acropora palmata development on top of buttress in high-relief spur and groove 

zone. (Depth: 4 m.) 
NUMBER 12 29 

FIGURE 19.—Low-relief spur and groove zone with scattered 

coral heads (foreground: Montastrea annularis) among octo 

corals on the rock pavement of a low spur. (Depth: 10 m.) 

FIGURE 20.—Sand-covered hard ground dominated by octocorals with islands of coral heads, near outer edge of lowrelief spur and groove zone. (Depth: 13 m.) 

posed of silt to medium-sized sand with scattered 

coarser debris, occurs in patches on this slope. 

Halimeda plates, mollusks, and echinoid spines 

make up most of the readily recognized coarse 

fraction. This inner reef slope is comparable to 

"the steep coral-veneered rock slope" that James 

and Ginsburg (1978:33-35) called the "reef step." 

The next section along the transect is a sand 

trough 40 m wide (575-615 m marks) and an 

average of 23 m deep (Figure 22). The substrate 

is a poorly sorted, silt size to very coarse sand 

sediment plain. This sediment is mainly very fine 

to fine sand, but coarser material consisting of 
30 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 21.—Platy coral development (Montastrea annularis) at base of inner reef slope; note 

brick with coral transplant (Graus and Macintyre, herein: 441). (Depth: 21 m; scale = 40 cm.) 

Halimeda plates, mollusks, benthic foraminiferans, 

and echinoids is scattered throughout. Probings 

indicated that sediment varies in thickness from 

about 1 m at the toe of the inner reef slope to 

more than 12 m in the axis of the trough. Pieces 

of rubble support gorgonians and sponges, as well 

as some corals that form several small, isolated 

coral patches, predominantly of Montastrea annularis and M. cavernosa (Figure 22a). This sand 

trough zone on the transect also encompasses one 

tall coral pinnacle in the western part of the 

trough that has a coral-gorgonian composition 

similar to that of the inner slope with which it is 

connected by a low Acropora cervicornis ridge (Figure 4). This zone correlates with the seawarddipping sediment terrace off Tobacco and Buttonwood cays, where the slope is not bordered by 

an outer ridge (James and Ginsburg, 1978). 

At Carrie Bow Cay an outer ridge runs parallel 

to the intertidal reef crest and delineates the 

continental shelf (Figure 4). On the transect it is 

formed mainly by a thicket of Acropora cervicornis 

(Figures 22b, 23a), but south of the Carrie Bow 

transect Montastrea annularis becomes the principal 

framework builder (Figure 23 b). The steep 45-

degree landward slope of the outer ridge (Figure 

22b) supports, among the branches of A. cervicornis, massive Diploria labyrinthiformis, Porites astreoides, and Stephanocoenia michelinii, gorgonians, 

large sponges—Pseudoceratina crassa (Hyatt), Xestospongia sp.—and conspicuous algae—Halimeda 

spp., Slypopodium zonale (Lamouroux), Peyssonelia 

sp. The top of the outer ridge lies between 625 

and 645 m along the transect and its depth ranges 

from 12-14 m (Figure 23a). Acropora cervicornis 

and gorgonians are the dominant organisms, with 
NUMBER 12 31 

- 4* 

a "/•?**• , 

^ - * •'>-:-• 

& 

FIGURE 22.—Sand trough: a, isolated coral patches at the base of inner reef slope, 23 m deep; 

b, view from the top of the outer ridge over Acropora cervicornis thicket, 12 m relief. 
32 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 
NUMBER 12 33 

FIGURE 24.—Fore-reef slope: a, looking up sand shoot from 25 m; b, platy coral-octocoral 

community on transect at 30 m. 

local accumulations of Montastrea annularis, Agaricia agaricites, A. tenuifolia, and Madracis mirabilis 

(Duchassaing and Michelotti). Fleshy green algae, and sponges of the genera Aplysina, Verongula, 

Callyspongia, Xestospongia, and Agelas are common 

associates (Plate 4: center right). Poorly sorted, very 

fine sand to gravel-sized sediments that are rich 

in Halimeda plates occur in small, 1 m deep depressions scattered along the crest of the ridge. 

An identical shelf-edge ridge occurs off South 

Water Cay (James and Ginsburg, 1978). 

Approximately 645 m along the transect, the 

fore-reef slope (Figure 24) drops at an angle of 

50°-70° from depths of 14 m down to 30-60 m, 

FIGURE 23.—Outer ridge: a, coral community dominated by 

Acropora cervicornis on transect line, 12 m depth; b, platy 

Montastrea annularis and Porites astreoides framework 100 m 

south of transect, 14 m depth. (Scale = 40 cm.) 

0 -

2 0 -

4 0 -

1

 60 1 

E 

8 0 -

100-

120-

/ 

WJ 

1 j j ; A\ , Shallow Reef 

/ Ikm 

m <• South Woler Coy 

V 

South Water Cut 

£/Com e Bow Cay 

Carr e Bow Cut 

140-

160-

t 

% 180 

E 

200-

220-

^ Fore Reef Slope 

= 3 Lower limit of reef community 

= j Deep Fore Reef 

s £ . Blocks of reef limestone 

\l 

. •%_ , Continental Slope 

• A%v Coral rubble in sand 

•^\ . Lime mud 

100m AAu" ^ A \ > 

FIGURE 25.—Profile and dominant sea-floor characteristics 

of fore reef off South Water Cay (adapted from James and 

Ginsburg, 1978). 
34 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 26.—Close-up view of representative fore-reef slope community of sponges, corals, and 

octocorals, including (clockwise from center bottom) Cinachyra kuekenlhali, Amphimedon compressa, 

Agaricia lamarcki, Mycale sp., Iciligorgia schrammi, Porites porites, Pseudopterogorgia sp., and encrusting 

Millepora sp. (Depth: 30 m.) 

where it diminishes to 40°-50°. According to 

previous observations from the research submersible Nekton off nearby South Water Cay (James 

and Ginsburg, 1978), a vertical wall continues 

from a depth of 60 m to about 110 m where a 

talus of large reef limestone blocks, coral rubble, 

and Halimeda sand grades into a gently sloping 

lime-mud bottom at about 200 m (Figure 25). 

Our surveys, which were restricted to a maximum 

depth of 30 m, show an abundance of platy 

Montastrea cavernosa and M. annularis, Gorgonacea, 

Agaricia fragilis, Leptoseris cucullata (Ellis and Solander), and Demospongea (Figures 246, 26, 27; 

Plate 4: bottom left, bottom right). Commonly associated are A. agaricites, A. lamarcki Milne Edwards 

and Haime, Porites porites forma furcata Lamarck, 

Diploria labyrinthiformis, Colpophyllia natans (Houttuyn), and Mycetophyllia danaana Milne Edwards 

and Haime. Abundance of gorgonians decreases 

with depth whereas that of sponges increases. A 

conspicuous gorgonian under 20 m is Iciligorgia 

schrammi Duchassaing (Figure 27). Quantitatively 

important sponges are rope-shaped species of 

Aplysina, Niphates, and Haliclona, tubular Agelas 

sp. and Verongula sp., massive Ectyoplasia ferox (Duchassaing and Michelotti) and Cinachyra kuekenlhali Uliczka, and coral-eroding Cliona delitrix 

Pang and Siphonodictyon coralliphagum Riitzler. Hal-
NUMBER 12 35 

FIGURE 27.—Vertical fore-reef slope with platy development of Montastrea cavernosa and 

abundant sponges and octocorals. (Depth: 40 m.) 
36 SMITHSONIA

N CONTRIBUTIONS TO THE MARINE SCIENCES 

<-» u to C <U ?! .S E o .£. + 4) C • - £ 5 c a. o o __ w> c T3 « 5 c — g 

(fl u o to "5

 u 

5 c 

C 3 3 & ° ^-i o ~ e (fl 

=o 35 

(fl + * + 

fir? " ^ -a 

c 2 a P c

 b ° 5 II u o ~ B S I 

j2 .a « 

(fl c 

to _C O >~ C 

t£ o 

^ >< V « ° Io s S..S K

 M> Ml 

a 

-a 

. c 

3 bo D "n 6S A S * tfl a 8 S I « "5 T3 

tfl a. 

w> bo + •-

cf -if

 u 

o 3 (A E (fl H 

u (fl s I ^ u- o > 3 o u (fl « 

S -l fd 

2 O

5 

^3 

s rier l OQ 

^ Si o 

-«2 ^j CO ! 

| s» 

1 

g>o 

1 

" ] 2 a &o « o 

<>s ^ J ^ > ~ ^ £ 3 i » 

O j ^ 

"*-> 

•^ ^ <£> 

^ 

* t-) 

£ C M y* OS 

> Si co A ; CO 

r l " 05 

K § £ got- <3 —i 

l < | 

to 13 

R 

t 

s Is 

,C ^M O yO , r s ^ CO —< • * £S C1 + + + + + + + + + 1 1 

+ 

+ + + 1 1 + + 

+ 

Co" CM

-

 CM* 

cxi TH •* 

1 + + + + + 

+ 

S" OC? CO

-

CM CO CO w ' ^ / W ' 

1 + + + + + + + + + CM CO CO CM O r«." ^^ c^ ^ 1 + + + + + + + 

*—. S- v ^—v Tf TT" r^ 

iii p~i •>*< 

1 + + + + + + + + + f ^ CC? i n t o m ~—' 1 + + 1 + + 

+ 

CO 

n 

1 

+ 

+ 

+ 

1 

1 

1 

1 

1 

1 

0 

o 

•~ 

V 

> 

o h be 

C 

(fl 

0. 

5 0 

0 -l < 

z 

0 

P 

u 

z 

3 

,L ^ O 

00, 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

1 

1 

1 

,L s O O, 

1 

+ 

ST 

c\i CM 

+ 

+ 

+ 

u CO 

(fl Iw be 

U 

c 'C 

(fl 

i 

+ 

+ 

+ 

+ co CO 

+ 

CT) O^ 

1 

+ 

r»> CO 

+ 

^-CM ^ CO 

+ 

+ 

(D CO 

+ 

+ 

+ 

(fl bp 

(fl 

6 

u 

U 

(fl 6 >^ to (U 

fc 

1 1 + 1 

+ + 1 

+ r~ I-I —J CM + + 00 S ^ in + + 

+ 

^—N ID <X5 CM LO — ' CO + + 

+ 

+ 

^-CO^ C ^M s 

(X) uS + + + + 

(S~ o . 1 1 

+ 

V 

(fl be (fl — V 

(fl (J P c ( j ^ r- (A ^ O S u O OJ (U 5i 1 5 

(fl Ui 

u u 

^^ i n 

52-

+ 

+ 

+ 

+ 

+ 

+ 

+ r^ CM 

+ + 

+ 

r-~ CM 

w 

+ 

1 

1 

1 

(fl 

u 

to 

C 

o CO 

o 

E 

U 

Q 

D 

> 

' t o to (fl 

s 

^ -N /•—s ^—s y^^v 

to CM r~~ r~ 

O r-' oS o 1 + + 1 + + + + + + 

1 11 + 

+ 11 1 + + + in o in oq CM o CO o — ' CO — s -

x 

+ + + 1 + + + + + q oo -< m_ i n iri (£> o 

" - ' CO

 v

~ '

 v

—' 

+ + + 1 + + + 

+ 

^_^ ^_^^ 

(£> — r~~ - ; —; o 

S2- ^^ ^ 

1 + + 1 + + 

+ 

,p s i f sn c f ^q i f s n CO • * CO o v JVJ - w + + + 1 + + + 

+ 

Co" Co" o^ o^ 

1 1 1 1 + + 

to OJ VI to c g

- o ' C 2 s

 (fl 

<fl (fl S CO > c o c 

tfl p bo C 

3 B s -s (3 ffi o o 
NUMBER 12 37 

imeda-rich, very fine sand to gravel-sized sediment 

occurs in pockets or small ledges between the 

living cover of this slope. Other components of 

this sediment include mollusks, echinoids, corals 

and benthic foraminiferans. 

Lagoon Environment 

Reef-forming organisms and other characteristic and quantitatively prominent associates observed on the barrier reef transect off Carrie Bow 

Cay are all sessile and require stable substrates to 

keep them from being washed away. In the protected lagoon fine sediments tend to bury such 

substrates and, by settling on organisms as well, 

constitute a selective stress factor for the living 

populations. Thus, we examined the lagoon 

within a radius of 2 km from Carrie Bow Cay in 

order to compare its characteristic benthic biota 

with those of the barrier reef (Table 2). A similar 

comparison of zooplankton is presented by Ferraris (herein: 143) and complemented by Riitzler 

et al. (1980). Although seagrass flats and mangroves were surveyed only qualitatively, patch 

reefs were examined both quantitatively and 

qualitatively. The Carrie Bow Cay reef flat (between the cay and the reef crest) is the subject of 

a separate detailed study (Riitzler, in prep.). 

SEAGRASS COMMUNITY.—An area of approximately 6 km of lagoon bottom immediately due 

west of Carrie Bow Cay was found to be less than 

6.5 m deep, commonly 4-6 m deep. More than 

90 percent of this area is flat soft bottom covered 

by Thalassia testudinum; the rest consists of rubble, 

reef patches, and large sponges. Most of the rubble originates from the commercial conch Strombus 

gigas. The shells, abandoned by fishermen in large 

piles or fields, provide substrate for a variety of 

algae (for instance, Amphiora fragilissima (Linnaeus) Lamouroux), sponges {Desmapsamma anchorata (Carter)), hydrocorals {Millepora sp.), corals {Porites sp., Siderastrea siderea), gorgonians 

{Plexaura sp.), and many less conspicuous organisms. The dark interior of these shells constitutes 

a well-vented miniature cave habitat because 

fishermen puncture the spire near the apex where 

they cut the retractor muscle to remove the soft 

parts of the snail. Extent of colonization depends 

on exposure time and position of aperture relative 

to the sediment substrate. Small reef fishes, encrusting coralline algae, foraminiferans {Homotrema rubrum (Lamarck)), sponges {Spirastrella sp., 

Clathrina sp.), bryozoans and ascidians, as well as 

anemones {Bartholomea annulata), crabs {Mithrax 

sp.), and ophiuroids {Ophiotrix spp.) are common 

inhabitants. Several species of Cliona excavate the 

walls of old shells, some of which are occupied by 

the hermit crab Petrochirus diogenes (Linnaeus). 

Distribution of living conchs is patchy and 

their depletion in large areas observed by us over 

a seven-year period indicates over fishing. On the 

other hand we noted concentrations near certain 

conch rubble patches comparable to those reported by Hesse (1979). Another large gastropod, 

Turbinella angulata, is a common associate. Sponges 

are quantitatively the most important organisms 

in the Thalassia meadows, even outside the rubble 

fields. In 1975 and 1978 many were found loose 

and only partly alive, together with gorgonians 

in similar condition (Muzik, herein: 303) and can 

be assumed to have been torn from the reef and 

washed into the lagoon by hurricane surge (Riitzler and Ferraris, herein: 77). The presence of 

recently sunken coconut trees confirms this assumption. Other sponges, however, were healthy 

and attached to small pieces of rubble and seagrass {Desmapsamma anchorata, Iotrochota birotula 

(Higgin), Aplysina fistularis (Pallas), Niphates erecta 

Duchassaing and Michelotti) or, like some very 

large forms (5-40 1 volume), were rooted in sand 

{Spheciospongia vesparium (Lamarck), Ircinia spp.). 

PATCH REEFS.—These lagoon reefs of low relief 

are clustered about 0.3-2.0 km to the west and 

southwest of Carrie Bow Cay (Figure 2). Favorable substrate conditions, together with tradewind and tide-induced currents passing through 

South Water and Carrie Bow cuts probably promoted the development of these structures that 

now exhibit a richness of reef fauna that is surpassed only by the outer barrier fore reef. Diversity and biomass of sponges, in particular, are 

higher than in any other habitat of similar depth 

because most species favor areas having a high 

rate of water exchange but lack resistance against 
38 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

. A _.:> * 

FIGURE 28.—Twin Cays: a, aerial view of mangrove island showing channel and intertidal mud 

flats behind red mangrove fringes; b, red mangrove (Rhizophora mangle) on mud bank bordering 

channel. 

detachment from substrates by undulating water 

movement. The patch reefs are circular or oval, 

some are arranged in string-of-pearl fashion, 5-60 

m in diameter and raised 0.2-1.0 m above the 

surrounding flat sand and Thalassia bottom in 

depths of 3-6 m. Abundance data were derived 

from two perpendicular transects across a characteristic patch reef, named "Spaghetti Reef," 

after a new species of stringy sponge of the genus 

Ulosa (Riitzler, 1981). Sand and coral rock make 
NUMBER 12 39 

FIGURE 29.—Seagrass community of the shallow lagoon at the entrance to Twin Cay channel: 

Thalassia testudinum, Halimeda incrassata, and Manicina areolata. (Depth: 1 m.) 

up 45% of the surface area. Gorgonians are the 

most abundant organisms (20%), followed by corals and milleporids (17%), sponges (9%), and 

Thalassia seagrass (on sand, 9%). Large algae are 

notably absent. Gorgonia ventalina and Pseudopterogorgia spp. are the most conspicuous octocorals on 

the patch reefs. Massive forms dominate among 

the corals {Siderastrea siderea, Diploria labyrinthiformis, D. strigosa, Montastrea annularis), the most common hydrocoral is Millepora alcicornis, which encrusts numerous gorgonian skeletons. Sponges, 

although they score comparatively low in the 

point counts, contribute most to the standing 

crop. The principal species are Ircinia spp., Xestospongia sp., Iotrochota birotula, Desmapsamma anchorata, Amphimedon compressa Duchassaing and Michelotti, and Callyspongia spp. (Plate 5: center left, 

center right). 

MANGROVE.—Toward Twin Cays from the 

southeast a belt of shallow (1 m) Thalassia flat 

grades into the larger entrance of the channel 

that divides this island (Figure 28a). The flat is 

a transition zone between the more agitated 

deeper lagoon and the protected mangrove where 

moderate tidal currents control water exchange. 

Most abundant among the Thalassia are some 

algae (for instance, Penicillus capitatus, Halimeda 

incrassata (Ellis) Lamouroux), sponges {Tedania 

ignis (Duchassaing and Michelotti), Oligoceras violacea (Duchassaing and Michelotti), Haliclona viridis (Duchassaing and Michelotti)), and the coral 

Manicina areolata (Linnaeus) (Figure 29). Millepora 

sp. encrusts large surface areas of submerged 

wood. Some more sponges, Aplysina fulva (Pallas) 

and the large loggerhead Spheciospongia vesparium, 

and numerous starfish {Oreaster reticulatus (Lin-
40 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

naeus)) occur on the bottom of the channel entrance. 

The meandering channel is 0.5-2.0 m deep and 

is lined by red mangroves, Rhizophora mangle Linnaeus, which grow on intertidal mud banks and 

extend their arched prop roots into subtidal water 

(Figure 28.6). In places the mud banks are washed 

out to form vertical walls and even overhangs 

and caves. Mud caves, probably of similar origin, 

are found outside the present mangrove margin 

just north of Twin Cays. They extend horizontally into mud banks surrounding 4 m depressions 

in the seagrass floor. The caves are large enough 

to harbor 2 m long sharks. 

Mangrove roots offer the only substrates that 

are not subject to accumulation of fine sediments. 

In this habitat strong competiton for the limited 

space takes place among diverse flora and fauna 

(Figure 30). On the light-exposed roots and bank 

edges, clusters of the algae Caulerpa verticillata 

J. Agardh and Halimeda spp. compete with the 

sponges Tedania ignis, Ircinia felix (Duchassaing 

and Michelotti), and Lissodendoryx sp. (Figure 

30a,b; Plate 5: bottom left, bottom right). The walls 

of the shaded overhangs are dominated by the 

sponge Ulosa ruetzleri Wiedenmayer and, at the 

north entrance of the channel, by Mycale sp. Also, 

locally important in biomass are the anemones 

Bartholomea annulata and Condylactis gigantea (Weinland), the tunicates Ecteinascidia turbinata Herdman and Ascidia nigra (Savigny), and sabellid 

polychaetes. The intertidal parts of the mangrove 

roots are dominated by the oyster Crassostrea sp., 

whereas the Thalassia mud bottom supports a 

dense population of Scyphomedusae {Cassiopea 

spp.), with specimens of Oreaster reticulatus interspersed (Figure 30 c). Mud caves and roots provide hiding places for a diverse fish fauna (Figure 

30-6). 

Summary and Conclusions 

This paper presents the first detailed description of the biological-geological zonation of the 

barrier reef complex off Belize. Despite some 

variation along the barrier reef (see Burke, 

herein), the zonation of the Carrie Bow Cay 

segment is typical of the entire reef platform 

(Figures 4, 31). Except for its large lagoon and 

greater distance from land, the Belizean barrier 

reef is comparable to well-investigated fringing 

barrier reefs off the north coast of Jamaica (Goreau, 1959; Goreau and Land, 1974). 

Seaward of the Thalassia-dormm.ted lagoon, 

the back reef occurs between an area of massive 

coral heads on rock pavement and the breakers 

of the reef crest (Figure 31). On our transect we 

did not consider this "reef flat" as part of the 

crest (Goreau, 1959:74) because, although this 

zone is intertidal, it is very narrow and almost 

always flooded by waves; instead, we defined 

"reef flats" herein as large intertidal areas between the eastern shores of South Water and 

Carrie Bow cays and their nearby crests (Larson 

and Larson, herein; Riitzler, in prep.). Only the 

narrow intertidal breaker zone is included in the 

reef crest. The inner fore reef begins at this point 

with a spur and groove (= buttress) zone of high 

relief—which in Jamaica is considered a part of 

the crest (Goreau, 1959)—but it changes abruptly 

into a gently sloping terrace of spurs and grooves 

having low relief ("seaward slope" or "upper forereef terrace" in Jamaica). The outer fore reef on 

our transect has a steep inner reef slope, a perpendicular sand trough parallel to the reef crest, 

and an outer coral ridge where the fore-reef slope 

begins to drop off to the deep (vertical) fore reef. 

The comparable feature off Jamaica is a lower 

fore reef having a lower fore-reef escarpment. 

Both the trough and the ridge are missing in 

Jamaica (Goreau and Land, 1974). 

Water movement (direction and force) appear 

to control the development of zonation patterns. 

Wave action determines not only reef types, by 

influencing coral zonation (Geister, 1977), but 

also influences the distribution of all other sedentary organisms, some of which—for instance, 

octocorals, sponges, and algae—have considerable ecological importance. The quantitative significance of many coral associates is undervalued 

by most field methods (point and chain-link 

counts, estimates of area coverage) because presence or surface area is measured but not biomass 

(massive sponges) or space occupied (swaying 
NUMBER 12 41 

FIGURE 30.—Underwater views of mangrove root system at edge of Twin Cay 

channel: a, mudbank stabilized by rootlets and overgrown by Halimeda mat; b, 

juvenile barracuda finding shelter and 

food among the Rhizophora roots; c, Oreaster reticulatus on mangrove roots penetrating sediment bottom of the channel. 
FIGURE 31.—Carrie Bow reef zonation: above, oblique aerial 

view looking toward lagoon; below, block diagram of transect. 

frs 

frs = fore-reef slope 

hsg = high spur and groove 

irs = inner reef slope 

lsg = low spur and groove 

or = outer ridge 

pr = patch reefs 

re = reef crest 

rf = reef flat 

sg = seagrass 

st = sand trough 
NUMBER 12 43 

gorgonians). Objective evaluation of nonframebuilding organisms is essential when comparisons 

are made between reef biota and nearby lagoon 

habitats that are not dominated by corals, or only 

partly dominated by them. 

We documented an impressive proliferation of 

Acropora cervicornis in the back-reef and lagoon 

zones during the four years following the destruction caused by hurricane Fifi in 1974. Our spring 

1979 survey indicated that destruction associated 

with hurricane Greta (September 1978) was 

mainly the breaking up and lagoonward transportation of the shallow-water A. cervicornis. Few 

colonies of the fragile branching coral escaped 

this movement and subsequent burial in the sand 

and rubble zone as well as in the patch reef zone. 

Almost all of this coral has been washed out of 

the rubble and pavement zone (Figure 11). Although many of the living and partly buried 

fragments will, in time, develop into large colonies, the number of living A. cervicornis in shallow 

water has been drastically reduced. This constant 

cycle of vigorous development, destruction, and 

resurgence gives rise to the commonly observed 

high proportion of A. cervicornis rubble in comparison to living A. cervicornis in many shallow-reef 

areas. 

Although the branching corals Acropora palmata 

and A. cervicornis suffer extensive mechanical damage during hurricanes, the transportation and 

reestablishment of living fragments are significant 

factors in the distribution of these corals in the 

shallow-water environment (Plate 2: center left; 

Highsmith et al., 1980). Similar observations have 

been made in Florida (Shinn, 1972; Gilmore and 

Hall, 1976) and in Jamaica (Tunnicliffe, 1980) 

where the dispersal of A. cervicornis in these reefs 

was reported to be largely related to asexual 

reproduction by regeneration of broken and 

transported branches. 

Our observations and those of James and Ginsburg (1978) have documented Halimeda as a major contributor to sediment in the Belizean barrier-reef complex. This calcareous green alga 

forms a major fraction of these sediments, extending from the shallow lagoon down to at least 200 

m on the fore-reef slope, well below its living 

depth range (approximately 100 m). Commonly 

comprising the dominant component of the extremely coarse fractions (2-4 mm), the readily 

identifiable calcareous plates of Halimeda are more 

characteristic of reef-derived sediments than are 

the fragments of any other organisms, including 

the corals. 

The narrow rock pavement that occurs directly 

shoreward of the reef crest off Carrie Bow Cay 

(Figure 4) is a characteristic substrate of shallow 

reef areas that are constantly having their sediment cover swept away by turbulent waters. The 

dates of 480±90 years and 534±90 years obtained 

from coral fragments embedded in a pavement 

off South Water Cay (James et al., 1976) indicate 

a long period of formation. These dates also 

support Macintyre's (1977) observation that submarine lithification is most highly developed in 

reef areas of high agitation and/or slow accumulation, where the substrate is exposed to normal marine conditions for long periods of time. 

James and Ginsburg (1978) speculated that a 

shelf-edge ridge off South Water Cay is a submerged reef similar to the relict shallow-water, 

Late Holocene reefs described by Macintyre 

(1967, 1972), Adey et al. (1977), and Lighty et al. 

(1978). In contrast, Burke (herein) proposed that 

these ridges are active accumulations of the rapidly growing coral Acropora cervicornis. Burke points 

out that not only is A. cervicornis dominant on 

these ridges, but that the ridges along the barrier 

reef complex are restricted to areas protected 

from long-period storm waves by the outlying 

atolls. The difficulty with which we probed this 

ridge (an average penetration of 1 m, maximum 

of 1.5 m) in contrast to the ease of probing 

through Acropora cervicornis at Rhomboid Shoals 

near Victoria Channel (Macintyre et al., 1977) 

indicates that this ridge does not have a similar 

open-frame network. Our earlier observation that 

Montastrea annularis constructs most of the modern 

framework of the ridge south of our transect also 

indicates that this shelf-edge ridge is not merely 

an accumulation of A. cervicornis. Core samples of 

the internal structure are needed to establish the 
44 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

relative importance of relict and modern framework in the construction of this ridge system. 

The reef off Carrie Bow Cay has a species 

composition and community zonation that is representative of the entire barrier reef system 

(Burke, herein). The structure of shallow zones of 

this central province reef, however, is somewhat 

more similar to that of the discontinuous reefs in 

the northern and southern provinces because current flow through South Water and Carrie Bow 

cuts influences the sediment and coral distribution patterns in the back reef and lagoon. The 

Carrie Bow Cay fore-reef structures, on the other 

hand, have a degree of development and flourishing coral communities that are characteristic of 

the central province. 

Literature Cited 

Adey, W. H. 

1977. Shallow Water Holocene Bioherms of the Caribbean Sea and West Indies. In D. L. Taylor, editor, 

Proceedings, Third International Coral Reef Symposium, 

2:xxi-xxiv. Miami, Florida: Rosenstiel School of 

Marine and Atmospheric Science. 

Adey, W. H., I. G. Macintyre, and R. Stuckenrath 

1977. Relict Barrier Reef System off St. Croix: Its Implication with Respect to Late Cenozoic Coral 

Reef Development in the Western Atlantic. In D. 

L. Taylor, editor, Proceedings, Third International 

Coral Reef Symposium, 2:15-21. Miami, Florida: 

Rosenstiel School of Marine and Atmospheric 

Science. 

Antonius, A. 

1973. New Observations on Coral Destruction in Reefs 

[Abstract]. In [Abstracts of] Tenth Meeting of the 

Island Marine Laboratories of the Caribbean, page 3. 

Mayaguez, Puerto Rico: University of Puerto 

Rico. 

1977. Coral Mortality in Reefs: A Problem for Science 

and Management. In D. L. Taylor, editor, Proceedings, Third International Coral Reef Symposium, 2:617-

623. Miami, Florida: Rosenstiel School of Marine 

and Atmospheric Science. 

Burke, R. B. 

1979. Morphology, Benthic Communities, and Structure 

of the Belize Barrier Reef. 78 pages. Master's 

thesis, University of South Florida, Tampa. 

Chevalier, J. P. 

1973. Geomorphology and Geology of Coral Reefs in 

French Polynesia. In O. A. Jones and R. Endean, 

editors, Biology and Geology of Coral Reefs, 1:113-141. 

New York: Academic Press. 

Dahl, A. L. 

1973. Surface Area in Ecological Analysis: Quantification of Benthic Coral-Reef Algae. Marine Biology, 

23:239-249. 

Geister, J. 

1977. The Influence of Wave Exposure on the Ecological 

Zonation of Caribbean Coral Reefs. In D. L. 

Taylor, editor, Proceedings, Third International Coral 

Reef Symposium, 1:23-29. Miami, Florida: Rosenstiel School of Marine and Atmospheric Science. 

Gilmore, M. D., and B. R. Hall 

1976. Life History, Growth Habits, and Constructional 

Roles of Acropora cervicornis in the Patch Reef Environment. Journal of Sedimentary Petrology, 46:519-

522. 

Goreau, T. F. 

1959. The Ecology of Jamaican Coral Reefs, I: Species 

Composition and Zonation. Ecology, 40:67-90. 

Goreau, T. F., and L. S. Land 

1974. Fore-Reef Morphology and Depositional Processes, North Jamaica. In L. F. Laporte, editor, 

Reefs in Time and Space. Society of Economic Paleontologists and Mineralogists, Special Publication, 18: 

79-89. Tulsa, Oklahoma. 

Hesse, K. O. 

1979. Movement and Migration of the Queen Conch, 

Strombus gigas, in the Turks and Caicos Islands. 

Bulletin of Marine Science, 29:303-311. 

Highsmith, R. C , A. C. Riggs, and C. M. D'Antonio 

1980. Survival of Hurricane-generated Coral Fragments 

and a Disturbance Model of Reef Calcification/ 

Growth Rate. Oecologia (Berlin), 46:322-329. 

James, N. P., and R. N. Ginsburg 

1978. The Deep Seaward Margin of Belize Barrier and Atoll 

Reefs. 201 pages. Miami Beach, Florida: University 

of Miami. 

James, N. P., R. N. Ginsburg, D. S. Marszalek, and P. W. 

Choquette 

1976. Facies and Fabric Specificity of Early Subsea Cements in Shallow Belize (British Honduras) Reefs. 

Journal of Sedimentary Petrology, 46:523-544. 
NUMBER 12 45 

Lighty, R. G., I. G. Macintyre, and R. Stuckenrath 

1978. Submerged Early Holocene Barrier Reef SouthEast Florida Shelf. Nature, 275:59-60. 

Macintyre, I. G. 

1967. Submerged Coral Reefs, West Coast of Barbados, 

West Indies. Canadian Journal of Earth Sciences, 4: 

461-474. 

1972. Submerged Reefs of Eastern Caribbean. American 

Association of Petroleum Geologists Bulletin, 56:720-

738. 

1977. Distribution of Submarine Cements in a Modern 

Caribbean Fringing Reef, Galeta Point, Panama. 

Journal of Sedimentary Petrology, 47:503-516. 

Macintyre, I. G., R. B. Burke, and R. Stuckenrath 

1977. Thickest Recorded Holocene Reef Section, Isla 

Perez Core Hole, Alacran Reef, Mexico. Geology, 

5:749-754. 

Purdy, E. G. 

1974. Reef Configurations: Cause and Effect. In L. F. 

Laporte, editor, Reefs in Time and Space. Society 

of Economic Paleontologists and Mineralogists, Special 

Publication, 18:9-76. Tulsa, Oklahoma. 

Purdy, E. G., W. C. Pusey III, and K. F. Wantland 

1975. Continental Shelf of Belize—Regional Shelf Attributes. In K. F. Wantland and W. C. Pusey III, 

editors, Belize Shelf-Carbonate Sediments, Clastic 

Sediments, and Ecology. The American Association of 

Petroleum Geologists, Studies in Geology, 2:1-52. Tulsa, 

Oklahoma. 

Riitzler, K. 

1975. The Role of Burrowing Sponges in Bioerosion. 

Oecologia (Berlin), 19:203-216. 

1978a. Photogrammetry of Reef Environments by Helium Balloon. In D. R. Stoddart and R. E. Johannes, editors, Coral Reefs: Research Methods. Monographs on Oceanographic Methodology, 5:45-52. Paris: 

Unesco. 

1978b. Sponges in Coral Reefs. In D. R. Stoddart and R. 

E. Johannes, editors, Coral Reefs: Research Methods. Monographs on Oceanographic Methodology, 5:299-

313. Paris: Unesco. 

1981. An Unusual Blue-Green Alga Symbiotic with Two 

New Species of Ulosa (Porifera: Hymeniacidonidae). Marine Ecology, 2:35-50. 

In prep. The Atlantic Barrier Reef Ecosystem at Carrie 

Bow Cay, Belize: The Reef Flat Community. 

Riitzler, K., J. D. Ferraris, and R. J. Larson 

1980. A New Plankton Sampler for Coral Reefs. Marine 

Ecology, 1:65-71. 

Shinn, E. A. 

1972. Coral Reef Recovery in Florida and the Persian Gulf. 9 

pages. Houston, Texas: Conservation Department, Shell Oil Company. 

Smith, F.G.W. 

1948. A Handbook of the Common Atlantic Reef and ShallowWater Corals of Bermuda, the Bahamas, Florida, the 

West Indies, and Brazil. 112 pages. Coral Gables, 

Florida: University of Miami Press. 

Stoddart, D. R. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays, October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 

Tunnicliffe, V. Z. 

1980. Biological and Physical Processes Affecting the 

Survival of a Stony Coral, Acropora cervicornis. 

Ph.D. dissertation, Yale University, New Haven, 

Connecticut. 

Wantland, K. F., and W. C. Pusey III 

1971. A Guidebook for the Field Trip to the Southern Shelf of 

British Honduras. New Orleans, Louisiana: New 

Orleans Geological Society. 
Tides at Carrie Bow Cay, Belize 

Bjorn Kjerfve, Klaus Riitzler, 

and George H. Kierspe 

ABSTRACT 

The tide at Carrie Bow Cay, Belize, is microtidal (mean range of 15 cm) and is of the mixed 

semidiurnal type. Comparison with conditions at 

Key West, Florida, indicates that high and low 

waters off Carrie Bow occur earlier than at Key 

West by 45 and 2 minutes, respectively. Because 

of differences in tidal type and meteorological 

conditions, corrections for height and time difference applied to the predicted tide at Key West 

yield only approximate tide predictions for Carrie 

Bow Cay. 

Introduction 

Because the tide in the Caribbean Sea is microtidal, it might be expected to have little influence 

on the water flow regime. Velocity measurements 

indicate, on the contrary, that tidal forcing is a 

major cause of currents in coastal regions of the 

Caribbean (Roberts et al., 1975). Study of a 

shallow reef flat and back reef shows that even 

small tidal fluctuations have strong influence on 

the distribution and succession of organisms 

(Glynn, 1973; Riitzler, in prep.). This note describes, characterizes, and predicts the tide at 

Carrie Bow Cay (16°48'N, 88°05'W), Belize, as 

a necessary first step in the investigation of flow 

and water exchange characteristics as well as 

Bjorn Kjerfve, Marine Science Program, Department of Geology, and 

the Belle W. Baruch Institute for Marine Biology and Coastal 

Research, University of South Carolina, Columbia, S. C. 29208. 

Klaus Riitzler, Department of Invertebrate Zoology, National Museum 

of Natural History, Smithsonian Institution, Washington, 

D.C 20560. George H. Kierspe, Harbor Branch Foundation, Inc., 

Fort Pierce, Fla. 33450. 

intertidal and shallow subtidal communities at 

this barrier reef location. 

ACKNOWLEDGMENTS.—Many persons helped to 

collect the tide data used in this study. We are 

especially grateful to J. D. Ferraris, Mount Desert 

Island Biological Laboratory, R. Larson and M. 

Carpenter, Smithsonian Institution, and J. Greer, 

R. L. Crout, S. Ferguson, and G. Pickler, University of South Carolina. J. E. Fancher and D. C. 

Simpson of the National Ocean Survey provided 

valuable advice on the analytical procedures. L. 

Kjerfve did the drafting. All computer work was 

performed on the University of South Carolina 

IBM 370/168 system. The Belle W. Baruch Institute for Marine Biology and Coastal Research 

has listed this paper as its Contribution Number 

269. 

Measurements, Analyses, and Results 

A Benthos 2820 submergible in situ tide recorder, which senses the hydrostatic pressure, was 

installed below the pier on the west side of Carrie 

Bow Cay in the barrier reef lagoon. This location 

is 120 m west of the reef crest, 24 km southeast of 

Dangriga (Stann Creek). Two major navigation 

cuts through the barrier reef, less than 1 km away, 

connect the barrier reef lagoon to the Caribbean 

Sea. The gauge intake was, on the average, 33 cm 

off the bottom, 59 cm below water, 153 cm below 

the top of the cement dock (the local reference 

datum) and has been operated intermittently 

since early 1976. 

Several 29-day tide records were digitized to 

hourly intervals and subjected to harmonic anal47 
48 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Carrie Bow Cay Tide 

(MSL) 

FIGURE 32.—Comparison of measured and computer-predicted tides at Carrie Bow Cay, 

Belize, 10 April-8 May 1976. 

ysis (Schureman, 1940; Dennis and Long, 1971) 

for the purpose of computing amplitudes and 

epochs for the 24 major tidal constituents, which 

correspond to more than 99% of the actual amplitude. The nine constituents with the greatest 

amplitudes for a typical 29-day series are compared in Table 3 with the same constituents at 

Key West, Florida. The 24 amplitude and epoch 

values for Carrie Bow Cay were entered into the 

National Ocean Survey tide prediction computer 

program (Pore and Cummings, 1967), which allows monthly prediction of both hourly tidal 

height values and times and heights of high and 

low water. Results indicated reasonable agreement between measured and predicted tides for 

a 29-day test series (Figure 32). The prediction of 

time of high and low water is in general more 

successful than reproduction of water elevation. 

The 29-day test record, 10 April-8 May 1976, 

was also tabulated and subjected to high-and-low 

water analysis on the basis of instructions for 

Form 2211 (National Ocean Survey, 1974:44-59). 

TABLE 3.—Comparison of values for the nine major tidal constituents at Carrie Bow Cay, 

Belize, and Key West, Florida (symbol in parentheses follows each constituent; columns 1 and 

2 computed for Carrie Bow Cay using Program "Harmonic" (Dennis and Long, 1971); columns 

3 and 4 supplied by D. Simpson, Predictions Branch, National Ocean Survery; columns 5 and 

6 from Defant, 1960: 267) 

Tidal constituents 

SEMIDIURNAL COMPONENTS 

Principal lunar 

Principal solar 

Larger lunar elliptic 

Luni-solar 

Larger lunar evectional 

DIURNAL COMPONENTS 

Luni-solar 

Principal solar 

Principal lunar 

Larger lunar elliptic 

(M2) 

(&) 

(N2) 

(K2) 

("2) 

(*i) 

(A) 

(Oi) 

(Oj) 

Carrie Bow 

Amplitude 

(cm) 

5.7 

3.5 

2.5 

1.0 

0.5 

7.9 

2.6 

2.5 

0.5 

Cay 

Epoch 

C) 

251 

228 

243 

228 

244 

185 

185 

244 

273 

Key 

Amplitude 

(cm) 

17.8 

5.4 

3.5 

1.5 

0.7 

8.9 

2.9 

9.2 

2.2 

West 

Epoch 

C) 

285 

302 

269 

302 

271 

282 

287 

284 

275 

Period 

(mean solar 

hour) 

12.42 

12.00 

12.66 

11.97 

12.63 

23.93 

24.07 

25.82 

26.87 

Theoretical 

coefficient 

ratio (M2 = 

100) 

100.0 

46.6 

19.2 

12.7 

3.6 

58.4 

19.4 

41.5 

7.9 
NUMBER 12 49 

The pertinent results of this analysis are summarized in Table 4. 

Discussion 

The tide at Carrie Bow Cay is microtidal and 

of the mixed semidiurnal type. Its mean range is 

15 cm and its semidiurnal and diurnal amplitudes 

are of approximately equal importance (Table 3). 

The form number F, an amplitude ratio between 

harmonic constituents, may be used to quantify 

the tide type (Defant, 1960:306-308). It is defined 

by F = (Ki + Ox)/{M2 + S2), where Kx is the 

diurnal luni-solar, 0\ the diurnal principal lunar, 

Mi the semidiurnal principal lunar, and Si the 

semidiurnal principal solar component (Table 3). 

If F< 0.25 the tide is semidiurnal; if 0.25 <F< 

1.50 the tide is mixed semidiurnal; if 1.50 < F< 

3.0 the tide is mixed diurnal; and F > 3.0 the 

tide is diurnal. The Carrie Bow Cay form number 

is 1.13 in comparison with 0.75 at Key West, 

Florida. Although both locations have the same 

tide type, the diurnal influence is greater at Carrie 

Bow Cay. 

From the amplitude of the various harmonics 

(Table 3) it is possible to compute additional 

statistics (Marmer, 1954). With respect to the 

semidiurnal tidal constituents, the spring tide 

range is 2{M2 + S2) or 18.4 cm and the neap 

range approximately seven days later of 2(M2 — 

S2) or 4.4 cm. The mean semidiurnal tide is 2.2 

M2 or 12.5 cm. With respect to the diurnal constituents, the tropic tide range measures 2(K\ + 

Oi) or 20.8 cm, the equatorial range is 2(K\ — Oi) 

or 5.4 cm, and the mean diurnal tide is \.b(K\ — 

Oi) or 15.6 cm. The values above are only approximate as the P\ and N2 constituents show 

values significantly larger than could have been 

expected from the theoretical coefficient ratio 

based on the magnitude of the constituents' tideproducing forces (Table 3) at this location. Of 

course, such discrepancies between the magnitude 

of the tide-producing force and actual response 

of the water mass is quite common and is due to 

basin resonance characteristics. 

The diurnal and semidiurnal partial tides are 

approximately equal; however, the spring-neapspring cycle of the semidiurnal tide is 29.5 mean 

solar days (synodic month) and is related to the 

phase of the moon. The tropic-equatorial-tropic 

cycle of the diurnal tide is somewhat shorter, 27.3 

mean solar days (sidereal month), and is related 

to the declination of the moon from the equator. 

Because of this time difference, several longer 

cycles are introduced. If the occasional times of 

high water in M2, S2, N2, K\, P\, and Oi occur 

simultaneously, the total tide range could be as 

great as 50 cm. Of course, wind tides are likely to 

cause extreme sea level changes more so than the 

astronomical forces. 

The epoch or phase relative to the Greenwich 

meridian yields the following information about 

the inequality of the timing of highs and lows in 

the component tides (Marmer, 1949). If epochs 

are expressed in degrees, the difference D = 

\M°2-{K0iA- 0°i)\ indicates whether tidal inequality is entirely in the high waters {D ~ 0°), 

is equally great in the high and low waters {D ~ 

90°), or is entirely in the low waters {D ~ 180°). 

If D is greater than 180°, its value is subtracted 

from 360°. For Carrie Bow Cay, the D value of 

120° indicates inequality in both high and low 

waters, which is especially pronounced in the low 

water elevations (refer to Figure 32). This socalled diurnal inequality on the average measures 

5.6 cm between low waters and 2.4 cm between 

high waters (Table 4). 

The epoch also yields information about the 

phase age and the diurnal age. The phase age 

TABLE 4.—Tide statistics for Carrie Bow Cay, corrected for 

the longitude of the moon's node and changes in the declination of the sun, based on computations using National 

Ocean Survey Form 2211 for 10 April-8 May 1976 (refer to 

Figure 32) 

Tide datum (mean low water)1 

Mean range (Mn) 

Mean tide level (MTL)2 

Diurnal high water inequality (DHQ) 

Diurnal low water inequality (DLQ) 

Greenwich lunitidal high water interval 

Greenwich lunitidal low water interval 

0 cm 

15.0 cm 

7.5 cm 

2.4 cm 

5.6 cm 

2.14 h 

8.44 h 

1

 101 cm below local reference datum. 

2

 Marked "0" on Figure 32, right hand scale. 
50 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 33.—Co-tidal lines of the predominant tides for the Caribbean Sea (after Defant, 1960); 

tide regression indicated by relative arrival time of high or low water, expressed in lunar hours: 

a, semidiurnal tide, M2 constituent, progresses along the Belizean coast from north to south; b, 

diurnal tide, K\ constituent, progresses along the Belizean coast from south to north. (Carrie 

Bow Cay indicated by arrow). 

refers to the semidiurnal tide and is the lag of the 

spring tide relative to full or new moon. The 

phase age is computed as 0.98 {S2 — M2) and 

equals —22 hours for Carrie Bow Cay, which 

indicates that spring tide leads the new and full 

moon by 22 hours. The diurnal age, on the other 

hand, is a measure of the timing of tropic tide 

relative to maximum declination of the moon. It 

is computed as 0.91 (A"? — 0?) and is 0 hours for 

Carrie Bow Cay, indicating a maximum diurnal 

tide range at the time of maximum lunar declination. 

Figure 33, which shows the main Caribbean 

semidiurnal {M2) and diurnal {K\) amphidromic 

systems (Defant, 1960) indicates that the M2 tide 

progresses from north to south along the Belizean 

barrier reef, whereas the K\ tide progresses in the 

opposite direction, from south to north, along the 

reef crest. 

Because it is inconvenient, though possible, to 

publish tidal predictions for Carrie Bow Cay, we 

instead computed correction factors and applied 

them to predicted tides for Key West, Florida, a 

National Ocean Survey reference gauge for which 

daily predictions are published. The Greenwich 

lunitidal intervals at Key West are 2.89 hours for 

high water (HW) and 8.48 hours for low water 

(LW) (D. Simpson, pers. comm.). Comparison of 

these figures with the lunitidal intervals for Carrie 

Bow (Table 4) indicates that on the average HW 

at Carrie Bow Cay leads HW at Key West by 45 

minutes, and LW at Carrie Bow Cay leads LW 

at Key West by 2 minutes. Using the predicted 

tides for Key West for April and May 1976, we 

also found that HW at Carrie Bow Cay is 23 cm 

below HW at Key West and that LW at Carrie 

Bow Cay is 4 cm above LW at Key West on the 

average. However, because Carrie Bow Cay and 

Key West can be expected to experience different 

meteorological and oceanographic conditions at 

given times, the resulting tide predictions for 

Carrie Bow Cay now and then will deviate significantly from actual tide variations. 
NUMBER 12 51 

Literature Cited 

Defant, A. 

1960. Physical Oceanography. Volume 2, 598 pages. London: Pergamon Press. 

Dennis, R. E., and E. E. Long 

1971. A User's Guide to a Computer Program for Harmonic Analysis of Data at Tidal Frequency. National Oceanic and Atmospheric Administration, Technical 

Report, 4L: 31 pages. Rockville, Maryland. 

Glynn, P. W. 

1973. Ecology of a Carribbean Coral Reef: The Porites 

Reef-Flat Biotope, Part I: Meteorology and Hydrography. Marine Biology, 20:297-318. 

Marmer, H. A. 

1954. Tides and Sea Level in the Gulf of Mexico. Fishery 

Bulletin, 55(89): 101-118. 

National Ocean Survey 

1974. Manual of Tide Observations. National Oceanic and 

Atmospheric Administration Publication, 30-1: 72 

pages. Washington, D.C. 

Pore, N. A., and R. A. Cummings 

1967. A Fortran Program for the Calculation of Hourly 

Values of Astronomical Tide and Height of High 

and Low Water. United States Weather Bureau [now: 

National Weather Service] Technical Memorandum, 

TDL-6: 17 pages. Silver Spring, Md. 

Roberts, H. H., S. P. Murray, and J. N. Suhayda 

1975. Physical Processes in a Fringing Reef System. 

Journal of Marine Research, 33 (2): 233-260. 

Schureman, P. 

1940. Manual of Harmonic Analysis and Prediction of 

Tides. United Slates Coast and Geodetic Survey, Special 

Publication, 98: 31 pages. Washington, D.C. 

Riitzler, K. 

In prep. The Atlantic Barrier Reef Ecosystem at Carrie 

Bow Cay, Belize: The Reef Flat Community. 
Water Currents Adjacent to Carrie Bow Cay, Belize 

Jeffrey E. Greer 

and Bjorn Kjerfve 

ABSTRACT 

Tide, wind, and waves significantly influence 

the water currents off Belize, as indicated by 

continuous current-meter measurements at four 

locations in or behind the barrier reef at Carrie 

Bow Cay. Tidal currents reaching speeds of up to 

40 cm s~ are dominant in the major reef entrances. In the lagoon behind the reef crest, the 

water is primarily wind driven, without an obvious tidal signature, and reaches maximum 

speeds of 33 cm s_1. On the leeward side of Carrie 

Bow Cay a slow wave drift accounts for the 

maximum 6 cm s_1 flow. During the late part of 

the study, a tropical depression forming just east 

of Carrie Bow Cay caused an abnormally high 

mean tide that increased the flow of water into 

the lagoon. 

Introduction 

Coral reef ecosystems are controlled by the 

complex interaction of biological, chemical, geological, and physical parameters (Odum and 

Odum, 1955), of which the least understood are 

probably the physical forces—waves, winds, tides 

(see for example Macintyre et al., 1974). Early 

studies of the physical oceanography of reefs focused primarily on Pacific atolls (Munk and Sargent, 1948; von Arx, 1948) and paid considerably 

less attention to coral reefs in the Caribbean. 

Jeffrey E. Greer, Marine Science Program, University of South 

Carolina, Columbia, S.C. 29208. Bjorn Kjerfve, Marine Science 

Program, Department of Geology, and the Belle W. Baruch Institute 

for Marine Biology and Coastal Research, University of South Carolina, Columbia, S.C. 29208. 

Although recent investigations have increasingly 

shifted to the Caribbean, mainly because of proximity for American workers, physical oceanographic investigations of the well-developed barrier reef of Belize are only beginning. Previous 

physical research in this area was concerned with 

waves and wave-related processes (Roberts, 1974; 

Roberts et al., 1975; Shinn, 1963; Storr, 1964; 

Wilson et al., 1973), so that the interaction of 

tides with other physical processes has received 

little attention. Recent work on Grand Cayman 

(Roberts et al., 1977) suggests strong tidal influence on physical processes within the shallow 

fore-reef region. Because the tide in the Caribbean is microtidal and of mixed semidiurnal type 

(Kjerfve et al., herein), it might be expected to 

have considerably less influence on the reef system 

than wind-driven currents and waves. Our study 

around Carrie Bow Cay indicates that just the 

opposite may be true—that is, tidal processes may 

play a major role in controlling the water movements within the reef system. 

ACKNOWLEDGMENTS.—We are grateful for the 

valuable field assistance given by L. Greczy, 

Louisiana State University; J. A. Proehl and F. 

M. Drescher, the University of South Carolina; 

and especially to L. Kjerfve. Vice consul J. Walsh 

of the Unites States Embassy in Mexico City and 

United States Senator S. Thurmond of South 

Carolina gave us invaluable logistic assistance 

during the 1978 Belize trip, which allowed us to 

complete the study. The Belle W. Baruch Institute for Marine Biology and Coastal Research 

has listed this paper as its Contribution Number 

270. 

53 
54 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Site, Measurements, and Analysis 

The barrier reef off Belize, which extends in a 

north-south direction for over 200 km, is separated from the mainland by a lagoon that is 10-

25 km wide and 5-20 m deep. The study area, 

Carrie Bow Cay (16°48'N, 88°05'W), a small, 

sandy cay approximately 120 m long and 30 m 

wide, lies 100 m west of a well-developed section 

of the barrier reef crest (Figures 34, 40). Two 

large cuts, 12 m deep and approximately 1 and 

2 km wide, respectively, separate the Carrie Bow 

reef section from Tobacco Reef to the north and 

Gladden Cay Reef to the south (Figure 2). Carrie 

Bow receives steady northeasterly trade winds 

and has a mixed semidiurnal tide with a mean 

range of 15 cm. 

Current, tide, wind, and atmospheric pressure 

were measured in June 1978. A Bendix Q-15 

recording current meter suspended from a 

^ \*. 

• 

LT 

L2, L4 • 

South Water 

fm^\ 

J- jronsect 

Carrie Bow 

L

3 * 88' 

Cut 

04.7' W 

Cut 

• 

t 

N 

0 100 

meters 

6°48.l ' Ndense coral 

Thalassia 

FIGURE 34.—Location of current meter stations around Carrie Bow Cay. 

moored boat was monitored at five locations (Figure 34) in the vicinity of Carrie Bow Cay for time 

periods ranging from 17 to 74 hours. At each 

location the Q-15 was positioned at mid-depth. 

At location LI it was placed in the lagoon 125 m 

WSW of the island, where the total water depth 

was 4 m (duration, 24 hours); at locations L2 and 

L4 it was placed in South Water Cut above a 

large, flat sand plain in depths of 11 m (duration, 

43 and 44 hours, respectively); at location L4 it 

was in Carrie Bow Cut above a 10 m deep sand 

channel (duration, 74 hours); and at location L5 

it was in the lagoon 150 m W of the main study 

transect (Riitzler and Macintyre, herein), where 

total water depth was 3 m (duration, 17 hours). 

Each record of current speed and direction was 

digitized at 10 min intervals, resolved into E-W 

and N-S components, and then averaged vectorially to obtain resultant hourly speeds and directions. The time-series plots of the data obtained 

in the South Water Cut and Carrie Bow Cut are 

shown in Figures 35 and 36. Current records for 

stations LI and L5 are presented as progressive 

vector diagrams in Figure 37. 

A Benthos 2820 submergible recording tide 

gauge was installed below the dock on Carrie 

Bow Cay to obtain the complete tidal record 

during the study (Figure 38). Wind speed and 

direction were read on a cup anemometer with 

vane every hour on the hour; readings were averaged by eye for 30 s to smooth high-frequency 

noise. The wind data are plotted on a frequency 

isopleth in Figure 39. The atmospheric pressure 

was measured using a Weather-Measure B211 

recording microbarograph. 

Results and Discussion 

Northeasterly winds with speeds of 4-5 m s~ 

were most frequent during the study period. 

These conditions seem to prevail during 70% of 

the year (Riitzler and Ferraris, herein). Sustained 

winds did not exceed 9 m s_1 and the wind 

direction varied little from northeast. During the 

latter part of the study (16-20 June), the average 

wind speed increased slightly, to 6.5 m s-1, and 

water levels became abnormally high. The in-
NUMBER 12 55 

South Water Cut (L2, L4) Current 

20 4 

20 Jun 

FIGURE 35.—Time-series of current speed and direction in South Water Cut at current meter 

locations L2 and L4, using vectorially averaged hourly values. 

50-

4 0 • 

3 0 -

2 0-

10-

0-

^Direction 

A i i 1 1 

Carrie Bow Cut(L3) Current 

Speed 

. _//, 

360 

288 

•216 

• 144 

•72 

16 0 

16 Jun 

8 0 

17 Jun 

16 0 

18 Jun 

FIGURE 36.—Time-series of current speed and direction in Carrie Bow Cut at current meter 

Location L3, using vectorially averaged hourly values. 

1100 09 Jun 

Meter Location LI 

1200 08 Jun 

I 0 I 

East-West Distance (km) 

FIGURE 37.—Progressive vector diagram of currents at current meter locations LI and L5, using vectorially averaged 

hourly values. 

creased wind speed and water level were probably 

related to a tropical depression that was developing east of Glover's Reef. The atmospheric 

pressure showed a net drop of 6 mb from 15 to 20 

June. 

The predominant flow feature observed in the 

major reef cuts was a strong tidal periodicity with 

flood currents greatly exceeding the ebb currents. 

Maximum current velocities in South Water Cut 

were 26 cm s"1 and 43 cm s_1 during the first (L2) 

and second (L4) study periods, respectively. In 
56 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Carrie Bow Cay Tide 

FIGURE 38.—Measured tide at Carrie Bow Cay dock, 8-22 June 1978; numbered segments 

indicate water elevations when Q-15 current meter was in operation at locations L1-L5. 

W— 

Density: 2.07 1.49 

Frequency (%) 

FIGURE 39.—Frequency isopleth diagram (modified from 

Seppala, 1977) of 312 hourly recordings of wind data at 

Carrie Bow Cay, Belize, 8-22 June 1978, indicating density 

of wind observations (in observations per m s_1 and 15° 

segments) and percentage of frequency corresponding to a 

given direction. 

Carrie Bow Cut, observed hourly peak velocities 

measured 40 cm s- . Directional changes during 

each half tide (rising or falling) were negligible in 

both South Water and Carrie Bow cuts. In South 

Water Cut the average resultant direction for 

inflow was toward 260°, and for outflow toward 

70°. In Carrie Bow Cut the flood flow was toward 

275° and the ebb flow toward 105°. The predominance of the flooding tide over the ebb during 

the study period indicates a slow, continual infilling of the lagoon, which is also indicated by 

data for the mean increase of the tide level (Figure 

38). The tidal pattern was due to the stationary 

tropical depression to the east of Carrie Bow, 

which had strengthened the northeast trade 

winds. 

Current patterns observed at lagoon stations 

LI and L5 are shown in Figure 39 on a progressive 

vector diagram. Neither location exhibited the 

obvious tidal activity that had occurred at stations L2, L3, and L4. Meter location LI showed 

weak water movement and little effects of tidal 

or wind forcing. Hourly averaged currents at LI 

did not exceed 6 cm s~ and were generally on 

the order of 2 cm s_1. This location, being on the 

lee side of the island, was shaded from direct 

wind influence. The net flow direction to the 

northeast at LI suggests that the current was due 

to wave drift. The wave crests that propagated 

into Carrie Bow Cut were greatly refracted by 

the reef crest and entered the Carrie Bow lagoon 

from the southwest, as indicated by waves break-
NUMBER 12 57 

FIGURE 40.—Vertical high altitude aerial photograph of Carrie Bow Cay (center) showing wave 

refraction patterns; island to the north (top) is South Water Cay, sand bank to the south 

(bottom) is Curlew Bank (picture area = 2.5 X 3.5 km). 

ing on the lee side of the island and refraction 

patterns on aerial photographs (Figure 40). 

Although higher water levels and increased 

wind speeds due to the developing tropical depression were atypical conditions for station L5, 

these conditions probably did not alter the typical 

flow patterns there, but merely modulated their 

intensity. Maximum hourly velocities at L5 were 

33 cm s_1, with a net flow direction due southwest, 

varying almost 180° from that at LI. The current 
58 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

at L5 did not have the strong tidal signature 

evident in the major reef cuts. Rather, water was 

being driven across the reef crest and into the 

lagoon by the northeasterly trade winds. While 

the higher than usual water levels and increased 

wind speeds tended to move more water across 

the crest than usual, the current at L5 appeared 

to be primarily wind-driven. 

Conclusions 

Current flow through the major reef cuts bordering Carrie Bow Cay appears to be tidally 

dominated despite a small tidal range. Measurements within the cuts showed a predominance of 

flooding over ebbing currents that was related to 

increased northeasterly winds associated with the 

development of a tropical depression to the east 

of the island. Depending on location, net currents 

inside the lagoon may be wave-driven, winddriven, or both. Current records from this area 

lack the clear tidal signature that was observed 

in Carrie Bow and South Water cuts. Because 

our study was conducted in the early summer, 

however, caution must be exercised in extrapolating the results to other seasons. 

Literature Cited 

Arx, W. S. von 

1948. Circulation Systems of Bikini and Rongelap Lagoons. Transactions, American Geophysical Union, 

29(6) :861-871. 

Macintyre, I. G., S. V. Smith, and J. C. Zieman, Jr. 

1974. Carbon Flux through a Coral Reef Ecosystem: A 

Conceptual Model. Journal of Geology, 82:161-171. 

Munk, W. H., and M. C. Sargent 

1948. Adjustment of Bikini Atoll to Ocean Waves. Transactions, American Geophysical Union, 29 (6):855-860. 

Odum, H. I., and E. P. Odum 

1955. Trophic Structure and Productivity of a Windward Coral Reef Community on Eniwetok Atoll. 

Ecological Monographs, 25:291-320. 

Roberts, H. H. 

1974. Variability of Reefs with Regard to Changes in 

Wave Power around an Island. In A. M. Cameron, 

B. M. Campbell, A. B. Cribb, R. Endean, J. S. 

Jell, A. O. Jones, P. Mather, and F. H. Talbot, 

editors, Proceedings, Second International Symposium on 

Coral Reefs, 2:497-512. Brisbane, Australia: Great 

Barrier Reef Committee. 

Roberts, H. H., S. P. Murray, and J. H. Suhayda 

1975. Physical Processes in a Fringing Reef System. 

Journal of Marine Research, 33(2): 233-260. 

1977. Physical Processes in a Fore-Reef Shelf Environment. In D. L. Taylor, editor, Proceedings, Third 

International Coral Reef Symposium, 2:507-516. 

Miami, Florida: Rosenstiel School of Marine and 

Atmospheric Science. 

Seppala, M. 

1977. Frequency Isopleth Diagram to Illustrate Wind 

Observations. Weather, 32(5): 171-175. 

Shinn, E. A. 

1963. Formation of Spurs and Grooves on the Florida 

ReefTract. Journal of Sedimentary Petrology, 33:291-

303. 

Storr,J. R. 

1964. Ecology and Oceanography of the Coral Reef 

Tract, Abaco Island, Bahamas. Geological Society of 

America Special Paper, 79: 98 pages 

von Arx. See Arx, W. S. von 

Wilson, W. S., P. G. Wilson, and J. A. Michael 

1973. Analysis of Swell near the Island of Aruba. Journal 

of Geophysical Research, 78:7834-7844. 
Water Exchange across the Reef Crest 

at Carrie Bow Cay, Belize 

Bjorn Kjerfve 

ABSTRACT 

Water movement across the reef crest off Carrie 

Bow Cay, Belize, is affected by at least five microgrooves, 2-4 m wide and 1-3 m deep, cutting 

across the reef crest and connecting the shallow 

reef lagoon and the open Caribbean Sea. The 

currents in these microgrooves flow into the lagoon during rising tides and out to sea during 

falling tide. Over one or more tidal cycles, however, net flow inside the microgrooves is directed 

toward the ocean and along the remainder of the 

reef crest it is directed toward the lagoon owing 

to water being pumped over the crest by breaking 

waves. 

Introduction 

Few environments are as hazardous or as difficult to sample as the reef crest on an exposed 

coast. Breaking waves that pound the outside 

edge of the reef dissipate vast quantities of energy, 

and cause water to be pumped across the crest. 

In addition, the water flow over the reef is typically influenced by tidal currents, hydraulic head 

differences (Tait, 1972), and persistent 4-8 m s~ , 

trade winds. Few studies have examined this 

complex of water motions. 

Munk and Sargent (1948) noted that "surge 

channels," cutting across the reef crest and reef 

flat on the windward side of Pacific atolls, are 

tuned to the average wave characteristics and 

Bjorn Kjerfve, Marine Science Program, Department of Geology, and 

the Belle W. Baruch Institute for Marine Biology and Coastal 

Research, University of South Carolina, Columbia, S. C. 29208. 

play an important role in the dissipation of wave 

energy. Although Roberts et al. (1975) found no 

evidence of surge channels in exposed fringing 

reefs around Grand Cayman and Barbados, a 

section of the Belizean barrier reef at Carrie Bow 

Cay in the western Caribbean (16°48'N, 

88°05'W) has at least five channels cutting across 

the reef crest. Because of their small size and 

other characteristics that differ from the atoll 

surge channels, the Carrie Bow channels are 

called "microgrooves" in the following description of their role in the water exchange across the 

reef crest. 

ACKNOWLEDGMENTS.—For help with the collection of field data in 1977 and 1978 I am especially 

indebted to J. B. Atkins, F. M. Drescher, J. E. 

Greer, and J. A. Proehl, University of South 

Carolina; L. Greczy, Louisiana State University; 

and to L. Kjerfve. J. Stallings, United States 

Geological Survey, and D. Middaugh, Environmental Protection Agency, donated Rhodamine 

dye and practice golf balls, respectively, for numerous qualitative flow experiments. The Belle 

W. Baruch Institute for Marine Biology and 

Coastal Research has listed this paper as its contribution Number 271. 

Results 

MICROGROOVES.—The Carrie Bow section of 

the barrier reef lies just off the south end of 

Tobacco Reef. It is separated from South Water 

Cay by a cut, 800 m wide and 12 m deep. The 

reef crest forms a half-ellipse 1000 m long (Figure 

41) that consists of dense growths of Acropora 

59 
60 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Reef Crest Microgrooves 

-16 

88 04.7 W 

FIGURE 41.—Schematic representation of the Carrie Bow 

Cay reef crest, indicating location of five microgrooves (AE), refraction of typical wave crests before breaking, (curved 

lines) and direction of predominant trade winds and deep 

water waves (arrows). (Scale in m.) 

palmata, Millepora complanata, Agaricia agaricites, 

Porites astreoides, and other corals. The elevation 

at the crest coincides approximately with the 

mean low water level. The crest measures 20-35 

m across from a 1 m deep sand-filled moat on its 

lagoon side to another 4 m deep sand moat on its 

seaward side. 

The lagoonward edge of the reef crest is irregularly indented and gives the impression that 

numerous channels open into the ocean. Most of 

these cuts, however, extend only part of the way 

across the crest, except for five locations (Figure 

41) where the channels traverse the crest and thus 

make it possible for a swimmer to reach the fore 

reef even during moderate wave conditions. 

These small passages are the microgrooves (2-4 

m wide and 1-3 m deep) that wind their way 

across the crest; they typically exhibit a constricted area somewhere between the lagoon and 

ocean where the water becomes significantly shallow compared with the rest of the microgroove. 

The bottom of the microgrooves consists of poorly 

sorted carbonate sand and large amounts of broken coral branches and rubble. Large numbers of 

schooling fish appear to be associated with the 

microgrooves. Microgroove E (Figure 41) across 

the well-developed northern section of the reef at 

Carrie Bow Cay is by far the deepest and widest 

one. 

The microgrooves differ from atoll surge channels (Munk and Sargent, 1948) in several ways: 

they are shallower and narrower; they extend 

across the reef crest with several bends and 

branchings; they usually do not align with a forereef groove; they appear less well tuned to the 

predominant waves. These differences, however, 

may all be due to the calmer wave climate in the 

Caribbean. Whereas the trade-wind-generated 

waves on Bikini Atoll are 2.0-2.5 m high with 

steady 7-9 s periods, the Carrie Bow reef usually 

experiences 1.0-1.5 m, 4-5 s waves with a great 

variability over time. 

WATER EXCHANGE.—As wave crests progress 

toward a reef they shoal and refract. Before waves 

reach the breaking point, refraction causes their 

crests to become almost parallel to the outer reef 

edge (Figure 41). When a wave breaks, water is 

pumped across the reef crest (Shinn, 1963) and 

some water surges back before the arrival of the 

next breaker. The available wave energy is transformed into kinetic energy and is dissipated, and 

thus helps to maintain a superelevated water level 

at the breaking point, in contrast to the still-water 

levels of the lagoon (Munk and Sargent, 1948). 

This reef set-up may be as much as 20% of the 

incident wave height (Tait, 1972). If the waves 

are high, the set-up can drive over the reef crest 

a steady net inflow, which becomes superimposed 

on the more or less symmetrical, oscillatory 

pumping and return-surge action due to breakers. 

Because of refraction, waves usually traverse the 

barrier perpendicular to the local crest axis and 
NUMBER 12 61 

along the mean direction of the microgrooves. As 

in the case of atoll surge channels, this pattern 

suggests a close relationship between microgrooves and wave action. 

Qualitative features of the reef-crest water exchange were assessed by means of current-meter 

measurements and dye drops during March 1977 

and June 1978. A Bendix B-10 impeller-type, 

ducted, bidirectional current meter was placed at 

approximately middepth in microgrooves A, B, 

and C, as well as in two locations on the reef crest 

between microgrooves B and C. At each location 

two 25 h time series were obtained via a 200 m 

cable connecting the sensor and a shore-based 

read-out unit and recorder. The signal was subjected to a 2.5 s RC low-pass filter, and was then 

read every 5 s for 3 min on the hour every hour. 

Statistically significant hourly mean values based 

on 36 readings were plotted versus time, and 

compared with the tide for the same period. The 

resulting plot for microgroove B is shown in Figure 42, that for the reef crest in Figure 43. In 

" 15 

10 

o 

9 " 

6 " 

3 -

U 

-6 ' 

Flow 

7* ^ 

* \ 

\ / 

In Microgroove B 

Tide 

Toward Ocean \ 

f\ ' ^ 

/ ' * f l /

 V 

1A / \ \ 1 \ i \ J \ 

/ \ / \ i \ 1 AV /• 

' \ / Current \j 

Toward Lagoon 

i i i i i i 

o 

-10 

12 00 

15 June 16 June 

FIGURE 42.—A 25 hour time series of low-pass filtered current 

in microgroove B, 15-16 June 1978, with simultaneous tide 

record at the Carrie Bow dock (positive current = lagoonWard flow, negative current = oceanward flow). 

-10 

"-2 0 

19 June 

FIGURE 43.—A 25 hour time series of the low-pass filtered 

current on top of the reef crest between microgrooves B and 

C, 19-20 June 1978, with simultaneous tide record at the 

Carrie Bow dock (positive current = lagoonward flow, negative current = oceanward flow). 

addition, Rhodamine B dye was released at approximately 50 locations along the Carrie Bow 

section of the barrier, in front of the reef, in the 

lagoon, on top of the crest, and in all microgrooves 

during both falling and rising tides. In addition, 

water movement was traced by means of 200 

plastic practice golf balls released at several locations oceanward of the reef and observed as 

they crossed the reef crest; these were subsequently collected. 

Conclusions 

Although these experiments yielded primarily 

qualitative results, several generalizations can be 

made. Currents in microgrooves as well as on top 

of the reef crest varied on at least two time scales: 
62 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

wave and tide frequencies. The wave-induced 

motions were largely oscillatory and perpendicular (back and forth, up and down) to the crest 

with typical instantaneous microgroove speeds of 

10-60 cm s_1. The hourly means, however, reflect 

the tidal forcing and were typically 1-20 cm s_1 

m in the microgrooves, directed oceanward (negative) during falling tide and lagoonward (positive) during rising tide. The current on top of the 

reef crest behaved similarly but exhibited a much 

greater variation than it did in the microgrooves. 

Both microgroove and reef-top currents clearly 

display the tidal signature. When averaged over 

a 25-hour period (2 semidiurnal or 1 diurnal 

cycle), however, the microgrooves showed a net 

oceanward flow, whereas the water on top of the 

crest is directed toward the lagoon (Figure 43). 

Dye experiments during the falling tide indicated a jet-like flow from the microgrooves into 

the forereef region. Once outside the reef, some of 

this water, is transported southward with the 

longshore current. Most of the dye seemed to 

return via wave-pumping over the reef crest both 

up- and down-stream of the microgroove exits, 

thus setting up a series of circulation cells along 

the outer reef edge. The currents are much more 

vigorous at microgrooves C, D, and E compared 

with B and A. Instantaneous and time-averaged 

values were greater here because in this region 

the incident waves have been subjected to a 

minimum of refraction. 

The features described thus far reflect trade 

wind conditions estimated to occur 70 percent of 

the time (Riitzler and Ferraris, herein). During 

minor storm events and other times of high wave 

activity, however, both reef crest and microgroove 

net currents flow toward the lagoon and do not 

display any tidal influence. At these times, the 

wave set-up is probably sufficiently greater to 

dominate any tidal influence by exceeding any 

differences in tidal hydraulic head. Water then 

leaves the shallow Carrie Bow lagoon both at the 

north and south of the island. 

Although Hernandez-Avila et al. (1977) found 

that during storms, coral rubble was transported 

from the deep fore reef to form coral boulder 

ramparts along the south coast of Grand Cayman, 

elsewhere in the Caribbean sand-sized material 

appears to be slowly transported away from the 

reef crest. These observations are not contradictory. In view of the measured ocean-directed net 

flow at Carrie Bow Cay, it can be assumed that 

the microgrooves act as a passway for sediment 

from the lagoon and reef crest during typical 

conditions. When a storm strikes, however, the 

wave drift is directed up the fore reef and the 

forces are great enough to transport boulders 

toward and over the reef. The rubble rampart 

that makes up the SE reef crest of Carrie Bow 

Cay (Riitzler and Macintyre, herein:9) undoubtedly owes its existence to wave transport during 

heavy storms. 

Literature Cited 

Hernandez-Avila, M. L., H. H. Roberts, and L. J. Rouse 

1977. Hurricane-generated Waves and Coastal Boulder 

Rampart Formation. In D. L. Taylor, editor, Proceedings, Third International Coral Reef Symposium, 2: 

71-78. Miami, Florida: Rosenstiel School Of Marine and Atmospheric Science. 

Munk, W. H., and M. C. Sargent 

1948. Adjustment of Bikini Atoll to Ocean Waves. Transactions, American Geophysical Union, 29(6):855-860. 

Roberts, H. H., S. P. Murray, and J. N. Suhayda 

1975. Physical Processes in a Fringing Reef System. 

Journal of Marine Research, 33(2):233-260. 

Shinn, E. 

1963. Spur and Groove Formation on the Florida Reef 

Tract. Journal of Sedimentary Petrology, 33(2):291-

303. 

Tait, R.J. 

1972. Wave Set-up on Coral Reefs. Journal of Geophysical 

Research, 77(12):2207-2211. 
Geology and Sediment Accumulation Rates 

at Carrie Bow Cay, Belize 

Eugene A. Shinn,J. Harold Hudson, Robert B. Halley, 

Barbara Lidz, Daniel M. Robbin, and Ian G Macintyre 

ABSTRACT 

A 24 km long transect of cores consisting of 

four rotary drill cores and seven vibrocores was 

drilled in a line extending from the seaward side 

of the reef crest at Carrie Bow Cay to the mainland at a point between Sittee Point and the town 

of Stann Creek. Two of the four rotary cores were 

drilled seaward of the reef crest, one through a 

spur to a depth of 7.6 m and the other into the 

adjacent groove to a depth of 18.3 m. The two 

cores show no evidence that the spur and groove 

system was erosional in origin; rather, they demonstrate that it was constructional. Submarine 

cementation, chiefly in the form of fine-grained 

high-magnesium calcite, was found mainly in 

cemented internal sediment in both cores drilled 

seaward of the reef crest. 

Two other rotary cores were drilled landward 

of the reef crest. One was drilled on the reef flat 

to a depth of 8.8 m, and the other was drilled on 

the southwest tip of Carrie Bow Cay to a depth 

of 17.7 m. Both cores encountered essentially 

uncemented carbonate reef sands with some coral 

rubble. Of the four rotary cores, only the Carrie 

Bow Cay core encountered Pleistocene bedrock. 

Radiocarbon dating of a large head of Siderastrea 

siderea, growing on bedrock from the Carrie Bow 

Cay core at a depth of 15.04 m below sea level, 

gave an age of 6960±110 years. The leached 

calcitic coralline bedrock, at a depth of 15.7 m 

below sea level in the Carrie Bow Cay core, 

contained root marks, and iron staining indicative 

of subaerial exposure. 

Eugene A. Shinn, J. Harold Hudson, Robert B. Halley, Barbara 

Lidz, Daniel M. Robbin, United States Geological Survey, Fisher 

Island Station, Miami Beach, Fla. 33139. Ian G. Macintyre, 

Division of Sedimentology, Department of Paleobiology, Smithsonian 

Institution, Washington, D.C. 20560. 

Five of the seven vibrocores, with sediment 

recoveries ranging from 1.6 to 5 m, contained 

peat. Generally, the peat was located near the 

bottom of the cores, and in the four most landward cores the peat overlay clay, silt, quartz sand, 

and in some a few quartzite pebbles. The peats 

are interpreted to record flooding of the coastal 

plain during the last transgression. Radiocarbon 

dates of peats overlying terrigenous sediments 

range from 6804±150 to 8808±600 years. 

Introduction 

This paper reports the results of rotary rock 

coring on and around Carrie Bow Cay and describes a transect of seven sediment vibrocores 

drilled in a line from Carrie Bow Cay to the 

mainland, 24 km to the west. 

Carrie Bow Cay, literally within a stone's throw 

of the reef at 16°48'10"N, 88°04'45"W, is situated 

between two tidal passes that cut through an 

otherwise continuous reef flat (Figure 2). Tidal 

passes through the Belize barrier reef are rare, 

but wherever they occur, they are adjacent to 

islands situated near the edge of the reef tract. 

Carrie Bow, like other islands on the shelf south 

of the Belize River, is a Holocene sedimentary 

accumulation. Although there are extensive exposures of Pleistocene limestones at the northern 

limits of the Belizean barrier reef complex, notably Ambergris Cay, nowhere within Belizean waters south of the Belize River do Pleistocene or 

older limestones extend to or above sea level to 

form an island, as is common in other coral reef 

areas of the Caribbean. Sediments, vegetation, 

63 
64 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

88"10' 

16 50 

Tobacco Cay 

Tobacco 

Reef 

CBf 

* V Twin Cays 

/ \l3 7 $ 

\f * • NJJB2 ^South Water Cay 

/jfcoV o 0 *>^CB1 ) 

'^'). o ° y ^^")0Came Bow Cay 

Columbus 

Reef 

-i i i i I 

5 km 

-16*50' 

88*10' 

FIGURE 44.—Map of Carrie Bow Cay study area, Belize, showing transect of sediment 

vibrocores (CB1-CB7). 

FIGURE 45.—Map of immediate Carrie Bow Cay area showing rotary core locations (RC1-RC4) and the easternmost 

sediment vibrocore (CB1). 

A 

CB1 

RC4^ 

JVansect 

« / ) 

Carrie Bow C ay 

RC3 ^ 

^ RC 1 

% _ RC2 

^ Reef Crest 

and effects of storms on Belizean islands, including Carrie Bow Cay, have been described by 

Stoddart (1962, 1963). 

Vibrocore locations are shown in Figures 44 

and 46 and rotary drill core locations are shown 

in Figures 45 and 46. Core RC2, immediately 

adjacent to RC1, was taken on a spur (Figure 

47). This core was compared with core RC1, 

taken in a groove, to determine whether the spur 

and groove system was formed by erosion, as 
NUMBER 12 65 

proposed by Cloud (1959) for Saipan reefs or by 

construction, as proposed for Jamaica reefs by 

Goreau (1959) and determined for a Florida reef 

by Shinn (1963). 

This work, conducted on and around Carrie 

Bow Cay, is part of a larger United States Geological Survey Belize project, the objectives of 

which are to determine (1) porosity distribution 

as controlled by submarine cementation, (2) factors that control reef distribution, (3) nature and 

origin of the numerous patch reefs lagoonward of 

the barrier reef, and (4) rate of sediment accumulation during the past 10,000 years. Only that 

work conducted on and around Carrie Bow Cay 

is reported here. 

ACKNOWLEDGMENTS.—We gratefully acknowledge the gracious cooperation of the Belize government, in particular the fisheries department. 

This work would not have been possible without 

the aid of R. Gaensslen, owner and captain of the 

M /V Sea Angel. We also acknowledge the invaluable aid of P. Shea, who acted as diver and 

technician. The adapted jackhammer used in 

sediment vibrocoring was designed and perfected 

by D. Lanesky of the Comparative Sedimentology Laboratory, University of Miami, Fisher Island Station, Miami Beach, Florida. The use of 

brand names in this report is for descriptive purposes only and does not constitute endorsement 

by the United States Geological Survey. 

RC4 RC3 

FIGURE 46.—Cross sections of study area: a, entire area shown in Figure 44, with vibrocore (CB) 

and rotary core (RC) locations; b, detailed cross section of Carrie Bow Cay and associated reef 

area comparable to Figure 45. Note distribution of corals seaward of reef crest, which is 

composed of coral rubble encrusted with Millepora sp. The inferred transition zone from in situ 

coral accumulation to sand and rubble is indicated by the zig-zag line. Staghorn corals shown 

near cores RC3 and RC4 are inferred. Staghorn corals in RC 1 and RC2 were recovered because 

of cementation. RCl and RC2 are at the same distance along the E-W section, RC2 being 

immediately to the N of RCl. (For explanation of symbols, see Figure 47.) 
66 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

5 -

10-

15-

2 0 -

25 J 

R C 2 

7175 ±100 

^c S ^ ^ - Coralline algae 

& Agaricia spp. 

>< Acropora cervicornis 

(2& Montastrea annularis 

^ Porites astreoides 

^ Diploria 

10m 

FIGURE 47.—N-S cross section between cores RCl (drilled in a groove) and RC2 (drilled on 

adjacent spur) showing distribution of major coral components. Radiocarbon dates indicate 

that the section is less than 7175 years old and that growth rates are rapid. See Table 6 for 

accumulation rates. Also note apparent date reversal in corals between 14 and 17 m. Both ages 

are within the margin of error for the method; the samples should therefore be considered 

approximately.the same age. These two dates indicate an extremely rapid rate of accumulation 

for that interval. 

Methods 

ROTARY DRILL CORING.—Core drilling into the 

reef (Figure 48a) was accomplished with a version 

of the underwater hydraulic rotary drill described 

by Macintyre (1975). Modifications include (1) a 

portable, lightweight power source (manufactured by Custom Hydraulics of Miami, Fla.) 

measuring 1 m in length, 60 cm in width, 80 cm 

in height, and weighing only 125 kg, which is 

easily operable from a small skiff; (2) a lightweight plastic hydraulic hose, Vi inch (12.7 mm) 

in diameter; and (3) a lightweight, collapsible 

aluminum drilling tripod (Figure 48a). Standard 

length (5 ft, 1.5 m) N drilling rods and both X 

and BX diamond-tipped core barrels were used. 

Although the spur and groove zone was drilled 

to a depth of 18 m (RCl), only nine meters of the 

reef flat could be penetrated because of collapsing 

sand. Drilling the sands of Carrie Bow Cay required drilling mud, but commercial drilling mud 

FIGURE 48.—Underwater coring equipment employed: a, 

diver-operated hydraulic drilling rig at location of RCl; b, 

hydraulic jackhammer in operation; c, equipment used for 

vibrocoring (A = hydraulic jackhammer; B = adapter that 

mates jackhammer to core tube; C = plastic cap for capping 

core before extraction; D = section of 3-inch (76 mm) 

diameter aluminum core tubing, with vents for letting out 

water as core is taken; E = aluminum clamps with handles 

used for extracting core from sediment). 

68 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

was not available; a mixture of clay and silt dug 

from an area near Stann Creek on the mainland 

and diluted with sea water sufficed to accomplish 

the drilling. Five garbage cans containing sediment were brought to the island and dumped 

into a pit lined with plastic. The diluted mixture, 

necessary to prevent collapse of the hole, was used 

to drill the site near the south tip of Carrie Bow 

Cay to a depth of 16.2 m, where Pleistocene 

limestone appeared. Total depth including Pleistocene bedrock was 17.7 m. 

VIBROCORING.—Sediment vibrocoring (Figure 

48b) was accomplished by adapting a 36 kg hydraulic jackhammer (manufactured by Fairmont 

Hydraulics of Fairmont, Minn.), similar to the 

familiar air-powered jackhammers used to break 

up concrete, so that it could use 6 and 9 m lengths 

of standard 3 in (7.6 cm) aluminum irrigation 

tubing. The advantages of this hydraulic equipment were reduced corrosion and the ability to 

utilize the same hydraulic power source as that 

used for rotary drilling. The jackhammer was 

joined to the thin-walled tubing by an adapter 

that was constructed by press fitting an internally 

fitted steel sleeve to a standard, hardened steel 

chisel made especially for the hammer. The sleeve 

is a friction-fitting device that slips approximately 

10 cm into the tubing. Further insertion is prevented by a collar (Figure 48c). 

Water is able to escape from the core tubing as 

it penetrates sediment by means of relief ports 

added near the upper end of the tubing (Figure 

48c). Slits were cut into the tube with a hacksaw 

and one side of each cut hammered inward, so 

that the resulting ports resemble gill slits. Initially, ports were made with an electric drill, but 

the saw and hammer method was found to be 

more efficient. 

Coring was accomplished by attaching either 

the 6 or 9 m lengths of tube to the jackhammer 

and floating the entire device with an inverted 

air-filled garbage can. When the core tube was 

vertical over the selected site, one diver released 

air from the garbage can while another diver 

guided the unit to the bottom. When the tube 

contacted bottom, the diver depressed the trigger 

and the machine-gun action of the hammer drove 

the core tube into the sediment (Figure 48b). 

Generally, 6 to 9 m of penetration was achieved 

in less than 30 seconds with an average of 80 

percent recovery. In very clear water the tube was 

first driven approximately

 xh m vertically into the 

bottom with a 1 kg sledge hammer. When the 

tube was correctly oriented and free standing, the 

jackhammer was floated into position and attached to the adapter, and coring began. Because 

the tube itself does not rotate during coring, 

orientation of the core can be determined by 

putting a scratch on one side of the soft aluminum 

tube before extracting the core from the bottom. 

Cores were removed from the bottom with the 

aid of adjustable clamps with handles that were 

attached to the tube at the sediment/water interface. Excess tubing was sawed off underwater 

by hacksaw. An externally fitting plastic cap 

(Caplug EC-48, manufactured by Caplugs Division of Protective Closures Company, Inc., Buffalo, N.Y.) was affixed to the protruding tube 

end, thereby creating a vacuum and preventing 

loss of core material from the buried end during 

extraction. The cores were extracted and brought 

to the surface by attaching the garbage can and 

filling it with air from the SCUBA regulator. 

Although core' catchers were not used, sediment 

was seldom lost during extraction. 

All vibrocores were dried and impregnated 

with plastic according to the methods of Ginsburg 

et al. (1966). Peats were removed prior to impregnation. Carbon-14 dating was carried out at the 

University of Miami's Radiocarbon Dating Laboratory. 

Sediment types are described according to the 

classification of Dunham (1962). In keeping with 

his scheme, loose sediment is described as if it 

were the rock it will eventually become. Thus, 

"packstone" is used to describe sand-size carbonate grains whose interstices are filled with lime 

mud; "grainstone" is used if mud is absent. In 

addition, mud-supported sediments are termed 

"mudstones" if they contain less than 10 percent 

sand-size grains and "wackestones" if they have 

more than 10 percent. 
NUMBER 12 69 

Results 

OUTER REEF.—Of the four cores drilled in the 

Carrie Bow Cay area, only RCl and RC2 contained in situ corals. Core RCl from a groove in 

7 m of water was by far the most difficult to drill 

(see Figures 45, 46, 47). The core was drilled to 

18.3 m before caving and other difficulties prevented further penetration. Pleistocene bedrock 

was not encountered. Table 5 shows that core 

recovery was about 36 percent. Core RC2, approximately 10 m to the north on the adjacent 

spur, was drilled to a depth of 7.6 m, the first 3.7 

m being in the spur. Because the spur is composed 

primarily of the lettuce coral, Agaricia sp., and 

little cementation or infill of internal sediment is 

present, there was practically no core recovery in 

this interval. Recovery of material began when 

the drill reached the level of the adjacent groove. 

Because the material from both groove and spur 

base was identical, we concluded that the spur is 

of constructional origin. We recognize, however, 

the possibility of a deep-seated erosional spur and 

groove in the Pleistocene strata below 18 m which 

might serve as a template for later growth. Nonetheless, the visible portion of the spurs in this area 

is clearly constructional, and the interval 3.7 to 

7.6 m (below the spurs) is probably also constructional. 

The most noticeable feature in core RCl and 

the lower 3.7 to 7.6 m of RC2 is the occurrence of 

both cemented and uncemented internal sediment. Fine-grained internal sediment was generally light gray in color and contained sedimentary 

laminations, commonly inclined. The cement is 

Mg-calcite, similar to that reported elsewhere 

(Ginsburg et al., 1967; Macintyre et al., 1968; 

James et al., 1976; Land and Goreau, 1970; Macintyre, 1977; Shinn, 1969, 1971). In partially 

filled small cavities (less than 1 cm across), internal sediment is commonly geopetal, that is, the 

upper surface of the fill is horizontal. In larger 

voids, however, such fillings are generally inclined 

(Figure 49). Very little uncemented sediment was 

recovered from large voids, but its presence was 

indicated by bursts of muddy drilling water asE A/**''1 

*• a 

FIGURE 49.—Two pieces of core from RCl: a, section from 

10.75 m showing cemented internal sediment at steep angle 

(arrow); note unusual network of voids in cemented internal 

sediment; cause of the void network is not known, although 

it does not appear to be caused by leaching; b, sample from 

7.4 m; gray internal sediment resting on Porites coral; note 

steeply inclined laminations within cemented internal sediment. Cement is Mg-calcite; origin of voids is not known, 

but they are thought to be organic borings. 

sociated with sudden drops of the drill bit and by 

small amounts of sediment recovered in the cores. 

Fibrous aragonite, probably the most diagnostic form of submarine cement, was present only 

in small cavities, usually within shell and coral 

chambers, and was visible only in thin sections. 

This observation is identical to that reported by 

Macintyre (1977) but is in strong contrast to the 

fibrous aragonite crystals 1 to 2 cm long described 

by Ginsburg and James (1976) and James and 

Ginsburg (1978) from the deep fore reef at Belize 

below 100 m. 

REEF FLAT AND ISLAND.—Core RC3 was drilled 

on the reef flat approximately 15 m behind the 

reef crest in water about 25 cm deep. The seabottom at the RC3 site consisted of a hard pavement 
70 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

composed of cemented sediment sparsely populated by small head corals. Drilling showed the 

pavement to be only 1 cm thick and underlain 

by coral rubble imbedded in coarse carbonate 

reef sand. From approximately 2 m to total depth 

(8.8 m), only carbonate sand with sparsely scattered coral fragments was encountered. Except 

for the thin cemented pavement exposed at the 

surface, no evidence of submarine cementation 

was found, nor did any corals appear to be in 

growth position. 

Core RC4 was drilled on the southwest tip of 

Carrie Bow Cay (Figures 45). Drilling revealed a 

section consisting mainly of coarse-grained carbonate sand. At a depth of 15.5 m, a single large 

head of Siderastrea siderea appeared in growth position on Pleistocene bedrock. Bedrock was 

reached at 16.2 m and cored to 17.7 m. It is 

coralline limestone that has been leached and 

contains brown caliche staining. The coral overlying the bedrock was dated by carbon-14 technique and is discussed below. There was no evidence of cementation of the Holocene sands beneath Carrie Bow Cay. 

TABLE 5.—Locations, water depths, core recovery, and general comments on rotary drill cores 

and sediment vibrocores taken in the Carrie Bow Cay area, Belize 

Core 

ROTARY DRILL CORES 

R Cl 

RC2 

RC3 

RC4 

VIBROCORES 

CB1 

CB2 

CB3 

CB4 

CB5 

CB6 

CB7 

Lat, Long. 

16°48'13"N, 

88°04'41"W 

16°48'13"N, 

88°04'41"W 

16°48'13"N, 

88°04'43"W 

16°48'10"N, 

88°04'45"W 

16°48'10"N, 

88°05'10"W 

16°47'50"N, 

88°06'30"W 

16°49'30"N! 

88°07'45"W 

16°50'20"N, 

88°08'40"W 

16°53'25"N, 

88°14'00"W 

16°54'15"N, 

88°15'50"W 

16°54'15"N, 

88°15'88"W 

Location 

Description 

About 60 m seaward 

of reef crest 

10 m N of core RCl 

On reef flat 15 m W 

of reef crest 

SW tip of Carrie Bow 

Cay 

About 150 m W of 

Carrie Bow Cay 

S of Twin Cays 

See Figures 44 and 45 

Just W of dropoff into 

Victoria Channel 

See Figures 44 and 45 

~2 km from mainland 

~300 m from shore 

Water 

depth 

(m) 

6.7 

3.7 

0.25 

+0.5 

2.5 

7.3 

11.5 

20.2 

12.2 

6.7 

4.3 

Core hole 

depth 

(m) 

18.3 

7.6 

8.8 

17.7 

4.6 

3.5 

5.4 

2.0 

3.4 

3.9 

3.1 

Recovery 

(m) 

6.55 

1.8 

=0.5 

=0.03 

3.7 

2.8 

5.0 

1.6 

2.7 

3.7 

2.4 

Comments 

Drilled in groove 

Drilled on spur; recovery only below base 

of spur 

Poor recovery due to 

uncemented sand 

No core recovery above 

15m; good recovery 

from Pleistocene 

16.2-17.7 m 

Carbonate sand bottom with scattered 

Thalassia species 

Peat extends from 

about 2.3 m to bottom of core recovery 

at 2.8 m 

Contains peat from 4.5 

m to bottom of core 

recovery at 5.0 m 

Sticky red clay prevented further core 

penetration 

Peat in bottom 50 cm 

of core; water visibility about 25 cm 

Peat in upper section 

of core; see Table 6 

Bottom 3/5 of core is 

peat 
NUMBER 12 71 

r% 

JA* 

10cm 

FIGURE 50.—Selection of plastic-impregnated sediment cores from transect shown in Figure 44: 

a, core CB1, interval 40 to 70 cm; note burrowing, grass roots, and coral fragment at upper 

right; smaller white fragments are Halimeda; b, same interval from core CB2; note large burrow; 

c, core CB2 from interval 2.2 to 2.5 m showing transition from lime sediment to peat; note coral 

fragments at top, Oculina sp., and large unidentified oysters and other mollusks at transition; d, 

core CB4 showing transition from carbonate grainstone to orange terrigenous clay, contact is at 

base of large gastropod, cracks in lower part of core are due to dessication in clay, which would 

not impregnate because of its fine-grain size; e, top of core CB6 (interval 0 to 30 cm); upper 

portion consists of coarse, poorly sorted quartz sandstone merging with peat near bottom; large 

white fragments are quartz pebbles. 

VIBROCORE TRANSECT.—Location of all seven 

vibrocores is shown in Figures 44 and 46, and 

brief comments on depth, location, and penetration appear in Table 5. Selected photographs of 

these plastic-impregnated cores are shown in Figure 50. 

Core CBl consists entirely of burrowed, reefderived grainstone but lime silt content increases 

slightly near the base, locally creating a packstone 

facies associated with burrows. Despite a few 

sticks oi Acropora cervicornis, the principal sand-size 

constituents are Halimeda plates and soritid foraminifera. Homotrema sp., a foraminifer generally 

abundant in reef-derived sands, is rare. 

Core CB2 is predominantly a packstone mottled by abundant burrows. Mollusks are the most 

obvious large fossils and soritid foraminifera are 

abundant, approximately as abundant as Halimeda plates. At 2 m, there is a transitional change 

to lime silt, grading into a mixture of peat and 
72 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

lime silt. Large oyster shells are mixed into the 

peat down to 2.7 m. Broken oyster shells extend 

deeper. Coral fragments of Oculina sp. occur between 2.1 and 2.2 m, just above the peat. 

Core CB3 consists of burrowed packstone and 

silty lime mudstone with pelecypods to 3.7 m. 

Sediment becomes darker at 3.7 m owing to 

abundance of peat flecks. Pelecypods occur between 4 and 5 m. Sediment is lime silt between 4 

and 4.5 m with an admixture of clay that gives it 

a gray-brown color. Peat begins at 4.5 m and 

extends to the bottom of the core at 5 m. Gastropods, such as Astrea sp. and various cerithids, 

occur just at the transition from lime sediment to 

peat. 

Core CB4 was taken in 20 m of water immediately landward of a continuous shallow rock 

ridge (see Figure 46). The upper 1 m of the core 

consists of burrowed mixed grainstone and packstone. The grains are Halimeda, mollusk fragments, and foraminifera. Soritid foraminifera are 

rare, and obvious darkened grains make up 1 to 

3 percent of the section. At 1.24 m the lime 

sediment changes into brownish-orange clay containing red-brown concretions. The unexpected 

occurrence of such coarse-grained sediment at 

this depth is probably explained by the nearby 

shallow ridge. Sediment is probably periodically 

swept landward off this ridge. 

Core CB5 consists of brown mud and wackestone with terrigenous clay containing scattered 

mica flakes. The predominant fauna is small, 

thin-shelled pelecypods. At 2.3 m there are large 

oyster shells, and at 2.5 m the sediment grades to 

a darker color clay and organic-rich mudstone. 

Peat was collected from 1.78 and 2.67 m for 

dating. 

Core CB6 consists of 30 cm of very poorly 

sorted quartz sandstone with quartzite pebbles 

up to 1 cm in diameter. The only carbonate is 

pelecypod fragments. Peat begins at 30 cm and 

extends to 1.3 m. The remainder of the core down 

to 3.65 m consists of massive, dirty sandstone with 

root marks. Between 3.65 m and the bottom of 

the core at 3.7 m, the sediment is gray clay with 

bright-red clay mottles. 

Core CB7 consists of almost 1 m of burrowed, 

poorly sorted quartz sandstone with pelecypod 

fragments, scattered Halimeda plates, and few gastropods. Sediment contains peat fragments and 

abundant quartzite pebbles at 75 cm. The section 

becomes peat at 0.92 m, and peat extends to the 

bottom of the core at 2.4 m. The peat contains 

thin stringers of quartzite pebbles. 

Peat was exposed on the bottom in 1 m of 

water landward of site CB7, and a peat-forming 

swamp is present at sea level all along the shore, 

just landward of a thin quartz-sand beach ridge. 

The change from coarse-grained carbonates to 

terrigenous elastics and peats in a shoreward 

direction follows the basic pattern described by 

Purdy et al. (1975). 

Although red mangroves are abundant on the 

seaward edge of the swamp, numerous hardwoods, grasses, and small palms indicate the peat 

is not entirely of mangrove origin. Nevertheless, 

the peat is accumulating at sea level and can be 

visually correlated with the submarine pet just 

offshore and is thought to be continuous with the 

peat in cores CB6 and CB7. 

RADIOCARBON AGE DATING—Corals from various depths within the rotary cores RC 1 and RC4 

were sampled and dated by the radiocarbon 

method, as were peats from sediment vibrocores 

CB5, CB6, and CB7. Dates, calculated accumulation rates, and sediment and water depths are 

provided in Table 6. In this table there are two 

columns of figures under the heading "Accumulation rates (m/1000 y)". The first column (core 

top to sample) lists the accumulation rates in 

meters/1000 years, assuming the top of the reef is 

growing and has a 14C age of approximately zero. 

The second column (intervals between samples) 

lists the calculated accumulation rates between 

dated samples. This figure is more meaningful for 

obvious reasons. All peat dates were obtained by 

D. S. Introne in 1978 at the University of Miami's 

Radiocarbon Dating Laboratory. Coral material 

was dated at the same laboratory by J. J. Stipp. 

It can be seen from Table 6 that the reef off 

Carrie Bow Cay has been growing upward at a 

rapid rate (ranging from approximately 1 to 6 

m/1000 y), considerably faster than accumulation of the lagoonal sediments overlying the peats 
NUMBER 12 73 

TABLE 6.—Radiocarbon dates and accumulation rates (m/1000 y) of selected material in 

rotary drill cores and sediment vibrocores (further explanation in text; location coordinates 

listed after each core number; note unexplained age reversal in two samples from RCl) 

RCl 

RC4 

CB5 

CB6 

CB7 

Core 

16°48'13"N, 

88°04'41"W 

16°48'10"N, 

88°04'45"W 

16°53'25"N, 

88°14'00"W 

16°54'15"N, 

88°15'50"W 

16°54'15"N, 

88°15'88"W 

Lab. no. 

UM-1009 

UM-1010 

UM-1011 

UM-1012 

UM-1013 

UM-1249 

UM-1250 

UM-1252 

UM-1251 

UM-1248 

Material 

dated 

Montastrea 

annularis 

Porites sp. 

M. annularis 

Porites sp. 

Siderastrea 

siderea 

Peat 

Peat 

Peat 

Peat 

Peat 

Depth 

in 

core 

hole 

(m) 

5.79 

8.23 

10.97 

17.67 

15.54 

1.78 

2.67 

0.30 

0.62 

0.92 

Interval between 

samples 

(m) 

2.44 

2.74 

6.70 

0.89 

0.32 

Water 

depth 

(m) 

6.7 

6.7 

6.7 

6.7 

+0.5 

12.2 

12.2 

6.7 

6.7 

4.3 

Depth 

from 

water 

surface 

to 

sample 

(m) 

12.49 

14.93 

17.67 

24.37 

15.04 

13.98 

14.87 

7.00 

7.32 

5.22 

Age 

difference 

between 

samples 

(y) 

540 

0 

1035 

618 

2004 

UC age 

(yBP) 

5625±85 

6165±90 

6140±90 

7175+100 

6960+110 

7619-320 

8237+270 

6804+150 

8808+600 

2861 + 190 

Accumulation 

(m/WOOy) 

Core 

top Interto 

sample 

1.03 

1.33 

1.79 

2.46 

2.23 

0.23 

0.32 

0.04 

0.07 

0.32 

vats between 

samples 

4.52 

0 

6.47 

1.44 

0.16 

(<1 to = 1.4 m/1000 y). This agrees with Purdy's 

(1974a) observation of an approximately 10 to 1 

difference in accumulation rate between the barrier platform and the shelf lagoon. Our reef accumulation rates exceed rates determined for the 

Florida reef tract. Shinn et al. (1977) determined 

accumulation rates in Florida to range from 0.38 

m/1000 y to 4.85 m/1000 y. More rapid accumulation rates have been reported at Alacran 

Reef (north of the Yucatan Peninsula in the Gulf 

of Mexico) by Macintyre et al. (1977), where the 

average of four dated intervals is 5.6 m/1000 y 

and one interval indicates a rate of 12 m/1000 y. 

Further discussion of reef accumulation rates, 

both in the Atlantic and Pacific, has been provided by Adey (1978). 

Discussion 

Figure 46 is an interpretation of facies changes 

across the Carrie Bow reef. Core RC3 on the reef 

flat just behind the reef crest clearly demonstrates 

that the reef flat is not composed of corals that 

have grown up to sea level, but instead is composed of reef sands and some rubble, which have 

probably been thrown up and over the reef crest 

during storms. In many areas along the lagoonward edge of the reef-flat sands, patch reefs are 

in the process of being buried as these sands 

accrete landward to extend the reef flat. Reefs off 

the Florida Keys are also growing landward 

(Shinn et al., 1977). From his work on Alacran 

Reef, Logan (1969:189) termed such accumulations off-reef clastic drape reefs (model 3). 

If underlying topography accounts for the location of Carrie Bow Cay, the proof cannot be 

found in our drilling. Previous work (Halley et 

al., 1977) proved conclusively, however, that Boo 

Bee Patch Reef, near Wee Wee Key about 8 km 

to the southwest, was initiated over a 6 m Pleistocene topographic high. Drilling on two mangrove islands situated on patch reefs also con-
74 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

firmed Pleistocene bedrock topography to be the 

controlling factor. In both cases Pleistocene coralline limestone reached to within 12 m of sea 

level under the island. Surrounding the island, 

the rock was more than 21 m below sea level. On 

the basis of his 1960 studies, Purdy (1974a,b) was 

the first to suggest that rock floor topography is 

the factor controlling island distribution on the 

Belize shelf. We suspect, therefore, that the reef 

off Carrie Bow Cay was initiated on a pre-existing 

rock high and that initial flooding, determined 

by

 14C age of coral growing directly on bedrock 

(see Table 6), occurred before approximately 

7000 y BP. Whether there has always been an 

island since early Holocene flooding or whether 

the island (actually little more than a sand spit) 

sprang into existence during the past few thousand years has not been determined. 

On the basis of our observations and drilling, 

the spurs and grooves appear to be relatively 

recent in age and clearly are not of erosional 

origin. Because the Holocene reef is greater than 

18 m thick, however, it seems unlikely that the 

present day spur and groove systems have existed 

in the same place throughout the reefs history. 

This conclusion cannot be borne out by only two 

core holes. We suspect that such a system has 

existed in the past but that the position of the 

spurs and grooves has shifted laterally as the reef 

built up. Furthermore, it seems unlikely to us that 

the present system is simply patterned over a 

Pleistocene spur and groove system more than 18 

m below the present reef surface. 

Determining the origin of the spur and groove 

system at Carrie Bow Cay reef was beyond the 

purpose of the expedition; however, such a determination should be made. We encourage coral 

reef researchers to drill spur and groove systems 

at many sites in the Caribbean or Pacific to 

determine the significant factors that lead to spur 

and groove development. 

Conclusions 

The Holocene coral reef seaward of the reef 

crest is in excess of 18 m thick. The present spur 

and groove system is constructional. The reef flat 

is composed primarily of reef-derived carbonate 

sand with scattered coral rubble. Carrie Bow Cay 

is Holocene and composed primarily of carbonate 

sand over 15 m thick. Leached calcitic limestone, 

which was subaerially exposed during the last 

glacial period, underlies the island at a depth of 

16 m. Flooding of the bedrock occurred prior to 

7000 years ago. Submarine cementation is, for 

the most part, restricted to the Holocene section 

seaward of the reef crest. Except for a thin exposed pavement directly shoreward of the reef 

crest, all reef flat and lagoonal sediments over a 

distance of 24 km are uncemented. Peat, representing former sea level, occurs in sediment cores 

in the lagoon. Peat in cores near the mainland 

overlie terrigenous clastic sediments; thus, the 

peats probably record the initial flooding of the 

coastal plain during the most recent transgression. 

Literature Cited 

Adey, W. H. 

1978. Coral Reef Morphogenesis: A Multidimensional 

Model. Science, 202(4370):831-837. 

Cloud, P. E. 

1959. Geology of Saipan, Mariana Islands, Part 4: Submarine Topography and Shoal Water Ecology. 

United States Geological Survey Professional Papers, 

280-K:361-445. 

Dunham, R. J. 

1962. Classification of Carbonate Rocks according to 

Depositional Texture. In W. E. Ham, editor, Classification of Carbonate Rocks. The American Association of Petroleum Geologists, Memoir, 1:108-121. 

Tulsa, Oklahoma. 

Ginsburg, R. N., H. A. Bernard, R. A. Moody, and E. E. 

Daigle 

1966. The Shell Method of Impregnating Cores of Un-
NUMBER 12 75 

consolidated Sediments. Journal of Sedimentary Petrology, 46:1118-1125. 

Ginsburg, R. N., and N. P. James 

1976. Submarine Botryoidal Aragonite in Holocene 

Reef Limestones, Belize. Geology, 4:431-436. 

Ginsburg, R. N., E. A. Shinn, and J. Schroeder 

1967. Submarine Cementation and Internal Sedimentation within Bermuda Reefs [Abstract]. In Program, 1967 Annual Meetings, The Geological Society of 

America, New Orleans, Louisiana, pages 78-79. 

Goreau, T. F. 

1959. The Ecology of Jamaican Coral Reefs, I: Species 

Composition and Zonation. Ecology, 40:67-90. 

Halley, R. B., E. A. Shinn, J. H. Hudson, and B. Lidz 

1977. Recent and Relict Topography of Boo Bee Patch 

Reef, Belize. In D. L. Taylor, editor, Proceedings, 

Third International Coral Reef Symposium, 2:29-36. 

Miami, Florida: Rosenstiel School of Marine and 

Atmospheric Science. 

James, N. P., and R. N. Ginsburg 

1978. The Deep Seaward Margin of Belize Barrier and Atoll 

Reefs. 201 pages. Miami Beach, Florida: University 

of Miami. 

James, N. P., R. N. Ginsburg, D. S. Marszalek, and P. W. 

Choquette 

1976. Facies and Fabric Specificity of Early Subsea Cements in Shallow Belize (British Honduras) Reefs. 

Journal of Sedimentary Petrology, 46(3):523-544. 

Land, L. S., and T. F. Goreau 

1970. Submarine Lithification of Jamaican Reefs. Journal of Sedimentary Petrology, 40(l):457-462. 

Logan, B. W. 

1969. Coral Reefs and Banks, Yucatan Shelf, Mexico. In 

B. W. Logan, J. L. Hardin, W. H. Ahr, J. D. 

Williams, and R. G. Snead, editors, Carbonate 

Sediments and Reefs, Yucatan Shelf Mexico. The 

American Association of Petroleum Geologists, Memoir, 

11:129-196. 

Macintyre, I. G. 

1975. A Diver-operated Hydraulic Drill for Coring Submerged Substrates. Atoll Research Bulletin, 185:21-

26. 

1977. Distribution of Submarine Cements in a Modern 

Caribbean Fringing Reef, Galeta Point, Panama. 

Journal of Sedimentary Petrology, 47(2):503-516. 

Macintyre, I. G., E. W. Mountjoy, and B. F. D'Anglejan 

1968. An Occurrence of Submarine Cementation of Carbonate Sediments off the West Coast of Barbados, 

W. I. Journal of Sedimentary Petrology, 38(2) :660-

664. 

Purdy, E. G. 

1974a. Karst Determined Facies Patterns in British Honduras: Holocene Carbonate Sedimentation 

Model. The American Association of Petroleum Geologists Bulletin, 58 (5):825-855. 

1974b. Reef Configurations: Cause and Effect. In L. F. 

Laporte, editor, Reefs in Time and Space. Society 

of Economic Paleontologists and Mineralogists, Special 

Publication, 18:9-76. 

Purdy, E. G , W. C. Pusey, and K. F. Wantland 

1975. Continental Shelf Attributes. In K. F. Wantland 

and W. C. Pusey, editors, Belize Shelf—Carbonate 

Sediments, Clastic Sediments and Ecology. The 

American Association of Petroleum Geologists, Studies in 

Geology, 2:1-40. 

Shinn, E. A. 

1963. Spur and Groove Formation on the Florida Reef 

Tract. Journal of Sedimentary Petrology, 33(2):291-

303. 

1969. Submarine Lithification of Holocene Carbonate 

Sediments in the Persian Gulf. Sedimentology, 12: 

109-144. 

1971. Aspects of Diagenesis of Algal Cup Reefs in Bermuda. Transactions, Gulf Coast Association of Geological Societies, 21:387-394. 

Shinn, E. A., J. H. Hudson, R. B. Halley, and B. Lidz 

1977. Topographic Control and Accumulation Rate of 

Some Holocene Coral Reefs: South Florida and 

Dry Tortugas. In D. L. Taylor, editor, Proceedings, 

Third International Coral Reef Symposium, 2:1-7. 

Miami, Florida: Rosenstiel School of Marine and 

Atmospheric Science. 

Stoddart, D. R. 

1962. Castastrophic Storm Effects on British Honduras 

Reefs and Cays. Nature, 196(4854):512-515. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays: October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 
Terrestrial Environment and Climate. 

Carrie Bow Cay, Belize 

Klaus Riitzler 

and Joan D. Ferraris 

ABSTRACT 

Severe hurricane activity during the past 20 

years has reduced Carrie Bow Cay (16°48'N, 

88°05'W) to half its pre-1960 size of 0.8 ha. At 

present, Carrie Bow Cay (0.4 ha area) is one of 

the smallest inhabited sand cays on the barrier 

reef of Belize. The island measures 120 X 36 m, 

rises 40 cm above mean tide level, and supports 

three wooden cottages with freshwater tanks. The 

only permanent terrestrial plants are about 60 

coconut trees. Other vegetation appears periodically and spreads until it is destroyed by intermittent storm tides. Conspicuous animals include 

a few birds, a lizard, and some supratidal crustaceans. About one-third of the island's surface is 

intertidal and occupied mainly by algae, crustaceans, and mollusks that are adapted to this 

habitat. The climate is oceanic and is dominated 

largely by northeasterly trade winds. 

Introduction 

Previous terrestrial investigation of Carrie Bow 

Cay was based on brief topographic and floristic 

surveys (Stoddart, 1963; 1969; 1974) and a shortterm meteorological study (Kjerfve, 1978). Our 

own first topographic survey was prompted by 

the severe impact of hurricane Fifi, in 197, on the 

shape and size of the island. From then on we 

monitored morphological changes of the cay, conKlaus Riitzler, Department of Invertebrate Zoology, National Museum 

of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. Joan D. Ferraris, Mount Desert Island Biological 

Laboratory, Salsbury Cove, Maine 04672. 

dition of the remaining coconut tree population, 

and recovery of the vegetation that had been 

entirely eliminated by salt water flooding. Our 

other observations on the terrestrial and intertidal 

flora and fauna of the islet are of casual qualitative nature and restricted to large and conspicuous organisms. Meteorological records were taken 

regularly during the months of our field work, 

mainly in spring and early summer, but are sporadic during the remaining parts of the year. 

Carrie Bow Cay has been the base of the Smithsonian Institution coral reef study since the initiation of the program in 1972. The small island 

provided the necessary support in close proximity 

of reef and lagoon habitats without having noticeable terrestrial effects on these environments. 

ACKNOWLEDGMENTS.—We thank H. Pulpan 

and M. Carpenter for their help in surveying 

Carrie Bow Cay. The following colleagues provided or confirmed identification of organisms 

mentioned in this report: S. H. Brawley (bluegreen algae); J. N. Norris (algae, except bluegreens); M. E. Hale (lichens); M.-H. Sachet 

(higher plants); J. C. den Hartog (cerianthids); 

W. A. Newman (barnacles); B. Kensley (crustaceans, except barnacles); R. S. Houbrick (mollusks); D. L. Pawson (echinoids); R. I. Crombie, 

F. J. Irish (reptiles); and W. Trivelpiece (birds). 

C. H. Sprinkle helped with information on meteorological monitoring practices. The following 

persons assisted in maintaining and evaluating 

weather records: H.T.A. Bowman III, M. Carpenter, B. Kjerfve, R. J. Larson, K. Leslie, A. 

77 
78 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Rath, D. S. Robertson, R. Sims and B. Spracklin. 

I. Jewett drafted the graphs. Special thanks are 

due P. J. Herbert for information on hurricane 

events in Belize, particularly for case histories of 

Fifi and Greta, and D. R. Stoddart for use of his 

1972 map of Carrie Bow Cay. 

Methods 

GEODETIC SURVEYS.—Tape measure, sighting 

compass, and sighting along permanent or temporary markers were used for determining shape 

and size of the island and position of trees and 

artificial structures. The concrete boat dock on 

the lagoon side and the wooden main house in 

the center of the cay provided the principal points 

of reference. Vertical photographs from a helicopter, in March 1976, helped to improve the documentation of the island's physiography. 

PLANT ABUNDANCE.—The first quantitative 

survey of plants made in 1978 consisted of subjective visual estimates of five categories of relative abundance. More objective measurements of 

area of plant coverage were made in 1979 by 

placing a 50 X 50 cm (0.25 m2) frame subdivided 

into 10 X 10 cm (100 cm2) fields over all surfaces 

of the island showing vegetation. The number of 

frames occupied by a given species was recorded 

to the nearest half frame (50 cm2). 

METEOROLOGY.—Generally, meteorological 

conditions were recorded three times a day (0600-

0800 h; 1200-1400 h; 1800-2000 h) whenever the 

laboratory on Carrie Bow Cay was in operation, 

most commonly during the periods January-June 

and October/November, 1976-1978. Temperature was measured with ±0.5° C accuracy in 

shaded air, in sun-exposed sand (5 cm below 

surface), in water on the reef flat (0.2 m average 

bottom depth), and below low-tide level at the 

boat dock (lagoon, 0.8 m average bottom depth). 

Some continuous analog chart recordings of solar 

radiation were made by pyranograph (WeatherMeasure B211). Wind speed and direction were 

read from a cup anemometer with air foil vane 

(WeatherMeasure W121). Precipitation was measured with two rain gauges (10 cm diameter), one 

installed on the cay, and the other one on the 

mainland at Pelican Beach Motel, Dangriga. The 

rainy season (June-September) of 1979 was also 

monitored by an unattended tipping-bucket raingauge (WeatherMeasure P501-I) with solar-powered event recorder on Carrie Bow Cay. Humidity 

was calculated from psychrometer (Psychro-Dyne 

PP100) readings. 

Physiography 

LOCATION.—Carrie Bow Cay (16°48'N, 

88°05'W) is a small sand island located on top of 

the barrier reef that lines the outer shelf edge of 

Belize (formerly British Honduras), Central 

America (Plate 1: center right). Its former name, 

Ellen Cay, is still recorded on many nautical 

charts. The nearest significant settlement is Dangriga (Stann Creek), a town of 7000 inhabitants 

on the mainland, 24 km due 320° (NW). The cay 

belongs to H.T.A. Bowman of Dangriga and is 

used as a vacation place for his family. The 

nearest islands are South Water Cay, 1.5 km due 

0° (N), a sand cay populated by a few fishermen 

and occasional vacationers, and Twin Cays (also 

known as South Water Range) 4 km due 323° 

(NW), an uninhabited mangrove development. 

Carrie Bow Cay is protected from open ocean 

waves by a crescent-shaped reef crest to the east 

and a 100 m wide reef flat that extends from the 

crest to the island's seaward shore (Plate 5: top 

left). 

SHAPE AND SIZE.—With a surface area of less 

than 0.4 ha, Carrie Bow Cay belongs among the 

smallest inhabited cays on the Belizean shelf 

(Stoddart and Fosberg, herein: 527) (Figure 51; 

Plate 1: bottom left). The island formerly was 

double its present size (H.T.A. Bowman, pers. 

comm.) and bordered by mangrove, but clearing 

of these trees in 1944 led to progressive erosion by 

storm tides. Stranded beachrock as far as 30 m 

east and south of the present seaward shore documents both a shift in dimensions and slow migration leewards. At present, the cay has an eliptical shape with approximate north-south exten-
NUMBER 12 79 

FIGURE 51.—Carrie Bow Cay, aerial view from the south, May 1973; note exposed beach rock 

on the reef flat east of the island. 

sion. The longer axis is directed due 30° (NNE) 

and measures 120 m between mean tide level 

(MTL) points; the greatest dimension perpendicular to this axis lies along a line transecting the 

center of the isle and measures 36 m. Surface area 

calculated from planimetry is 0.36 ha to MTL, 

0.25 ha if only dry-land (supratidal) area is measured. Highest elevation, which is approximately 

40 cm above MTL, occurs at the central portion 

of the island. 

SUBSTRATES.—Reef-derived carbonate sand 

and rubble on a base of Pleistocene bedrock 

(Shinn et al., herein: 63) make up the entire 

natural substrate of the cay (Figure 52a). Accumulation of beach sand varies with the direction 

and force of wind and currents. Under the influence of the predominant northeasterly trade 

winds, sand is deposited at the north point and 

northwest beach and around the south tip; at 

times separate intertidal sand spits are formed to 

the north. Concrete block seawalls and rubble 

and coral rock landfills built up over many years 

to delay erosion dominate the northwest (Figure 

52b) and southeast shorelines, which also have a 

few small sandy beaches here and there. Conch 

shells abandoned by generations of local fishermen are accumulated along the southwest coast. 

STRUCTURES.—Other than seawalls, artificial 

structures on Carrie Bow Cay include two docks 

and three buildings with water vats (Figure 51; 

Plates 1: bottom right, 5: top left). The main or boat 

dock to the west (lagoon side) of the cay is 26 m 

long and built of concrete. A smaller wooden 

dock over the reef flat (SE) serves the two outhouses. The buildings are wooden and include 

the main house, 14 X 12.5 m, "Junior's House," 

13 X 3.5 m (now serving as our project's laboratory), and a small cabin, 5 X 5 m (Figure 60). 
80 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 52.—Intertidal substrates: a, sandand rubble (mainly conch shells) on western shore, 

looking north (coconut palms felled by hurricane Greta); b, northwest seawall, coated by algae 

Cladophoropsis and Oscillatoria, toppled by Greta. 

Flora and Fauna 

Carrie Bow Cay's small size, lack of a freshwater lens, and exposed location near the open 

ocean are responsible for the absence of a complex 

permanent terrestrial environment. We distinguish between an intertidal zone along the shore 

and a central island area above the high-tide 

beach undercut, which, in our experience, has 

been flooded by sea water only during hurricane 

tides (see below). 

SUPRATIDAL ORGANISMS.—The most conspicuous, and only permanent, plants, except perhaps 

for the lichens, are coconut palms {Cocos nucifera 

L.) most of which were planted during the past 

35 years in at least five recognizable N-S rows. In 
NUMBER 12 II 

May 1979 this population consisted of 58 healthy 

trees, of which 38 were mature and showed either 

nuts or flowers, 8 were immature (one or more 

years established), and 12 were freshly planted 

after hurricane Greta (less than one year established). The remaining vegetation observed in 

May 1978 and May 1979 (before and after hurricane Greta; Table 7) recolonized Carrie Bow 

Cay after salt water flooding associated with hurricane Fifi (September 1974) had washed away 

TABLE 7.—Systematic list and relative abundance of Carrie Bow Cay plants, excluding coconut 

and others artificially introduced; figures for 1978 show recolonization of the island after 

hurricane Fifi (September 1974), when all vegetation was destroyed, and are visual estimates; 

data for 1979 reflect minor changes and losses (indicated by dash) in plant cover caused by 

hurricane Greta (September 1978) and are compiled from quadrat counts (see "Methods"); 

likely methods of dispersal are indicated for each plant (B = bird, D = drift, W = wind); 

approximation of size and frequency of plants is given for 1978 

Species 

(Family) 

Paspalum distichum L. 

(Gramineae) 

Sesuvium portulacastrum (L.) L. 

(Aizoaceae) 

Philoxerus vermicularis (L.) Beauvois 

(Amaranthaceae) 

Suaeda linearis (Elliott) Moquin 

(Chenopodiaceae) 

Portulaca oleracea L. 

(Portulacaceae) 

Coccoloba uvifera L. 

(Polygonaceae) 

Cakile lanceolata (Willdenow) O. E. Schulz 

(Cruciferae) 

Rhizophora mangle L. 

(Rh izophoraceae) 

Euphorbia blodgettii Engelmann ex Hitchcock 

(Euphorbiaceae) 

Euphorbia mesembrianthemifolia Jacquin 

(Euphorbiaceae) 

Ipomoea pescaprae brasiliensis (L.) van Ooststroom 

(Convolvulaceae) 

Ipomoea stolonifera (Cyrillo) Gmelin 

(Convolvulaceae) 

Tournefortia gnaphalodes (L.) Kunth 

(Boraginaceae) 

Eclipta alba (L.) Hasskark 

(Compositae) 

Unidentified seedling 

1 

2 

Total plant cover 

1978 

m

2 

> 5 

0.02-0.50* 

0.02-0.50* 

<0.01 

1-5** 

<0.01 

0.5-1.0f 

-

1-5** 

1-5** 

>5 

<0.01 

<0.01 

<0.01 

<0.01 

<0.01 

not 

determined 

m

2 

0.010 

2.585 

-

0.015 

4.825 

-

3.275 

0.010 

0.185 

1.820 

0.190 

-

0.035 

-

0.015 

-

12.965 

1979 

% total 

0.08 

19.94 

-

0.12 

37.22 

-

25.26 

0.08 

1.43 

14.04 

1.47 

-

0.27 

-

0.12 

-

100.03 

rank 

9a 

3 

8a 

1 

-

2 

9b 

6 

4 

5 

-

7 

8b 

-

Method of 

dispersal 

D? 

D?, B? 

D, B? 

D, B? 

B? 

D, B? 

D 

D 

B? 

B? 

D 

? 

D 

W, B? 

D 

D 

* few medium-sized plants, each 0.02-0.30 m 

** numerous small plants, each 0.01-0.02 m2 

f few large plants, each >0.3 m 
82 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

or killed all plants except the majority of coconut 

trees (Figure 53). In addition to the species listed 

in Table 7, two were recently artificially introduced: Casuarina equisetifolia L. (Casuarinaceae) 

and Hymenocallis littoralis (Jacquin) Salisbury (Liliaceae, sensu lato). At least three species of lichens 

are common on the northeast surfaces of wind 

exposed palm trunks: Lecanora subfusca (L.) Acharius, Pyxine cocoes (Swartz) Nylander, and Chiodecton 

sp. 

Although we have noted a variety of insects 

and a few spiders on Carrie Bow Cay, we have 

not determined the species and do not know 

whether they are breeding resident populations. 

Some ants, cockroaches, and spiders are no doubt 

FIGURE 53.—Plant cover on north point, May 1978: Ipomoea 

pescaprae, Cakile lanceolata (foreground, with 0.25 m frame), 

and freshly planted Casuarina tree. 

introduced by supply boats carrying produce. 

Fleas and ticks have been left behind by dogs, the 

former at times plaguing sensitive investigators. 

Flying insects are commonly blown over from 

land or larger islands during westerly winds. Most 

of the island's invertebrate fauna, however, consists of three crustaceans: the hermit crab Coenobita 

clypeatus (Herbst), and the crabs Ocypode quadrata 

(Fabricius) and Gecarcinus lateralis (Freminville). 

Only the lizard Anolis sagrei Dumeril and Bibron, a species widespread in the West Indies and 

apparently expanding its range onto Caribbean 

Mexico and Middle America (R. Crombie, pers. 

comm.), occurs as resident population of vertebrates on Carrie Bow Cay. A sea turtle, Caretta 

caretta (L.), was last seen laying eggs on the island 

on 28 May 1972 (A. Antonius, pers. comm.). 

Birds that feed regularly around the cay are the 

Brown Pelican {Pelecanus occidentalis L.), Frigatebird {Fregata magnificens Mathews), and Osprey 

{Pandion haliaetus (L.)). Other birds commonly 

seen include the Boat-tailed Grackle {Cassidix 

mexicanus (Gmelin)), Common Tern {Sterna hirundo 

L.), Brown Booby {Sula leucogaster (Boddaert)), 

Snowy Egret {Leucophoyx thula (Molina)), and 

Barn Swallow (Hirundo rustica L.). An assortment 

of involuntary visitors from the mainland, such 

as warblers and flycatchers, arrive exhausted on 

the island after periods of strong westerly winds. 

All birds, except the grackle and the swallow, 

roost eleswhere, most likely on South Water Cay. 

The grackle may even breed on Carrie Bow Cay 

because a female was observed gathering materials for nest building. 

INTERTIDAL ORGANISMS.—The mean tidal range 

at Carrie Bow Cay is only 15 cm (Kjerfve et al., 

herein: 47, Table 4). The observed maximum 

range, however, partly because of wind forcing is 

more than 40 cm. With a shoreline slope of 90° 

to 4° the width of the intertidal zone on Carrie 

Bow Cay ranges between 40 cm on vertical cinder 

block walls and 6 m at the flat northern point, on 

the average between 0.5 and 2.0 m. Only during 

spring tides are wide areas on the reef flat exposed 

(Plate 1: bottom right). 

Sandy beaches have a diverse and rich intertidal meiofauna (Kirsteuer, in prep.) but only one 
NUMBER 12 83 

benthic macro-organism, the cerianthid Arachhanthus nocturnus den Hartog, could be observed at 

low tide buried in exposed sand on the northeast 

shore. Ocypode quadrata crabs, however, temporarily establish burrows in sand areas exposed at low 

tide. 

Rocky substrates support a more varied intertidal flora and fauna but differences in abundance can be observed between the leeward (west) 

and windward (east) sides of the island. Coral 

rock, rubble, and concrete blocks of the leeward 

sea wall are thickly covered by algae (Figures 

52b, 54a). Oscillatoria submembranacea Ardissone 

and Strafforella and Schizothrix mexicana Gormont 

(Cyanophyta), and Cladophoropsis membranacea (C. 

Agardh) B0rgesen (Chlorophyta) occupy the upper zone, Padina Jamaicans (Collins) Papenfuss 

(Phaeophyta) and Neomeris annulata Dickie (ChloFIGURE 54.—Intertidal organisms; a, algae Cladophoropsis and 

Oscillatoria (top), Neomeris and Padina (bottom) on concrete 

block seawall; b, gastropod Liltorina ziczac on coral boulder. 

(Picture width, a = 70 cm; b = 5 cm.) 

rophyta) the zone below. On the windward side 

only the calcareous green alga Halimeda opuntia 

(L.) Lamouroux, red Laurencia papillosa (Forsskal) 

Greville, and some of the Oscillatoria were found 

exposed. 

A few specimens of the actinian Stoichactis anemone (Ellis) and barnacle Tetraclita stalactifera (Lamarck) were also encountered on the windward 

side. The most abundant crustaceans on the lagoon shore are the hermit crab Clibanarius tricolor 

(Gibbes), which clusters in great numbers on 

intertidal rock and rubble, and the elusive isopod 

Ligia olfersii Brandt, which is particularly common around empty conch shells near the concrete 

boat dock. Several crabs are common among 

rubble and concrete blocks all around the cay. 

Grapsus grapsus L. is the largest and most abundant; other crabs include Pachygrapsus transversus 

(Gibbes), Cyclograpsus integer Milne Edwards, and 

Petrolisthes quadratus Benedict. Among the mollusks only gastropods occur intertidally at Carrie 

Bow Cay. On the windward side Nerita peloronta 

L., N. versicolor (Gmelin), Littorina nebulosa Lamarck, L. ziczac (Gmelin) (Figure 54b), and Tectarius muricatus (L.) are found on vertical coral 

rock and concrete block surfaces of the seawall. 

Several size classes of juvenile Cittarium pica (L.) 

cluster among rubble or on beach rock below. 

Nerita versicolor, L. ziczac, and T muricatus also 

occur on the leeward seawall but are less abundant there. A few specimens of a single species of 

echinoderm, the echinoid Echinometra viridis Agassiz, are found here and there under tide-exposed 

rocks. 

Climatic Parameters 

The climate of Belize is subtropical to tropical, 

with temperatures ranging from 10° to 36° C 

(average range in Belize City, 23°-33° C), and 

rainfall averaging 125-450 cm a year. Temperatures are lowest in the highlands and during the 

cool period of the year (November to March). 

Average rainfall increases from north to south; 

the rainy season lasts from June to October. The 

overall climate of the country, particularly of the 

outer cays, is influenced by northeasterly trade 
84 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

winds that prevail at velocities of 4-5 m/s during 

about 70% of the year. Our meteorological records from Carrie Bow Cay, although not continuous, indicate major patterns of temperatures, 

solar radiation and cloudiness, wind, and rainfall 

and allow some comparisons with conditions prevailing on the mainland. In addition, 12-day 

continuous measurements of radiation, evaporation, wind, and air-water-sand temperatures in 

June 1975 were reported by Kjerfve (1978). 

TEMPERATURE AND SOLAR RADIATION.—Figure 

55 presents monthly temperature records, except 

for July and December. Data for January, February, and August to October are the result of a 

single year's readings; other data were derived 

from at least three consecutive years of observation. Values are plotted against a background of 

ten-year average minimum and maximum temperature readings provided by the Melinda Forest 

Station near Dangriga, on the mainland of Belize. 

Temperature conditions on the cay follow closely 

those on land, where the highest averages occur 

during May and August (33° C) and the lowest 

during January and February (22° C, 21 C). 

Solar radiation measurements are only available 

for the cay and for the months of March through 

June, and November. The highest total radiation 

reaching the ground on a single day was recorded 

during April and May and amounted to 490 cal/ 

cm2. Monthly averages of daily radiation related 

to this value give an indication of cloudiness and 

haze (Figure 55). 

WIND.—Measurements of wind direction, 

speed, and frequency on Carrie Bow Cay are 

summarized in Figures 56 and 57. Values for 

March-June show the typical situation: northeasterly trade winds predominate and compare well 

with published wind roses from the open ocean 

surface off Belize (United States Naval Oceanographic Office, 1963). Our observations on wind 

for the rest of the year are sparse and may not be 

representative of long-term averages. Speed values for the infrequent winds from the northwest 

sector are somewhat low because of the shading 

effect of the big house and of coconut trees. 

RAINFALL AND HUMIDITY.—Long-term rainfall 

°c 

40 -, 

35 -

30 -

/83% \ /86% \ /77% \ f70%\ (£) {JJ {£) (j^) 

25 

20 -

Id 

A l l 

n M 

( ^ 

m 

i- 40 

- 35 

- 30 

- 25 

- 20 

1 1 1 I I I I I I I 1 

JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC 

'Ai r • Sand A Reef Flat Lagoon W!\ Mainland 

FIGURE 55.—Monthly temperatures (mean, range) 1976-1980 and solar radiation (percentage 

of maximum) 1978-1980 for Carrie Bow Cay; monthly temperature range (shaded area) at 

Melinda Forest Station on mainland, averaged over a ten-year period, 1965-1975. 
NUMBER 12 85 

Jan /Feb 

(25) 

40.6 

13.8 

• 20 

30.7 

<=- ( 371 

Jun 

(98) 

12.2 5.| >vl4.3 

43.9 

28.8 

Oct/Nov 

(66) 

19.7 

FIGURE 56 (above).—Wind roses for Carrie Bow Cay indicating direction, speed, and frequency (figures in parentheses 

are numbers of observations during 1976-1980). 

FIGURE 57 (right).—Monthly summaries of wind speed frequencies during 1976-1980, Carrie Bow Cay. 

data taken at the Melinda Forest Station indicate 

an average annual accumulation of 218 cm for 

the Dangriga district. The range is from 4.4 cm 

in March to 30.4 cm in September. Values for 

Carrie Bow Cay are presented in Figure 58 and 

compared with the mainland averages. The is36.6 

140.8 

Legend 

1-5 6-8 9-14 >I4 m/s 

-c 

i r 

20 

— i 1 1 

40 60% 

Frequency 

Legend 

/ColrnX 

( ~i 

0 

i 

20 

-5 

Speed 

6 -8 

1 • 

Frequency 

1

 1 ' 

40 ' 60 

9- 14 > I 4 IT 

80 

i 

/s 

100% 

Jan/Feb 

Mar & 

Apr 

May 

Jun 

Oct/Nov 

^ 

I I I 1 L_ I I I I I 

20 40 60 80 100% 
86 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

mm 

300 

250 -

200 -

150 

100 -

50 -

0 £*d 1. Wm T: W. w 

-30 0 

- 250 

200 

M M N 

- 150 

i-100 

- 50 

0 

0 Daily Maximum Mainland 

FIGURE 58.—Monthly average (total bar) and daily maximum rainfall on Carrie Bow Cay 

(1976-1980), compared with mainland monthly rainfall (Melinda Forest Station) averaged 

over a 71-year period (1906-1977). 

land receives, on the average, only 42 prcent of 

the mainland rainfall if one excludes February 

and December for which comparative data are 

lacking. The high value (279 mm) for November 

may be a peculiarity of the year (1979) in which 

the record was taken. On the other hand, a second 

but incomplete measurement of 213 mm (1978, 

November 14-26) indicates a similarly high or 

even higher rainfall during that month. 

Humidity measured between March and June 

averaged 78 percent, with a range of 58-96 percent. 

Recent Hurricane Effects on Carrie Bow Cay 

Computer files of the United States National 

Hurricane Center (P. J. Hebert, pers. comm.) 

indicate that at least 20 hurricanes and 45 tropical cyclones have passed within 100 nautical miles 

(185 km) of Belize City (17°30'N5 88°18'W) during the last century (records date back to November 1889). From these data it can be determined 

that nine hurricanes and seven tropical storms 

have passed Carrie Bow Cay within a 50 km 

radius. Storm activity in this area seems to have 

increased recently as six of the hurricanes and the 

most violent of tropical storms (Laura) have occurred within the last 20 years (Table 8). 

Hattie is the only storm for which the longterm effects on Belizean reefs and cays, including 

Carrie Bow Cay, have been monitored (Stoddart, 

TABLE 8.—Hurricanes passing within 50 km radius of Carrie 

Bow Cay, 1960-1980, including name, date, and maximum 

sustained wind speed while storm center was within 50 km 

of Carrie Bow Cay 

Name 

Abby 

Anna 

Hattie 

Francelia 

Laura* 

Fifi 

Greta 

Month/ Year 

Jul 1960 

Jul 1961 

Oct 1961 

Aug 1969 

Nov 1971 

Sep 1974 

Sep 1978 

Wind speed 

(km/h) 

128 

148 

259 

182 

111 

176 

176 

Officially declared a tropical storm. 
NUMBER 12 87 

1963, 1969, 1974). Other recent hurricane reports 

include a brief eyewitness account of tropical 

cyclone Laura passing over Glover's Reef and 

Stann Creek (Dangriga) (Antonius, 1972), and 

observations on the impact of hurricane Greta on 

the reef community near Carrie Bow Cay (Highsmith et al., 1980). 

HURICANE FIFI (14-22 September 1974).—A 

tropical depression south of Puerto Rico and 

Hispaniola moving westward developed into hurricane Fifi on 17 September. Fifi, as reported by 

Hope (1975), acquired its maximum sustained 

winds of 95 kt (176 km/h) while it moved along 

the coast of Honduras, 18-19 September, where 

heavy rains caused a high number of deaths by 

inland flooding of rivers. The hurricane crossed 

the barrier reef approximately 20 km south of 

Carrie Bow Cay and reached the coast of southern 

Belize during the afternoon of 19 September. 

Our observations on the effects of Fifi on Carrie 

Bow Cay rely on a survey in December 1974 as 

no eyewitness reports are available. Storm surge 

flooded the entire island, and most of the unconsolidated sand was either piled up high inside the 

buildings or carried away, leaving a surface of 

coral rubble and exposed palm tree roots. A 

comparison of photographs (Figure 59) and a 

map of Carrie Bow Cay prepared by D. R. Stoddart in 1972 (Figure 60a) indicate that coastal 

erosion was strongest to the north, northeast, and 

%JE w 

mtt •** 

FIGURE 59.—Carrie Bow Cay silhouettes looking east: a, February 1972; b, December 1974, 

three months after hurricane Fifi. Note reduction of island size, in number of trees, and density 

of leaves caused by the hurricane. 
SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

a 

Old New 

AAAA 

Mean Tide Level 

Undercut Shoreline 

Submerged Sand Spit 

Coconut Palms 

• Live 

o Planted after Fif 

& Stump 

FIGURE 60.—Maps of Carrie Bow Cay showing hurricane effects and poststorm recovery: a, 

changes 1972-1974, caused principally by hurricane Fifi, September 1974; b, recovery, December 1974 to May 1978. 
NUMBER 12 89 

»-*» 

FIGURE 61.—Effects of hurricane Fifi: a, eroded north point of Carrie Bow Cay with fallen 

coconut palms, looking northeast; b, submerged bank of sand originating from northern portion 

of the island, looking northwest. 
90 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

south of the island. About 30% of the island's 

surface area was lost, but some of it was regained 

by subsequent deposition along the western 

shoreline. Most island sand, however, settled as a 

subtidal sand bank to the northwest of the cay 

(Figure 61 b). The eastern seawall and outhouse 

dock were destroyed and a huge tree stump, 

situated for years on the reef flat to the east, was 

floated to the new south tip of the cay. At least 

14 coconut trees, predominantly from the north 

point and northeast shore were uprooted by erosion and either fell in place (Figure 61a) or were 

carried into the lagoon and sank. Others lost their 

tops in the storm or withered from overexposure 

to salt water. All other plants previously recorded 

(Stoddart, 1969), such as low Tournefortia bushes, 

Euphorbia, Ipomoea, and Sesuvium ground cover, and 

grasses, disappeared and did not recover to the 

approximate prehurricane condition until spring 

1978. By that time, with the help of seawalls and 

rubble fills, much of the prehurricane island outline was restored (Figure 60b). 

HURRICANE GRETA (13-23 September 1978).— 

The track of Greta was almost identical to that of 

Fifi, and both storms occurred at almost the same 

time in September. The meterorological history 

of Greta is described by Lawrence (1979). A 

depression formed northwest of Trinidad on 13 

September. Hurricane force with sustained winds 

of 115 kt (213 km/h) developed at a position 

south of Jamaica on 16 September. Moving over 

the Bay Islands off the north coast of Honduras, 

Greta weakened and made its landfall with 80 kt 

(148 km/h) winds near Dangriga on the evening 

of 19 September. Greta was a much more severe 

hurricane than Fifi but despite locally heavy rain 

it did not cause devastating river floods (P. J. 

Hebert, pers. comm.). 

The eye of hurricane Greta passed Carrie Bow 

Cay about 6 km to the north and brought the 

island winds of approximately 95 kt (176 km/h). 

Although direct observations are lacking, a considerable storm tide (about two meters above 

normal, estimated from events at Dangriga) must 

have flooded the island because the smallest cottage disappeared and the ocean-side wall of the 

laboratory caved in. Despite damage to buildings, 

coastal erosion was considerably lower than during Fifi. About 20 coconut trees were lost, most of 

these from the leeward side of the island. Other 

plants were much less affected by Greta than by 

the 1974 hurricane as only one common and four 

minor species disappeared (Table 7). 

Summary and Conclusions 

From its position, structure, and flora, Carrie 

Bow Cay can be classified as a reef-derived sand 

cay. It is located at the seaward margin of the 

barrier reef, is composed of reef rubble and sand, 

and is held together primarily by coconut rootlets 

and, to a lesser degree, by ground cover and 

artificial structures. The island measured a little 

over two acres (0.8 ha) when it was bought by 

the present owners in 1943. Today it is less than 

half that size and exposed beachrock on the windward side indicates westward (leeward) migration, which confirms the view that sand cays of 

this nature are slowly migrating sand waves (Milliman, 1973). 

Because of its small size, low elevation, and 

porous substrate, Carrie Bow Cay lacks a freshwater lens and it has not developed a complex 

terrestrial environment. Considering the occasional salt water flooding during storm tides, the 

island may be described as a supralittoral habitat. 

The climate, too, is dominated by the surrounding ocean and by northeasterly trade winds; it is 

moderate in comparison with the nearest mainland. 

Clearing of vegetation during this century and 

increased hurricane activity in the area during 

the past two decades are mainly responsible for 

the rapid shrinking of Carrie Bow Cay. Captain 

Owen, who mapped the island as "Jack Ellin's 

Cay" in 1830, noted "tops of bushes 20 feet" 

(Stoddart, 1963), presumably seagrape, baycedar, 

and mangrove. Coconuts planted in the early 

1900s, and repeatedly again since, may not be 

equally effective in holding the sand, also, they 

do not protrude into the intertidal to trap sediments or break the power of waves or currents. 
NUMBER 12 91 

Although physiographic change of the cay was 

minor during hurricane Hattie (Stoddart, 1963, 

1969), later storms, Fifi in particular, took severe 

toll. Recovery of plant cover destoyed by Fifi took 

about four years. Colonization of sand cays is 

thought to be primarily by floating seeds or by 

seeds carried by birds (Stoddart, 1960), or by 

wind, but direct observations on these processes 

are sparse. Possible means of dispersal judged 

from seed type are listed in Table 7 (data provided by M.-H. Sachet). Our own findings suggest that only four of 16 species of plants—coconut, red mangrove, and two unidentified seedlings—arrived by sea and sprouted. None of them 

survived beyond two years because of the unsuitable location of settlement. Experimental studies 

on natural means of island colonization should be 

the next step in elucidating the terrestrial development of Carrie Bow Cay. 

Literature Cited 

Antonius, A. 

1972. Hurricane Laura, Witnessed in British Honduras. 

Atoll Research Bulletin, 162:11-12. 

Highsmith, R. C , A. C. Riggs, and C. M. D'Antonio 

1980. Survival of Hurricane-generated Coral Fragments 

and a Disturbance Model of Reef Calcification/ 

Growth Rate. Oecologia (Berlin) 46:322-329. 

Hope, J. R. 

1975. Atlantic Hurricane Season of 1974. Monthly 

Weather Review, 103:285-293. 

Kirsteuer, E. 

In prep. Horizontal and Vertical Migrations of Ololyphlonemertes (Nemertinea) in a Sandy Beach at Carrie 

Bow Cay, Belize. 

Kjerfve, B. 

1978. Diurnal Energy Balance of a Caribbean Barrier 

Reef Environment. Bulletin of Marine Science, 28: 

137-145. 

Lawrence, M. B. 

1979. Atlantic Hurricane Season of 1978. Monthly 

Weather Review, 107:477-491. 

Milliman, J. D. 

1973. Caribbean Coral Reefs. In O. A. Jones and R. 

Endean, editors, Biology and Geology of Coral Reefs, 

1:1-50. New York and London: Academic Press. 

Stoddart, D. R. 

1960. The Reefs and Sand Cays of British Honduras. 

In Cambridge Expedition to British Honduras, 1959-

1960, General Report, pages 16-22. Cambridge, 

England. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays. October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 

1969. Post-Hurricane Changes on the British Honduras 

Reefs and Cays: Re-survey, 1965. Atoll Research 

Bulletin, 131:1-31. 

1974. Post-Hurricane Changes on the British Honduras 

Reefs: Re-survey, 1972. In A. M. Cameron et al., 

editors, Proceedings of the Second International Coral 

Reef Sympsoium, 2:473-483. Brisbane, Australia: 

The Great Barrier Reef Committee. 

United States Naval Oceanographic Office 

1963. Oceanographic Atlas of the North Atlantic Ocean, 

Section IV: Sea and Swell. United States Naval 

Oceanographic Office, Publication, 700: 227 pages. 

Washington, D.C. 
Distribution of Microborers within Planted 

Substrates along a Barrier Reef Transect, 

Carrie Bow Gay, Belize 

Jeffrey A. May, Ian G Macintyre, 

and Ronald D. Perkins 

ABSTRACT 

A diverse assemblage of endolithic microorganisms was identified in a series of carbonate substrate stations planted along a transect across the 

barrier reef near Carrie Bow Cay, Belize. These 

tropical endoliths at the sediment-water interface 

include the cyanophytes, Hyella tenuior Bornet and 

Flahault, Mastigocoleus testarum Lagerheim, and 

Plectonema terebrans Bornet and Flahault, the chlorophytes, Ostreobium brabantium Weber Van-Bosse 

and Phaeophila engleri Reinke, the rhodophyte Porphyra sp. (Conchocelis-phase), and various fungi. 

This assemblage was subdivided into an upper 

photic zone assemblage dominated by Mastigocoleus, Hyella, Phaeophila, and Ostreobium species, and 

a lower photic zone assemblage dominated by 

Porphyra sp. 

Subsurface endolithic activity detected at the 

shallow lagoon station included filamentous irregular polygonal networks, irregular flattened 

masses, and regular crenulate discoids, which differed from and were less diverse than the assemblage at the sediment-water interface. Affinities 

of these subsurface microborings are unknown 

but they resemble endolithic traces and organic 

scars variously attributed to fungi, bacteria, and 

Actinomycetes. The regular discoids and irregular 

masses occurred only in association with the filamentous form, and therefore may be related reproductive bodies. For reasons not fully underJeffrey A. May, Department of Geology, Rice University, Box 1892, 

Houston, Texas 77001. Ian G. Macintyre, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560. Ronald D. Perkins, Department of 

Geology, Duke University, Box 6665, College Station, Durham, 

N. C 27708. 

stood, microborings were not present in the second subsurface station, in fore-reef sand at a 

depth of 24 m. 

Introduction 

Endoliths are microorganisms (generally less 

than 1 jtim to 100 jum in diameter) that penetrate 

calcareous substrates by chemical and/or mechanical means and that leave post-mortem microscopic networks. They are distinguished from 

"epiliths," which live only on a substrate's surface, and from "chasmoliths," which adhere to 

surfaces of fissures or cavities within the substrate 

(Golubic et al., 1975). Endoliths include cyanophytes, chlorophytes, rhodophytes, fungi, and 

possibly bacteria and sponges. 

The most diverse and ecologically important 

microborers occur in the marine setting; their 

boring patterns—size, mode of branching, spatial 

arrangement, and growth directions—are taxonomically characteristic (Golubic, 1972). The marine endoliths have been subdivided on the basis 

of bathymetric and regional assemblage distributions, which are controlled by geographic, climatic, photic, and environmental factors (Perkins 

and Halsey, 1971; Perkins, 1972; Rooney and 

Perkins, 1972; Golubic et al., 1975; Green, 1975). 

Various studies of ancient and modern forms 

have indicated that endoliths may be used to 

interpret paleoclimatic conditions and to recon93 
94 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

struct depositional environments (for example, 

see Swinchatt, 1965; Gatrall and Golubic, 1970; 

Perkins and Tsentas, 1976). As well, microboring 

organisms have been used to establish the positions of relict shorelines (Perkins and Halsey, 

1971; Edwards and Perkins, 1974) and to detect 

sediment transport (Rooney and Perkins, 1972). 

Microborers modify both lithified and unlithified carbonate coasts (Purdy and Kornicker, 

1958; Hodgkin, 1970; Schneider, 1976), in the 

colonization of shifting upper sublittoral substrates—where they create finer carbonate sediments, preferentially remove certain components, 

and initiate micrite rind formation (Bathurst, 

1966; Golubic, 1969; Alexandersson, 1972; Rooney and Perkins, 1972; Perkins and Tsentas, 

1976)—and in their alteration of the deep-sea 

sedimentary record (Zeff and Perkins, 1979). Endolithic algae not only dissolve carbonates, but 

also induce precipitation of calcium carbonate 

within shallow marine corals (Schroeder and 

Ginsburg, 1971; Schroeder, 1972a, b; Scherer, 

1974). Furthermore, their biologically related 

physico-chemical processes may influence lowtemperature sedimentary mineralization within 

boring networks (Taylor, 1971; Kobluk and Risk, 

1977). 

Recently, planted substrates have been used to 

identify modern microboring assemblages and to 

establish rates of infestation (Golubic, 1969; 

LeCampion-Alsumard, 1975; Perkins and Tsentas, 1976). The present investigation examined 

and identified endoliths within planted substrates 

in order to determine the distributional patterns 

of these organisms at the sediment-water interface 

along a reef transect. Two stations planted below 

the sediment surface allowed a cursory examination of subsurface microboring activity. 

ACKNOWLEDGMENTS.—The authors thank M. 

R. Carpenter (Smithsonian Institution) and 

R. G. Zingmark (University of South Carolina) 

for field assistance, ar\d G. W. Lynts and R. B. 

Searles (Duke University) for helpful comments 

on the manuscript. Support for part of this research was provided by National Science Foundation Grant OCE74-12555 A01 to Duke University. 

Methods 

This study examined two types of carbonate 

substrates planted at and below the sedimentwater interface along a reef transect of Carrie 

Bow Cay (Figure 62): (1) crushed, fresh inner 

shell parts of the queen conch, Strombus gigas 

Linnaeus, and (2) cleaved calcite rhombohedra. 

Samples of each type were retained as controls. 

Fragments from 1 to 10 mm in size were attached 

to 15 cm2 plexiglass plates (one type per plate) 

and 40 cm long polyvinyl chloride (PVC) pipes 

(types mixed) by a thin film of epoxy resin. 

Fourteen substrate-covered plexiglass plates were 

mounted on short lengths of protruding PVC 

pipe to maintain the samples above shifting substrata (Figure 63). These plates were placed in 

pairs at seven locations extending from the lagoonal Thalassia zone (depth 1.2 m) to the forereef slope (depth 27.4 m). One subsurface pipe 

station (consisting of one pipe) was inserted into 

the sea floor in the Thalassia zone (Figure 64) and 

another into the Halimeda-rich sand of the forereef sand trough (depth 24 m). Exposure time of 

substrates ranged from 21 to 24 months. 

After being harvested, the planted substrates 

were preserved in 4 percent formaldehyde in 0.1 

M phosphate neutral buffer. Fragments intended 

for light microscopic study were carefully scraped 

to remove epilithic organisms, then dissolved with 

5 percent EDTA solution (van Reine and van 

den Hoek, 1966) at a pH of 6. Although the threedimensional configuration of the microborers is 

lost because of their collapse, organic structure 

and color are not damaged by this slow-dissolving 

solution. Extracted tissues were then mounted on 

glass slides. 

Scanning electron microscopic (SEM) analysis 

was based on the casting-embedding technique of 

Golubic et al. (1970). An alcohol dehydration 

series was followed by an infiltration series using 

Durcupan ACM Araldite Base Embedding 

Agent. After polymerization within plastic holders, the substrate fragments were exposed by 

means of a rotary grinding tool, then etched with 

3 percent hydrochloric acid. This technique revealed plastic casts of the microboring networks, 
NUMBER 12 95 

DEPTH AND 

tOCATION | 2 m 

OF SUBSTRATE ' 

STATIONS ft ft 

SPORANGIAL STRUCTURES 

( Fungal filaments found in all substrate stations 

FIGURE 62.—Idealized cross section of Carrie Bow Cay transect showing locations of substrate 

stations planted at sediment-water interface and the distribution of endoliths collected from 

these plates (hatched squares = great abundance; hatched rectangles = presence observed). 

which retained the original spatial relationships 

of the endoliths and their three-dimensional configurations. The mounted plastic blocks were 

scanned with an International Scientific Instruments Super II electron microscope after vacuum 

shadowing with gold-palladium alloy. Casts of 

boring networks were correlated with the endoliths isolated by acid dissolution. 

Results 

ENDOLITHS AT THE SEDIMENT-WATER INTERFACE.—Blue-green algae were ubiquitous and 

consisted of Hyella tenuior Bornet and Flahault, 

Mastigocoleus testarum Lagerheim, and Plectonema 

terebrans Bornet and Flahault. Green algal microborers, likewise abundant, included Ostreobium brabantium Weber Van-Bosse and Phaeophila engleri 

Reinke. Red algae were much less abundant and 

were represented only by the Conchocelis-stage of 

Porphyra sp. Fungal forms were found in almost 

all samples. 

Plectonema terebrans was the most common cyanophyte at all stations. Diagnostic are its 

smooth, elongate, thread-like filaments 2 to 4 jum 

in diameter, which may run along the interior 
96 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 63.—Patch reef zone substrate station, Carrie Bow Cay, showing location of plates 

above the sediment-water interface. 

FIGURE 64.— Thalassia-zone subsurface substrate station, Carrie Bow Cay, showing a closeup of the buried substratecovered 40 cm long PVC pipe. Note that only the protective 

collar and a small portion of the pipe protrude above the 

sediment-water interface. 

surface of the substrate or may form dense, interwoven meshworks (Figure 65a,b). Also ubiquitous 

was Mastigocoleus testarum, which is composed of 

sharply curved to elongate filaments 5 to 8 /xm in 

diameter that have numerous short lateral 

branches and heterocysts (Figure 65c,d). The morphology of M. testarum penetrating inorganic substrates is much more affected by the rhombohedral microstructure than is the morphology of 

P. terebrans; this observation corresponds with 

findings of LeCampion-Alsumard (1975). Hyella 

tenuior, which is less common, appears as a cluster 

of slender, elongate, relatively straight to bent 

filaments 5 to 8 /xm in diameter (Figure 66a,b). 

These filaments grow subperpendicular to the 

surface of the substrate. 

FIGURE 65.—Scanning electron and transmitted light photomicrographs of endolithic algae: a, Plectonema terebrans forming a typical dense network of filaments, acid-etched mollusk 

fragment, Thalassia zone (note the smooth, elongate, fine 

nature of the plastic casts); b, P. terebrans isolated by dissolution of a mollusk fragment, reef-crest zone; c, characteristic 

heterocyst development of Mastigocoleus testarum shown on 

plastic casts, acid-etched mollusk fragment, Thalassia zone; 

d, M. testarum isolated by dissolution of a mollusk fragment, 

Thalassia zone (note the heterocyst development marked by 

arrows). (Scale = 50 /xm for a, d; 40 jLtm for b; 25 fim for c.) 
NUMBER 12 97 
98 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

1 

( j <^ 

^ 9 k i ' v^*vi ;>' ** 

d 
NUMBER 12 99 

Phaeophila engleri was the most abundant and 

widespread chlorophyte. This species is characterized by rectilinear branching and pronounced 

bulbous or irregularly ovoid, 15 to 20 jum swellings at points of branching or along the irregular 

5 to 10 /im filaments (Figure 66c,d). The largest 

boring species detected, Ostreobium brabantium has 

digitate growths of single or bilobate branches 

(Figure 67a,b). Single plants up to 1 mm in length 

radiate into the substrate; individual branches of 

40 to 60 jum may enlarge up to 120 /xm before 

bifurcation. The Conchocelis-stage of the rhodophyte Porphyra sp. is characterized by rectilinear 

branching of long and fine, 2 to 3 /xm filaments 

running along slightly beneath the substrate surface (Figure 67 c,d). 

Extremely fine filaments from less than 1 up to 

4 /xm in diameter occur in a wide variety of forms, 

from non-branched to extensively branched and 

fused, sparse to massive networks (Figure 68a,b). 

These filaments probably represent fungal hyphae. The hyphae appear to be directed towards 

algae and can be observed penetrating these organisms, presumably in the act of feeding (Figure 

68b). Three different types of structures observed 

with the scanning electron microscope were attributed to fungal spore cases. Form A has 5 to 

15 /xm ovoid to pyriform reproductive bodies, 

from the bases of which radiate long and threadlike, i to 2 jum hyphae (Figure 69a). These hyphae 

are typically unbranched and connect spore cases. 

Form B has 8 to 20 /xm, globose to oblong fruiting 

bodies with connective hyphae attached at their 

FIGURE 66.—Scanning electron and transmitted light photomicrographs of endolithic algae: a, elongate casts of the 

blue-green alga Hyella tenuior directed away from the substrate surface, some branching near their bases, plastic cast 

within an acid-etched calcite rhomb, Thalassia zone (very 

little control is exerted by the substrate microstructure); b, 

filaments of//, tenuior displaying elongate, somewhat rectangular cells, isolated by dissolution of the molluscan substrate, 

Thalassia zone; c, green alga Phaeophila engleri exhibiting 

characteristic ovoid swelling at points of branching and 

rectangular branching pattern, plastic cast of etched mollusk 

fragment, Thalassia zone; d, P. engleri demonstrating probable 

sporangia (arrows) and swellings at branching points, filaments isolated by dissolution of a molluscan substrate, Thalassia zone. (Scale = 50 ju,m for a; 25 iim for b, c; 40 ttm for d.) 

sides, so that these spore cases appear positioned 

along filaments (Figure 696). The irregular hyphae of less than 1 jum diameter become branched 

and fused, and form interconnected networks. 

Fungal form C has 5 to 15 /xm globose to discoid 

sporangial bodies with indented ends that appear 

"doughnut-shaped" (Figure 68c). These bodies 

occur alone or in clusters directly upon the surface 

of endolithic algae; connective hyphae are lacking. 

ENDOLITHS BELOW THE SEDIMENT-WATER INTERFACE.—In contrast to the diverse assemblage of 

microorganisms boring into the substrates 

planted at the sediment-water interface, only a 

few endolithic forms were found below the surface. These boring organisms were present only 

in the molluscan fragments planted within the 

lagoonal Thalassia zone. Classification is problematic as these forms have not previously been 

described. No microboring activity was recorded 

at the subsurface station in the fore-reef sandtrough zone. 

An extremely irregular, polygonal network of 

variable and intermittent filaments 5 to 7 /xm in 

diameter (Figure 70a,b) commonly crosses the 

regular and parallel lines representing remnants 

of the organic matrix that separated the inorganic 

crystals of the gastropod shell. This network of 

variously spaced filaments bores just below the 

surface of the molluscan substrates. The filaments 

are curvilinear, have a twisted appearance, and 

branch sideways. This form is recurrent in approximately 10 percent of the molluscan fragments throughout the pipe planted in the Thallasia zone, and was not found in the control 

samples or in samples from the sediment-water 

interface. Possibilities for taxonomic assignment 

include marine fungi (Phycomycetes, Ascomycetes, and Deuteromycetes) and filamentous bacteria (Actinomycetes). 

Another group occurs as regular patches or 

regular crenulate discs associated only with the 

irregular polygonal networks described above 

(Figure 70c,d). These may be reproductive structures or separate endolithic forms; they range 

from 125 to 200 /xm in diameter and average 2 

/xm in thickness. Although they may be remnants 
100 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Ms©!;** - * 
NUMBER 12 101 

FIGURE 68.—Scanning electron photomicrographs of endolithic fungi: a, little-branched, thin 

plastic casts believed to be fungal hyphae, intertwined with the larger blue-green alga Plectonema 

terebrans, etched mollusk fragment, Thalassia zone; b, network of fine fungal borings covering 

and possibly feeding (arrows) upon the underlying alga, plastic casts within an etched mollusk 

substrate, Thalassia zone. (Scale = 5 /xm for a; 25 /xm for b.) 

of the organic matrix, no analogous structures 

were found in any control sample or in any 

sample infested at the sediment-water interface. 

FIGURE 67.—Scanning electron and transmitted light photomicrographs of endolithic algae; a, large, radiating growth 

form of the green alga Ostreobium brabantium, plastic cast 

within an acid-etched mollusk fragment, Thalassia zone (note 

both the single and bilobate branches, background forms are 

algae and fungi); b, 0. brabantium filaments isolated by dissolution of a molluscan substrate, patch-reef zone; c, characteristic rectilinear pattern of the Conchocelis-stage of the red 

alga Porphyra sp., plastic cast, enhanced on a microscale by 

boring within a calcite rhomb, fore-reef slope zone (larger 

forms are an unidentified alga with strong microstructural 

control upon its boring pattern); d, Conchocelis-stage of Porphyra sp. displaying fine filaments in the typical rectilinear 

pattern, isolated by dissolution of a molluscan substrate, 

fore-reef slope zone. (Scale = 200 /xm for a; 100 jum for b; 25 

/xm for c, d.) 

Discussion 

ENDOLITHS AT THE SEDIMENT-WATER INTERFACE.—The bathymetric distribution of Belizean 

endoliths was compared with similar tropical microboring assemblages recovered from artificial 

substrate stations planted in reefs off St. Croix 

and Jamaica (Green, 1975; Perkins and Tsentas, 

1976). The distribution of autotrophic endolithic 

organisms is related to light penetration in the 

sea—both the intensity of illumination and spectral composition (Golubic et al., 1975). Although 

endolithic organisms cannot be assigned to absolute depths—owing to variations in water clarity, 

currents, and other environmental factors—Perkins and Tsentas (1976) pointed out that "clearwater" assemblages might be used to estimate 

maximum depths for endolithic algae. Their di-
102 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 69.—Scanning electron photomicrographs of endolithic fungi: a, form A, 

characterized by ovoid to pyriform reproductive bodies, distributive unbranched 

casts of hyphae radiate from the base of 

the fruiting bodies just below the surface 

of the substrate, plastic cast of an etched 

calcite rhomb, fore-reef slope; b, reproductive bodies of fungal form B laterally 

connected by hyphae, etched mollusk 

fragment, Thalassia zone (these bodies are 

typically globose to oblong, note twisted 

or irregular appearance of the hyphal 

casts); c, "doughnut-shaped" or indented 

sporangial bodies of fungal form C, plastic cast in an etched mollusk substrate, 

Thalassia zone (note that these occur in 

close proximity to the outer substrate surface or to algal borings). (Scale = 10 /xm 

for a, b; 25 /xm for c.) 
NUMBER 12 103 

vision of such assemblages into an upper photic 

zone of Mastigocoleus, Hyella, Phaeophila, and Ostreobium species, and a lower zone dominated by 

Porphyra sp. in its Conchocelis-phase parallels, to 

some extent, the zonation found in the present 

study (Figure 62). 

Ostreobium brabantium was observed only in the 

shallowest (1.2 and 1.5 m) sites of the Thalassia 

and patch-reef zones off Belize. In St. Croix, 

although present to 30 m, 0. brabantium was predominant in depths less than 15 m (Perkins and 

Tsentas, 1976). Also, only the three shallowest 

sites off Belize contained Hyella tenuior, which was 

found down to 45 m in Jamaica (Green, 1975). 

Four species of algae occurred at all depths 

along the Carrie Bow Cay transect. Phaeophila 

engleri was most abundant from 1.2 m to the 7.6 

m spur and groove zone of the inner fore reef; 

similarly, this species is very common in the 

shallower zones of St. Croix and Jamaica. Mastigocoleus testarum was most abundant in the upper 

12 m of the Belize sites, as was the case in St. 

Croix. Off Belize, however, it decreased slightly 

in the reef-crest and Thalassia zones. Plectonema 

terebrans was ubiquitous off Belize as well as Jamaica and St. Croix, except that in Belize it was 

less abundant in the Thalassia and reef-crest 

zones. Patchily distributed below 1.5 m, Porphyra 

sp. was most common in depths of 12 to 27 m, 

the deepest zone examined off Belize. This pattern has also been reported from the Australian 

Barrier Reef (Rooney and Perkins, 1972), the 

Puerto Rico shelf (Perkins, 1972), and Woods 

Hole, Massachusetts (Golubic et al., 1975), as 

well as Jamaica and St. Croix (Green, 1975; 

Perkins and Tsentas, 1976). 

Although thin filaments believed to be fungal 

hyphae were present at all depths, the distribution of different sporangial structures was depth 

dependent. Fungal form A, with pyriform to 

ovoid bodies, from which basal hyphae radiate, 

was found only in the sample from 27 m on the 

fore-reef slope. Similarly, Perkins and Tsentas 

(1976) found a reticulate fungal form at 30 m in 

St. Croix. Forms B and C with laterally arranged 

globose spore cases and isolated "doughnutshaped" bodies, respectively, were found only in 

the shallow reef-crest, patch-reef, and Thalassia 

zones off Belize. 

ENDOLITHS BELOW THE SEDIMENT-WATER INTERFACE.—Fungi may be as important as bacteria in 

breaking down organic matter into a nutrient 

source for other organisms. Marine fungi may be 

as versatile and potent in their feeding activity as 

their terrestrial and fresh-water counterparts, and 

are probably able to attack the entire spectrum 

of plant and animal detritus (Johnson and Sparrow, 1961). Endolithic fungi in living and dead 

shells at the sediment-water interface produce 

intertwined, anastomosing, and branched networks, and are able to use the energy contained 

in organic conchiolin and chitin matrix and to 

parasitize algal microborers (Kohlmeyer, 1969; 

Calvaliere and Alberte, 1970; Green, 1975; Zeff 

and Perkins, 1979). 

Although marine fungi are generally believed 

to be restricted to the aerobic surface layers of 

bottom sediments, the Thraustochytriacea and 

Chyritidiaceae commonly occur well below the 

sediment surface (Clokie, 1970; Bremer, 1976; 

Johnson, 1976). The similarity between irregular 

polygonal networks observed in this study and 

microborings attributed to fungi leaves little 

doubt that these polygonal networks are of fungal 

origin. 

The associated irregular masses and crenulate 

discoids may be the reproductive bodies of these 

boring fungi. Kohlmeyer (1969) described irregular black peritheca and conidia 100 to 125 /xm 

in diameter as fruiting bodies for endolithic marine Ascomycetes and Deuteromycetes. Alternatively, these masses may be separate endolithic 

fungal colonies, similar to irregular patches and 

to crenulate rosettes attributed to fungal colonies 

attacking spores, pollen, and other organic microfossils (Elsik, 1971). 

In addition, two other groups of organisms, the 

bacteria and Actinomycetes, could produce endolithic scars resembling these forms. Similar 

scars on modern and ancient spores and pollen 

have also been attributed to bacteria and Actinomycetes (Moore, 1963; Elsik, 1966, 1971; Hav-
104 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 
NUMBER 12 105 

inga, 1971). Both groups are abundant at all 

levels within bottom sediments (ZoBell and Feltham, 1942) and at all depths sampled in lakes 

and the ocean (ZoBell and Rittenberg, 1938; 

Weyland, 1969; Willoughby, 1976). Boring by 

bacteria and Actinomycetes into carbonate substrates has not yet been investigated. 

The heterotrophic or chemoautotrophic mode 

necessitated by endolithic life within buried sediments may explain the occurrence of these forms 

only in the molluscan shell fragments planted in 

the Thalassia zone. The lack of endoliths within 

inorganic calcite rhombs planted below the sediment-water interface may indicate parasitic or 

saprophytic requirements of these organisms. Boring might be an effort to obtain nutrients from 

the organic matrix of the molluscan substrates. 

No reasons are known for the lack of these endoliths in the deeper fore-reef sand-trough zone of 

Carrie Bow Cay. Although both this environment 

and the Thalassia zone consist of a carbonate sand 

bottom, the latter may contain more interstitial 

detrital organic matter or may be a more reducing 

environment. The availability of nutrients may 

control the distribution of these endoliths and 

therefore may explain the paucity of these forms 

within coarse sediments lacking necessary nutrient sources. Eh conditions within sediments may 

also be a controlling factor. Endolithic boring in 

this case may not be an "active" search for organic matrices in carbonate substrates, but a 

"passive" result of metabolic reaction or a form 

of protection from interstitial grazers. 

FIGURE 70.—Scanning electron and transmitted light photomicrographs of unidentified microborings collected in subsurface station in the Thalassia zone: a, irregular polygonal 

network of varying intermittent borings, plastic cast within 

an etched mollusk fragment, 20 cm below the sedimentwater interface; b, irregular polygonal endolithic network 

within molluscan substrate, collected 35 cm below the sediment-water interface; c, irregular, flattened aggregates associated with a polygonal network, these patches might be 

related reproductive bodies or separate colonial forms, plastic 

cast of acid-etched molluscan substrate; d, crenulate discoid 

associated with a polygonal network, this also might be a 

separate organism or related reproductive body, plastic cast 

of an etched mollusk fragment, 5 cm below the sedimentwater interface. (Scale = 50 /xm for a, c, d; 25 /xm for b.) 

PALEOECOLOGIC AND GEOLOGIC SIGNIFICANCE.— 

Endoliths and their borings found in carbonate 

sediments that are exposed on the sea floor may 

provide valuable information for the study of 

paleoecological conditions of carbonate sediments. The microborers and their distributional 

patterns at the sediment-water interface of Belize 

closely resemble those of assemblages previously 

examined in St. Croix, Jamaica, and Florida 

(Perkins and Tsentas, 1976). It thus becomes 

possible to identify endoliths typical of a tropical 

shallow marine setting and to establish their occurrence in upper photic and lower photic zones. 

Microborings commonly are preserved within ancient carbonates (Hessland, 1949; Gatrall and 

Golubic, 1970; Golubic et al, 1975), but they have 

not been examined in relation to recent zonations 

for the interpretation of paleoenvironmental conditions. 

Conclusions 

Carbonate substrates, both conch shell fragments and cleaved calcite, planted just at the 

sediment-water interface in various depths along 

the reef transect off Carrie Bow Cay, Belize, 

contained a diverse assemblage of microboring 

forms. The blue-green alga Hyella tenuior and the 

green alga Ostreobium brabantium were restricted to 

the shallowest sample sites of the upper photic 

zone, which is also characterized by the abundant 

blue-green alga Mastigocoleus testarum and the 

green alga Phaeophila engleri and very little of the 

Conchocelis-stage of the red alga Porphyra sp. The 

lower photic zone, below approximately 12 m, is 

characterized by abundant Porphyra sp. and considerably less M. testarum and P. engleri. The bluegreen alga Plectonema terebrans was abundant at all 

depths examined. Hyphae of fungal endoliths 

were present at all sample sites, although different 

sporangial forms were bathymetrically restricted. 

This study of algal endoliths supports previous 

findings of a distinct tropical assemblage that 

may provide a basis for paleoecological studies of 

ancient assemblages. 

Below the sediment-water interface off Belize, 

endoliths infested only the molluscan (conch) 
106 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

fragments in the Thalassia-zone station and were 

not present in material buried in a deeper forereef sand-trough station. Restriction of infestation 

to the molluscan shell fragments suggests that 

these subsurface endoliths require organic matrices as nutrient sources. Three types of endoliths 

occur below the sediment-water interface: (1) 

irregular filamentous networks, (2) irregular flattened amorphous masses, and (3) regular crenulate discoids. These microborers have unknown 

taxonomic affinities but they closely resemble 

endolithic traces and scars attributed to fungi, 

bacteria, and Actinomycetes. Further studies are 

needed to explain the origin and geological importance of these possible environmental indicators and post-depositional carbonate degraders. 

Literature Cited 

Alexandersson, T. 

1972. Micritization of Carbonate Particles: Processes of 

Precipitation and Dissolution in Modern ShallowMarine Sediments. Uppsala University Geological Institutions, Bulletin, new series, 3:201-236. 

Bathurst, R.G.C. 

1966. Boring Algae, Micrite Envelopes, and Lithification of Molluscan Biosparites. Geological Journal, 5: 

15-32. 

Bremer, G. B. 

1976. The Ecology of Marine Lower Fungi. In E.B.G. 

Jones, editor, Recent Advances in Aquatic Mycology, 

pages 313-333. New York: John Wiley and Sons. 

Cavaliere, A. F., and R. S. Alberte 

1970. Fungi in Animal Shell Fragments. Journal of the 

Elisha Mitchell Scientific Society, 86:203-206. 

Clokie, J. 

1970. Some Substrate Relationships of the Family 

Thraustochytriaceae. Verbffentlichungen des Instituts 

fur Meeresforschung in Bremerhaven, 12:329-351. 

Edwards, B. B., and R. D. Perkins 

1974. Distribution of Microborings within Continental 

Margin Sediments of the Southeastern United 

States. Journal of Sedimentary Petrology, 44:1122-

1135. 

Elsik, W. C. 

1966. Biological Degradation of Fossil Pollen Grains and 

Spores. Micropaleontology, 12:515-518. 

1971. Microbiological Degradation of Sporopollenin. In 

J. Brooks, P. R. Grant, M. Muir, P. van Gijzel, 

and G. Shaw, editors, Sporopollenin, pages 480-511. 

New York: Academic Press. 

Gatrall, M., and S. Golubic 

1970. Comparative Study on Some Jurassic and Recent 

Endolithic Fungi Using Scanning Electron Microscope. In T. P. Crimes and J. C. Harper, editors, 

Trace Fossils. Journal of Geology, Special Issue, 3:167-

178. 

Golubic, S 

1969. Distribution, Taxonomy, and Boring Patterns of 

Marine Endolithic Algae. American Zoologist, 9: 

747-751. 

1972. Scanning Electron Microscopy of Recent Boring 

Cyanophyta and Its Possible Paleontological Application. In R. V. Desikachary, editor, Proceedings 

of the Symposium on Taxonomy and Biology of Blue-Green 

Algae, pages 167-170. India: Bangalore Press. 

Golubic, S., G. Brent, and T. LeCampion 

1970. Scanning Electron Microscopy of Endolithic Algae and Fungi Using a Multipurpose CastingEmbedding Technique. Lethaia, 3:203-209. 

Golubic, S., R. D. Perkins, and K. J. Lukas 

1975. Boring Microorganisms and Microborings in Carbonate Substrates. In R. W. Frey, editor, The Study 

of Trace Fossils, pages 229-259. New York: Springer. 

Green, M. A. 

1975. Survey of Endolithic Organisms from the Northeast Bering Sea, Jamaica, and Florida Bay. 112 

pages. Master's thesis, Department of Geology, 

Duke University, Durham, North Carolina. 

Havinga, A. J. 

1971. An Experimental Investigation into the Decay of 

Pollen and Spores in Various Soil Types. In J. 

Brooks, P. R. Grant, M. Muir, P. van Gijzel, and 

G. Shaw, editors, Sporopollenin, pages 446-478. 

New York: Academic Press. 

Hessland, I. 

1949. Investigation of the Lower Ordovician of the Siljan District, Sweden, II: Lower Ordovician Penetrative and Enveloping Algae from the Siljan 

District. Uppsala University Geological Institutions, 

Bulletin, 33:409-428. 

Hodgkin, E. P. 

1970. Geomorphology and Biological Erosion of Limestone Coasts in Malaysia. Geological Society of Malaysia, Bulletin, 3:27-51. 
NUMBER 12 107 

Johnson, T. W. 

1976. The Phycomycetes: Morphology and Taxonomy. 

In E.B.G. Jones, editor, Recent Advances in Aquatic 

Mycology, pages 193-211. New York: John Wiley 

and Sons. 

Johnson, T. W., and F. K. Sparrow 

1961. Fungi in Oceans and Estuaries. 668 pages. Weinheim: 

J. Cramer Publishing. 

Kobluk, D. R., and M. G. Risk 

1977. Algal Borings and Frambiodal Pyrite in Upper 

Ordovician Brachiopods. Lelhaia, 10:135-143. 

Kohlmeyer, J. 

1969. The Role of Marine Fungi in the Penetration of 

Calcareous Substances. American Zoologist, 9:741-

746. 

LeCampion-Alsumard, T. 

1975. Etude experimentale de la colonisation d'eclats de 

calcite par les Cyanophycees endolithes marines. 

Cahiers de Biologie Marine, 16:177-185. 

Moore, L. R. 

1963. Microbiological Colonization and Attack on Some 

Carboniferious Miospores. Palaeontology, 6:349-

372. 

Perkins, R. D. 

1972. Microboring Organisms and Environmental Indicators and Sediment Tracers: S. W. Puerto Rico 

Shelf [Abstract]. Geological Society of America, Abstracts with Programs, 1972 Annual Meeting, page 625. 

Perkins, R. D., and S. D. Halsey 

1971. Geologic Significance of Microboring Fungi and 

Algae in Carolina Shelf Sediment. Journal of Sedimentary Petrology, 41:843-853. 

Perkins, R. D., and C. I. Tsentas 

1976. Microbial Infestation of Carbonate Substrates 

Planted on the St. Croix Shelf, West Indies. Geological Society of America, Bulletin, 87:1615-1628. 

Purdy, E. G., and L. S. Kornicker 

1958. Algal Disintegration of Bahamian Limestone 

Coasts. Journal of Geology, 66:97-99. 

Reine, W.F.P. van, and C. van den Hoek 

1966. Isolation of Living Algae Growing in the Shells of 

Molluscs and Barnacles with EDTA (Ethylenediaminetetraacetic Acid). Blumea, 14:331-332. 

Rooney, W. S., Jr., and R. D. Perkins 

1972. Distribution and Geologic Significance of Microboring Organisms within Sediments of the Arlington Reef Complex, Australia. Geological Society of 

America, Bulletin, 83:1139-1150. 

Scherer, M. 

1974. The Influence of Two Endolithic Microorganisms 

on the Diagenesis of Recent Coral Skeletons. Neues 

Jahrbuch fiir Geologie und Palaeontologie, Monatshefte, 

9:557-566. 

Schneider, J. 

1976. Biological and Inorganic Factors in the Destruction of Limestone Coasts. Contributions to Sedimentology, 6:1-112. 

Schroeder, J. H. 

1972a. Calcified Filaments of an Endolithic Alga in Recent Bermuda Reefs. Neues Jahrbuch fiir Geologie und 

Palaeontologie, Monatshefte, 7:16-33. 

1972b. Fabrics and Sequences of Submarine Carbonate 

Cements in Holocene Bermuda Cup Reefs. Geoligische Rundschau, 61:708-730. 

Schroeder, J. H., and R. N. Ginsburg 

1971. Calcified Algal Filaments in Reefs: Criterion of 

Early Diagenesis [Abstract]. American Association of 

Petroleum Geologists, Bulletin, 55:364. 

Swinchatt, J. P. 

1965. Significance of Constituent Composition, Texture 

and Skeletal Breakdown in Some Recent Carbonate Sediments. Journal of Sedimentary Petrology, 35: 

71-90. 

Taylor, B. J. 

1971. Thallophyte Borings in Phosphatic Fossils from 

the Lower Cretaceous of Southeast Alexander Island, Antarctica. Palaeontology, 14:294-302. 

van Reine. See Reine, W.F.P. van 

Weyland, H. 

1969. Actinomycetes in North Sea and Atlantic Ocean 

Sediments. Nature, 223:858. 

Willoughby, L. G. 

1976. Freshwater Actinomycetes. In E.B.G. Jones, editor, 

Recent Advances in Aquatic Mycology, pages 673-698. 

New York: John Wiley and Sons. 

Zeff, M. L., and R. D. Perkins 

1979. Microbial Alternations of Bahamian Deep-Sea 

Carbonates. Sedimentology, 26:175-201. 

ZoBell, C. E., and C. B. Feltham 

1942. The Bacterial Flora of a Marine Mud Flat as an 

Ecological Factor. Ecology, 23:69-78. 

ZoBell, C. E., and S. C. Rittenberg 

1938. The Occurrence and Characteristics of Chitinoclastic Bacteria in the Sea. Journal of Bacteriology, 

35:275-287. 
Production of Some Benthic Communities 

at Carrie Bow Cay, Belize 

Paul E. Hargraves 

ABSTRACT 

The primary production of Thalassia testudinum, 

mixed algae on coral rubble, and coral sand in 

the vicinity of Carrie Bow Cay, Belize, was estimated from in situ oxygen flux (by titrimetry) in 

enclosed chambers. Thalassia beds were autotrophic, yielding a daily net production in the 

range of 2.22-42.3 mg O2 per square meter (approximately equivalent to 0.67-12.7 g C per 

square meter). The other habitats were heterotrophic despite considerable microalgal development, with a production/respiration ratio (P/R) 

less than unity. Contribution by the phytoplankton was negligible. Diurnal oxygen flux in open 

waters over Thalassia beds was typical for similar 

tropical areas. 

Introduction 

In tropical coastal waters, coral reef ecosystems 

are generally considered to be highly productive 

in contrast to plankton communities, whose primary production is quite low or even negligible 

(Lewis, 1977; Milliman and Mahnken, 1969; 

Sournia, 1969). The metabolism of coral reefs as 

a unit has been the subject of numerous studies 

beginning with Sargent and Austin (1949) and 

Odum and Odum (1955). The remarkable fertility of reefs, their dynamic balance, and susceptibility to environmental perturbation have often 

been noted (see reviews by Lewis, 1977, and 

Johannes in Wood and Johannes, 1975). 

The plants responsible for primary production 

in tropical coastal waters include mangroves, sea 

Paul E. Hargraves, Graduate School of Oceanography, University of 

Rhode Island, Kingston, R. I. 02881. 

grasses, macroscopic algae, boring and epipelic 

algae, zooxanthellae, and phytoplankton. The 

relative importance of each group varies from 

area to area, but except for special environments 

such as enriched lagoons and upwelling areas 

(Sournia, 1969; Steeman-Nielsen, 1975), the contribution of phytoplankton is low. The techniques 

of investigating primary productivity have varied 

among investigators according to whether information is required on the productivity of the total 

community or its components, and according to 

the facilities and methods available. In general, 

chemical or polarographic measurements of oxygen flux are sufficiently precise for nonplanktonic 

biotopes, and these techniques have provided 

considerable information on the metabolism of 

tropical benthic communities (for instance, Lewis, 

1977; McRoy and McMillan, 1977). 

Reported here are results of production measurements in several habitats at and near Carrie 

Bow Cay, Belize. Topographic, oceanographic, 

and ecological characteristics of the area are presented elsewhere in this volume. 

ACKNOWLEDGMENTS.—The technical assistance 

of D. Hargraves and K. Zimmerly is gratefully 

acknowledged. Partial support was provided by 

National Science Foundation Grant OCE-76-

82280. N. Marshall reviewed an early draft of the 

manuscript. 

Methods 

Oxygen flux was measured in three habitats: 

seagrass beds (Thalassia testudinum Banks ex Konig 

with small amounts of Syringodium filiforme Kiitzing) at depths of 1 to 3 m on the lagoon side of 

109 
110 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Carrie Bow Cay; mixed algal communities on 

coral rubble at depths of 0.5 to 1.0 m in the 

rubble and pavement zone behind the reef crest; 

and coral sand at a depth of 1 m in the back-reef 

zone. The coral rubble was composed of cobblesized pieces of coral mixed with sand and with 

small (unidentified) filamentous, crustose, and 

thallose seaweeds. The coral sand first seemed 

devoid of organisms, but microscope examination 

revealed considerable numbers of blue-green algae, diatoms, Foraminifera with symbiotic algae, 

filamentous algae, and a multitude of other representatives of the micro- and meiobenthos. 

Experiments were conducted over periods of 

dawn/noon, noon/sunset, and 24 h cycles with 

determinations at 1 or 2 h intervals, all during 

April/May 1977 and January 1978. For comparative purposes, the production of phytoplankton 

was also assessed. 

For each experiment three transparent and two 

opaque polystyrene boxes of 500 cm area were 

inverted over the substrate. The edges were 

pushed into the sediment to prevent leakage 

around the margin of the container. Water was 

withdrawn through sampling ports with a 50 ml 

capacity syringe. The total volume of enclosed 

water was 4.2 liters. For diurnal fluctuations in 

oxygen and phytoplankton production measurements, 300 ml biological oxygen demand (BOD) 

bottles were used. 

Dissolved oxygen was measured titrimetrically 

by the Winkler method, using phenylarsine oxide 

(PAO) as titrant. PAO is superior to thiosulfate 

in its longer shelf life and resistance to bacterial 

decomposition, and produces results with comparable precision. 

During the experiments water temperature 

varied from 26° to 29°C (by mercury immersion 

thermometer) and salinity varied from 34.4 to 

35.2 %o (by Endeco refractometer type 102). As 

much as possible, all experiments were conducted 

on days that were cloud free, or nearly so. 

Results and Discussion 

Results of a preliminary experiment to determine the diurnal variation in dissolved oxygen in 

the open water over a Thalassia bed are shown in 

Figure 71. During predawn hours, characteristically, minimum oxygen concentrations are exhibited. The subsequent rapid increase of dissolved oxygen reflects photosynthetic activity (as 

light intensity increases). The daily oxygen maximum occurs at or somewhat after noon, at which 

time minute streams of oxygen bubbles frequently 

issue from some Thalassia blades. A gradual decline in oxygen concentration occurs during the 

rest of the day, becoming more pronounced in 

the late afternoon. Minimum concentration is 

again reached in the predawn hours. This pattern 

is typical and well documented, not only for 

tropical seagrass communities (for instance, 

Odum et al., 1959; Qasim and Bhattathiri, 1971) 

but for coral reefs" as well (for instance, Sournia, 

1976a). For part of the day, the water is supersaturated, often in excess of 150%, with respect to 

oxygen. Some consequences of these conditions 

are discussed below. 

Table 9 summarizes the production of the different communities. Oxygen data were converted 

to carbon assuming a photosynthetic quotient 

(PQ) of 1.25 (McRoy and McMillan, 1977; Westlake, 1963). The standing stock of Thalassia blades 

varied from about 34 to 126 g/m2 dry weight, 

with a mean of about 47 g. Other organisms were 

not analyzed quantitatively. 

Compared to published data for seagrass beds, 

summarized in Lewis (1977) and McRoy and 

Time (h) 

FIGURE 71.—Changes over a 24-hour period in dissolved 

oxygen over a Thalassia bed near Carrie Bow Cay, 28-29 

April 1977. 
NUMBER 12 111 

TABLE 9.—Oxygen production of communities at Carrie Bow Cay, 26 April-5 May 1977, unless 

otherwise indicated (gross and net carbon production assumes P Q = 1.25 (Westlake, 1963); 

phytoplankton assumes 1 m water depth) 

Community 

Thalassia beds 

Thalassia beds 

(Jan 1978) 

Mixed algae/ 

coral rubble 

Coral sand 

Phytoplankton 

Trials 

(n) 

5 

3 

3 

2 

2 

Production 

(mg 02/m2/d) 

Gross Net 

19.3-118.1 

14.9-33.2 

60.0-94.3 

75.5, 107.9 

0.48, 1.00 

6.03-42.30 

2.22-14.50 

163.0-31.6 

-11.3,-32.1 

0.14,-0.07 

Respiration 

(mg 02/m2/d) 

13.27-75.80 

12.68-18.70 

223.0-125.9 

86.8, 140.0 

0.34, 1.07 

P/R 

1.45-1.56 

1.18-1.78 

0.27-0.75 

0.87,0.77 

1.41,0.93 

Production 

Gross 

5.79-35.43 

4.47-9.96 

18.00-28.29 

22.65, 32.37 

0.14,0.30 

(g C/m2/d) 

Net 

1.81-12.69 

0.67-4.35 

-48.90-9.48 

-3.39, -9.63 

0.04, -0.02 

McMillan (1977), the data for Thalassia at Carrie 

Bow Cay fall within previously noted ranges for 

gross and net production. The standing stock is 

rather low compared to several other localities in 

the Caribbean and Gulf of Mexico (refer to table 

1 in McRoy and McMillan, 1977). In contrast to 

some Thalassia habitats in Puerto Rico (Odum et 

al., 1959), in the present case all measurements 

indicate a consistently autotrophic community, 

that is, ratio of production to respiration (P/R) 

in excess of 1. The wide range in measurements, 

in part attributable to density variations in Thalassia and seasonal differences, make extrapolations to an annual production uncertain. For 

comparative purposes the mean production for 

all Thalassia measurements may be estimated at 

1800 g C/m2/y net production. This amount is 

higher than those reported from seagrass beds in 

Puerto Rico (Odum et al., 1959), but considerably less than those reported from Cuba and 

Florida (Buesa, 1972; Odum, 1957, 1963). 

Differences in production were noted between 

late spring and winter. The extent to which these 

differences are significant in unknown because of 

the small number of replicates analyzed. Such 

variables as photoperiod (Marmelstein et al., 

1968) and water temperature (citations in McRoy 

and McMillan, 1977) affect Thalassia production 

and add to the uncertainty of annual rate comparisons based on extrapolation. 

A considerable area of the back-reef zone at 

Carrie Bow Cay consists of coral rubble and 

pavement rock overgrown with a variety of small 

but conspicuous filamentous and thallose algae. 

Despite a gross production generally exceeding 

that of the Thalassia beds (Table 9), the community as a whole is heterotrophic, having a P/R 

less than 1. This habitat harbors a diverse and 

abundant fauna of burrowing invertebrates that 

contribute to the high community respiration. 

Grazing effects of fishes and large invertebrates, 

not considered here, are probably also significant 

in the consumption of primary producers (Marsh, 

1976). The relative contribution of microalgal 

species in this habitat has generally been neglected, although Qasim et al. (1972) reported 

net production rates of 365-800 g C/m2/y for 

similar types of algae from the Laccadive Archipelago. Overall, the algal oxygen production in 

this habitat is less than oxygen consumption by 

herbivores and other animals. 

Although the extensive coral sand areas north 

and northwest of Carrie Bow Cay appear to have 

few plant producers, microscopic examination 

revealed large numbers of benthic pennate diatoms and filamentous blue-green algae (or cyanobacteria), smaller numbers of Foraminifera with 

apparent endosymbiotic algae, and large numbers of micro- and meiobenthos metazoans. Community respiration slightly exceeded gross production, with P/R less than 1 (Table 9), although 

this habitat was closer to an autotrophic condition than the rubble-pavement habitat. The contribution of bacteria to community respiration 
112 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

was not evaluated but it may be significant (see 

for instance, Edwards, 1978). The gross production approached that of the Thalassia beds, and 

therefore indicated the high photosynthetic activity here. 

That such sand areas are not necessarily heterotrophic has been convincingly demonstrated 

by Sournia (1976b). Extensive development of 

the blue-green alga Oscillatoria limosa in sand at 

Moorea Island lagoon was responsible for an 

average gross production equivalent of over 6000 

mg 0 2 / m2/ d with P/R = 1.5-3.0, a highly autotrophic habitat. Although blue-green algae are 

not abundant at Carrie Bow Cay, the potential 

contribution of coral sand areas should not be 

ignored in calculations of production in reef areas. 

Under some circumstances Foraminifera with endosymbiotic algae may dominate coral sands. 

Sournia (1976a) described sands from Takapota 

Atoll with a net production of 115-354 mg 

0 2 / m2/ h (= 43-133 mg C/m2/h) and with populations consisting primarily of Foraminifera with 

endosymbionts. Small numbers of these protozoa 

were also present in sands at Carrie Bow Cay, but 

pennate diatoms dominated as primary producers. 

These sands could potentially assume the autotrophic role that Sournia (1976a) described. A 

minor perturbation in the environment of organisms already under natural stress (by light and 

temperature thermal, among others) may shift 

community composition to one of low density 

consisting of eurytolerant species (Wood and Johannes, 1975). Several different species of bluegreen algae, which certainly qualify as eurytolerant, can be observed in the sands of Carrie Bow 

Cay. Coral sand habitats have been largely neglected by ecologists, so that it would be useful to 

stress them experimentally in various ways in 

order to determine whether and how their trophic 

status is modified. Perturbation in the form of 

nutrient enrichment was attempted in one microatoll, resulting in significantly increased production (Kinsey and Domm, 1974). 

As expected, the production of the phytoplankton was very low, and near the limits for accuracy 

of the technique used (Table 9), supporting the 

conclusion that phytoplankton in the vicinity of 

coral reefs contribute a negligible amount to net 

production (Milliman and Mahnken, 1969; Sournia and Ricard, 1976). Although such shallow 

areas generally have, in addition to the normal 

complement of typically planktonic forms, a 

higher number of benthic diatoms swept into the 

water column by turbulence, net production remains negligible. For offshore Caribbean waters 

net production can be higher or lower than in 

reef areas, depending on locale. Beers et al. (1968) 

found 0.03-0.28 g C/m2/d offshore from Jamaica; Steeman-Nielsen and Jensen (1957) found 

0.14 and 0.19 g C/m2/d in the south-central 

Caribbean Sea. In contrast, Ricard (1977) found 

higher production in the lagoon than in the open 

ocean at Tahiti, but the converse was true at 

Lakeba lagoon. Higher productivity is unquestionably possible under localized enriched conditions (Gordon et al., 1971; Margalef, 1975). 

The limitations of technique and procedures 

used in this work deserve brief mention. Production in tropical benthic communities has been 

studied in several ways: by flow respirometry, 

which measures changes in the flux of carbon 

dioxide or oxygen (upstream-downstream); by in 

situ light/dark bottle methods using oxygen flux 

(titrimetric and polarographic) and various radioactive tracers; and by direct measurement of 

changes in standing stocks. All these methods are 

subject to errors and various assumptions (for 

discussion, see Lewis, 1977; Sournia, 1976c; Vollenweider, 1974) that will not be examined here, 

except to note that the technique used here appears to be satisfactory for benthic habitats. Unquestionably, accuracy decreases in the case of 

water supersaturated with oxygen; similarly, respiration of animals is affected by reduced oxygen 

levels. Recycling of oxygen in lacunae of Thalassia 

blades is another potential source of error (Hartmann and Brown, 1967). 

The accuracy of conversion of oxygen flux to 

its equivalent carbon varies widely depending on 

the photosynthetic quotient assumed. This creates a problem in comparing the production rates 
NUMBER 12 113 

calculated by different authors. Photosynthetic 

quotients ranging from 0.86 to 3.00 are possible. 

This work follows Westlake (1963) in the assumption that P Q = 1.25 is valid for natural tropical 

communities in favorable conditions. 

Two important sources of tropical primary production have not been considered here, namely 

the coral zooxanthellae and the larger seaweeds. 

The complex role of the former in reef trophic 

structure has been discussed by Lewis (1977) and 

Marsh (1976) among others, and the contribution 

of zooanthellae at Carrie Bow Cay is certainly 

worthy of investigation. Under some conditions 

the larger seaweeds contribute significantly to 

primary production, even more so than Thalassia 

beds (Doty, 1971; Wanders, 1976), but their development around Carrie Bow Cay is not extensive. 

Conclusions 

Of the benthic habitats investigated at Carrie 

Bow Cay, Thalassia beds were most productive, 

yielding a maximum gross production of 35.4 g 

C/m / d and a maximum net production of 12.7 

g C/m2/d. Coral sand habitats and mixed algae 

with coral rubble habitats were heterotrophic in 

nature, despite considerable algal development. 

Net production by phytoplankton was negligible. 

The diurnal oxygen content of water flowing over 

Thalassia beds fluctuates in a manner typical of 

similar habitats previously described. 

Given the problems inherent in comparing a 

wide variety of techniques and procedures, the 

pattern of primary production at Carrie Bow Cay 

is typical of similar tropical areas. 

Addendum 

Since the writing of this contribution three 

pertinent papers have appeared. Two center on 

reef community metabolism elsewhere in the Caribbean (Puerto Rico, U.S. Virgin Islands) and 

also employ the Winkler method for oxygen determinations (Rogers, 1979; Rogers and Salesky, 

1981). The third deals with primary production 

by microalgae in North Sea sediments comparing 

various techniques, including platinum electrode 

measurements that allow recording of dissolved 

oxygen microprofiles (Revsbech et al., 1981). 

Literature Cited 

Beers, J. 

1968. 

Buesa, R 

1972. 

Doty, M. 

1971. 

Edwards 

1978. 

R., D. M. Steven, and J. B. Lewis 

Primary Productivity in the Caribbean Sea off 

Jamaica and the Tropical North Atlantic off Barbados. Bulletin of Marine Science, 18:87-104. 

•JProduccion primaria de las praderas de Thalassia 

testudinum de la platforma noroccidental de Cuba. 

Cuba Centro de Investigaciones Pesquera, Revista Traba7'oj,3:101-143. 

S. 

The Productivity of Benthic Frondose Algae at 

Waikiki Beach, 1967-1968. University of Hawaii, 

Botanical Science Paper, 22: 119 pp. 

R.R.C. 

Ecology of a Coastal Lagoon Complex in Mexico. 

Estuarine and Coastal Marine Science, 6:75-92. 

Gordon, D. C , R. O. Fournier, and G. J. Krasnic 

1971. Note on the Planktonic Primary Production in 

Fanning Island Lagoon. Pacific Science, 25:228-

233. 

Hartman, R. T., and D. L. Brown 

1967. Changes in Composition of the Internal Atmosphere of Submerged Vascular Hydrophytes in 

Relation to Photosynthesis. Ecology, 48:252-258. 

Kinsey, D. W., and A. Domm 

1974. Effects of Fertilization on a Coral Reef Environment—Primary Production Studies. In A. M. 

Cameron, B. M. Campbell, A. B. Cribb, R. Endean, J. S. Jell, O. A. Jones, P. Mather and F. H. 

Talbot, editors, Proceedings of the Second International 

Coral Reef Symposium, 1:49-66. Brisbane, Australia: 

The Great Barrier Reef Committee. 
114 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Lewis, J. B. 

1977. Processes of Organic Production on Coral Reefs. 

Biological Review, 52:305-347. 

Margalef, R. 

1975. Fitoplancton invernal de la laguna costera de 

Alvarado (Mexico). Anales del Instituto Botanico Antonio Jose Cavanilles, 32:381-387. 

Marmelstein, A. D., P. W. Morgan, and W. E. Pequegnat. 

1968. Photoperidism and Related Ecology in Thalasia 

testudinum. Botanical Gazette, 129:63-67. 

Marsh, J. A. 

1976. Energetic Role of Algae in Reef Ecosystem. Micronesica, 12:13-22. 

McRoy, C. P., and C. McMillan 

1977. Production Ecology and Physiology of Seagrasses. 

In C. P. McRoy and C. Helfferich, editors, Seagrass 

Ecosystem, Scientific Perspective, pages 53-87. New 

York: Marcel Dekker, Inc. 

Milliman, J. D., and C. V. Mahnken 

1969. Appendix: Reef Productivity Measurements. Atoll 

Research Bulletin, 129:23-26. 

Odum, H. T. 

1957. Primary Production of Eleven Florida Springs and 

a Marine Turtle Grass Community. Limnology and 

Oceanography, 2:85-97. 

1963. Productivity Measurements in Texas Turtle Grass 

and the Effects of Dredging an Intracoastal Channel. Publications of the Institute of Marine Science, 

University of Texas, 9:45-58. 

Odum, H. T., P. R. Burkholder, and J. Rivero 

1959. Measurements of Productivity of Turtle Grass 

Flats, Reefs, and the Bahia Fosforescenti of Southern Puerto Rico. Publications of the Institute of Marine 

Science, University of Texas, 6:159-170. 

Odum, H. T. and E. P. Odum 

1955. Trophic Structure and Productivity of a Windward Coral Reef on Eniwetok Atoll. Ecological 

Monographs, 25:291-300. 

Qasim S. Z., and P.M.A. Bhattathiri 

1971. Primary Production of a Seagrass Bed on Kavaratti Atoll (Laccadives). Hydrobiologia, 38:29-38. 

Qasim, S. Z., P.M.A. Bhattathiri, and C.V.G Reddy 

1972. Primary Production of an Atoll in the Laccadives. 

Internationale Revue des gesamten Hydrobiologie, 57: 

207-225. 

Revsbech, N. P., B. B. J0rgensen, and O. Brix 

1981. Primary Production of Microalgae in Sediments 

Measured by Oxygen Microprofile, H14C03-Fixation, and Oxygen Exchange Methods. Limnology 

and Oceanography, 26:717-730. 

Ricard, M. 

1977. Phytoplankton Contribution of Primary Productivity in Two Coral Reef Areas of Fiji Islands and 

French Polynesia. In D. L. Taylor, editor, Proceedings of the Third International Symposium on Coral Reefs, 

1:343-348. Miami, Florida; Rosenstiel School of 

Marine and Atmospheric Science. 

Rogers, C. S. 

1979. The Productivity of San Cristobal Reef, Puerto 

Rico. Limnology and Oceanography, 24:342-349. 

Rogers, C. S., and N. H. Salesky 

1981. Productivity oi Acropora palmata (Lamarck), Macroscopic Algae, and Algal Turf from Tague Bay 

Reef, St. Croix, U.S. Virgin Islands. Journal of 

Experimental Marine Biology and Ecology, 49:179-187. 

Sargent, M. C , and T. S. Austin 

1949. Organic Productivity of an Atoll. Transactions of the 

American Geophysical Union, 30:245-249. 

Sournia, A. 

1969. Cycle annuel du phytoplancton et de la production primaire dans les mers tropicales. Marine Biology, 3:387-303. 

1976a. Primary Production of Sands in the Lagoon of an 

Atoll and the Role of Foraminiferan Symbionts. 

Marine Biology, 37:29-32. 

1976b. Ecologie et productivity d'une Cyanophycee en 

milieu corallien: Oscillatoria limosa Agardh. Phycologia, 15:363-366. 

1967c. Oxygen Metabolism of a Fringing Reef in French 

Polynesia. Helgolander Wissenschaftliche Meeresuntersuchungen, 28:401-410. 

Sournia, A., and M. Ricard 

1976. Phytoplankton and its Contribution to Primary 

Productivity in Two Coral Reef Areas of French 

Polynesia. Journal of Experimental Marine Biology and 

Ecology, 21:129-140. 

Steeman-Nielsen, E. 

1975. Marine Photosynthesis. 141 pages. New York: Elsevier Scientific Publishing Company. 

Steeman-Nielsen, E., and A. Jensen 

1957. Primary Oceanic Production: The Autotrophic 

Production of Organic Matter Production in the 

Ocean. Galathea Report, 1:49-136. 

Vollenweider, R. A., editor 

1974. A Manual for Measuring Primary Production in Aquatic 

Environments. 213 pages. Oxford and Edinburgh: 

Blackwell Scientific Publications. 

Wanders, J.B.W. 

1976. The Role of Benthic Algae in the Shallow Reef of 

Curacao (Netherlands Antilles), II: Primary Productivity of the Sargassum Beds on the Northeast 

Coast Submarine Plateau. Aquatic Botany, 2:327-

335. 

Westlake, D. F. 

1963. Comparisons of Plant productivity. Biological Reviews, 38:385-425. 

Wood, E.J.F., and R. E. Johannes, editors 

1975. Tropical Marine Pollution. 192 pages. Amsterdam, 

Oxford, New York: Elsevier Scientific Publishing 

Company. [Elsevier Oceanography Series, 12.] 
Macrobenthic Invertebrates in Bare Sand 

and Seagrass {Thalassia testudinum) 

at Carrie Bow Cay, Belize 

David K. Young 

and Martha W. Young 

ABSTRACT 

The generally accepted view that seagrasses 

support a more dense and diverse invertebrate 

fauna than sand areas devoid of such vegetation 

was tested in Thalassia testudinum beds of Carrie 

Bow Cay lagoon, Belize. Mechanisms regulating 

the distribution of invertebrates were also examined. 

Core samples from five stations, ranging from 

bare sand to dense seagrass, indicated no clearcut effect of seagrass versus bare sand on densities 

of polychaetes and mollusks. The numbers of 

species from all seagrass stations were not significantly higher than those from sand stations. Species diversities for all stations were influenced 

mainly by species richness, except for the station 

in dense seagrass where species evenness was high. 

The most abundant polychaete and mollusk species sampled were found at all stations. 

The proportion of fine-grained sediments in 

the substrate increased progressively from bare 

sand into the seagrass. Encrusting coralline algae 

provide a major contribution to the fine sediments of the grass beds. Seagrass standing crop 

increased from the small seagrass patch to the 

dense Thalassia bed. Inorganic components made 

up 49% of live and 70% of dead seagrass. 

This study found no relationship between seagrass standing crop and species densities of invertebrates. Intense predation pressure is proposed 

as the primary regulating mechanism of species 

David K. Young, Naval Ocean Research and Development Activity, 

Department of the Navy, National Space Technology Laboratory 

Station, Miss. 39529. Martha W. Young, U.S. Fish and Wildlife 

Service, National Coastal Ecosystem Team, Slidell, La. 70458. 

densities in the Carrie Bow lagoon. Experimental 

field studies are needed to clarify predator-prey 

interactions. 

Introduction 

Seagrasses are widely thought to support a 

more dense and diverse invertebrate fauna than 

bare sediment devoid of such vegetation. Only 

recently has this generalization been subjected to 

rigorous quantitative sampling and experimental 

testing (Heck and Wetstone, 1977; Orth, 1977; 

Reise, 1977; Young and Young, 1978). The mechanisms of distribution examined by these workers 

primarily concern (1) food, (2) living space, (3) 

predation, and (4) sediment stability. 

FOOD.—Because seagrasses decompose rapidly 

relative to other vascular plants in the marine 

environment, they are considered to be major 

contributors to the detrital food web of seagrass 

ecosystems (Fenchel, 1977). Heck and Wetstone 

(1977) suggested from their work in Panama that 

food reserves are abundant in seagrass ecosystems 

and that food is not a limiting factor. Young and 

Young (1978), however, have experimentally 

demonstrated that numbers of certain depositfeeding invertebrates are food-limited in a seagrass detritus-rich habitat in Florida and that 

significant increases in overall macrobenthic densities result from addition of detrital food with 

high nitrogen content. 

LIVING SPACE.—Increased surface area afforded 

115 
116 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

by topographically complex living and nonliving 

substrata has been defined as an important ecological parameter on the study transect at Carrie 

Bow Cay (Dahl, 1973; Riitzler and Macintyre, 

herein: 9). Dahl (1973) has estimated that the 

functional surface area of bare sand is increased 

over four times by the presence of seagrass in the 

Carrie Bow Cay lagoon. Young and Young (1978) 

have experimentally shown no significant decreases in overall species densities of macrobenthos following the clipping and removal of seagrass blades from an area in a Florida seagrass 

bed, although the abundance of several seagrassassociated species diminished. 

PREDATION.—Heck and Wetstone (1977) suggested that plant blades provide protection to 

epifauna from visually searching fish predators. 

Orth (1977) and Reise (1977) provided experimental evidence that seagrass roots and rhizomes 

decrease the effectiveness of crab predators upon 

infauna. Reise (1977) has also shown that coarsegrained sediments afford protection to infauna 

from crab predation more than fine-grained sediments. Because finer sediments accumulate more 

in seagrass beds relative to adjacent areas bare of 

vegetation (Orth, 1977), a clear-cut assessment of 

the effect of sediment size versus that of seagrass 

upon intensity of predation cannot be discerned 

in most seagrass areas. 

SEDIMENT STABILITY.—Seagrass roots and rhizomes bind loose sediments together, and their 

blades baffle waves; both actions assist in preventing sediment resuspension and erosion (Ginsburg and Lowenstam, 1958). Orth (1977) has 

demonstrated that the overall sediment stabilizing effect of seagrasses is a major factor in increased densities and diversity of infauna of seagrass beds in Chesapeake Bay and Bermuda. 

Our primary intent in this study was to determine whether densities of macrobenthic invertebrates in the Carrie Bow Cay lagoon were related 

to the presence of vegetation and, within the time 

constraints, which mechanisms were regulating 

the distribution of species. 

ACKNOWLEDGMENTS.—We thank B. Spracklin 

for his invaluable assistance during our stay in 

Belize. Appreciation is due T. Wolcott for sediment analyses. This study was funded by a grant 

from the Atlantic Foundation to the Smithsonian 

Institution and by the Harbor Branch Foundation, Inc. Laboratory space was provided by Mississippi State University Research Center and 

administrative support by the Naval Ocean Research and Development Activity. 

Study Area and Methods 

The lagoon floor of Carrie Bow Cay is covered 

by calcareous sediment comprised largely of coral, 

encrusting coralline algae, Halimeda, molluscan, 

foraminiferal, and echinoid material. The sand 

surface in areas lacking vascular vegetation is 

here and there covered with a fine filamentous 

algal "felt" (Dahl, 1973). The seagrass Thalassia 

testudinum Banks ex Konig grows profusely in these 

sediments, and grades from small, thinly vegetated patches (from the back-reef region) to the 

extensive, dense beds of the lagoon. The Thalassia 

is interspersed with sparse stands of the seagrass 

Syringodium filiforme Kiitzing. Thalassia blades are 

heavily encrusted with coralline algae and conspicuously devoid of epizoans, with the exception 

of Foraminifera. 

We extended the established transect (Riitzler 

and Macintyre, herein) from its beginning point 

at the buoy (0 m) lagoonwards 5 m into a dense 

bed of Thalassia (Figure 72). Station 1 (+23 m) 

was an area of bare sand containing no vegetation, station 2 (+18 m) was a small patch of 

Thalassia, station 3 (+12 m) was in bare sand, 

station 4 (+5 m) was in Thalassia at the edge of 

an area densely covered with seagrass, and station 

5 (—5 m) was inside the Thalassia bed. Because of 

the depth of water at the sampling locations 

(approximately 2 m) and the small tidal variation 

(0.2 m tidal range, Kjerfve, 1978), seagrasses in 

this area of the lagoon are never exposed at low 

tide. Salinity was 34 %o throughout the sampling 

period and water temperature varied from 27° to 

28° C. The grass area sampled at station 2 is 

approximately 80 m from the nearest patch reefs. 

The sand areas (stations 1 and 3) are at the same 

depth as adjacent seagrass stations. These bare 
NUMBER 12 117 

Lagoon 

5 

(-5m) 

x— 

Buoy 

(Om) 

4 

(+5m) 

3 

(+12m) 

2 

(+18m) 

Reef 

1 

(+2 3 m) 

FIGURE 72.—Diagrammatic representation of the transect in Carrie Bow Cay lagoon showing 

sampling stations for this study; the buoy marks the beginning of the main transect (Riitzler 

and Macintyre, herein), which was extended, for purposes of this study, 5 m into dense Thalassia 

(distance from zero point in parentheses). 

sand stations show no traces of scour or ripple 

marks. 

Samples were taken from 23 to 31 March 1976 

with a PVC corer (9.45 cm in diameter) to a 

depth of 20 cm (= 1405 cm3) at five stations 

along a 28 m long transect. Each sample consisted 

of four replicate cores taken perpendicular to the 

transect at each sampling station. The number of 

replicate cores per sample was determined by 

analyses of pilot samples taken in dense Thalassia 

from the lagoon. Four replicate cores contained 

90% or more of the species of polychaetes and 

mollusks expected according to the test of Gaufin 

et al. (1956). Samples taken in Thalassia included 

blades, roots, and rhizomes. 

Samples were sieved through 1.0 mm mesh 

Nytex screen, narcotized with 0.15% propylene 

phenoxytol in sea water, stained with rose bengal 

and fixed in 5% to 10% formalin in sea water. 

Organisms were transferred to 70% ethyl alcohol 

after 24 hours for sorting, identification and storage. In this study, the term "macrobenthos" refers 

to those benthic invertebrates retained on a 1.0 

mm mesh screen. Sampling efficiency of 1.0 mm 

mesh was determined by screening one of the 

replicates from each station through both 0.5 mm 

and 1.0 mm mesh screen. Only whole animals, 

anterior fragments of polychaetes and live shells 

of mollusks were identified as to species and 

counted. All other macrobenthos were identified 

only to higher taxonomic categories and counted. 

One sediment sample (comprising four replicate cores) was taken with the PVC corer at each 

station. Particle sieve size was determined using 

a Ro-Tap and analyzed by the method of Folk 

(1974). Samples from each particle size category 

were examined microscopically. 

All seagrass blades were clipped at the sediment 

surface and shoots were counted from one-metersquare areas adjacent to stations 2, 4, and 5. Erect 

live seagrass blades and prostrate dead blades 

were collected separately from each clipped plot. 

Live and dead seagrass dried at 100° C for 48 

hours and weighed provided an estimate of seagrass standing crop at each station. Organic and 

inorganic composition was determined from the 

ash of both live and dead fractions processed at 

550° C for 2 hours. 

A random sample of Thalassia blades that 

showed evidence of grazing by herbivorous fishes 

was collected, and bite marks were counted and 

measured. 

Data were analyzed (log2) using Shannon's 

information measure of species diversity, H' (Pielou, 1966); species richness, d, was determined by 

the method of Margalef (1958), and species evenness by E' (Buzas and Gibson, 1969). A one-way 

analysis of variance (ANOVA) using non-transformed data compared numbers of species as well 

as species densities at the 5 stations. The a posteriori Student-Newmans-Keuls (SNK) multiple 

comparison test was run, since in both cases the 

ANOVA indicated significant differences among 

means (Sokal and Rohlf, 1969). Significance for 

the ANOVA and SNK tests was chosen at the 

95% level of confidence. 
118 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

. 3 

E 5 ^ 

CM CM CM CT) O") CO COC O r^t-~ to to ID in in in- * 

CO CO CO <T> <J> O O — —' —c CM CM CM CO CO CO Th LO in io 

CO 

CM 

CO 

^f 

o 

O 

<£> 

<JD 

O 

O 

CO 

CN 

i n 

CM 

o 

o 

o 

uO 

CM 

CM 

o 

Th 

o 

o 

i n 

CM 

o 

o 

o 

•tfl 

Th 

CM 

-

<* 

o 

CO 

o 

CM 

-

o 

r~ 

o 

o 

CM 

o 

o 

o 

o 

1 ^ 

CM CMCM CM CMCM CMCM CMCM CM CMCM CM CM CM CM 

O — O O O O 

o — <r> —i co —i 

O CO O O O CO 

o 

o 

-

CM 

CM 

o 

CM 

o 

CO 

o 

o 

o 

o 

Th 

o 

CM 

CM 

o 

o 

o 

o 

-

-

-

o 

CO 

o 

o 

o 

o 

- S 

S "2 •&. o 

o 

£ g E S B 

v -s ~3 •? -c 

re -o 

re a 

<D ^ 

rS -=,-~ 

2 e < CO 5 w 3 2 

x j 

2 g 

j . u © 

OT e« -C 

^ j -Si 

U ^ 

- V 

*"* CM • ^ 

OJ 3 re 

R 

&> re - 2 

c

 £ 

11i||i S C 2d b 

'£l 

3 .B •« -2 u 

8 ^ h 

2 5 

» » « 

5 3 >- -S — 

g *8 J= -a > 

•S « .S I 3 

<5/) s_ 

q H 

1 J 

is 

«5 

co CD O I-H CM CO i n UD i-» co 

r ~ — .- < in oi in 

CM CT) CM 

Th 

Th 

o 

CO 

CM 

CO 

CM 

O 

CO 

o 

CM 

CM O Th 

cDincocoino—'• * — ooco o 

r-cr>inino~)CMco-^<X)—'^-^^i n 

cMinocMmof-^cooc o — — o 

coinincO'—icoococo^mr^— ' 

cMinocoocMoooThvo — io 
NUMBER 12 119 

co m <£> 

CO COC O CO CM CM CM CM 

CO COCO CO CT) O) <T, CT> o o o o 

inocMoo—'OOCM O 

CMinT h — — o —• — o o 

(MO O — — o— ' —' 

o o — 

O —i —' 

CM uo — co 

o o o o o o o o 

O O O O —• —' o o 

—< o o o o o o o O O —< —' 

O O —' O O — — O O O —• —" o o o o 

O — CM —. o o — — o o — — o o o o o o 

£N 

a J3 

r^> flj 

>-> 

•3s 

* w "S. 

-c 

«, 

CUS 

u 

ifl 

s 

pustu 

a ^ a 

*o 

^ 

bling; 

X 

S -2 be ia ~ c 

bo a 

be •§ 

C 

1 0 

>*•§ -M 2 a •« 8 -s » -s •« 

l&i iala j lis I aid!<5 

•5 ' — ^-w r^* ^** "^C fr Cs sii 

« re 

53 OT 

-3-9 

03 5 § Q fc 0 

R.S 

r ^ r~~ t^- » ^ r ~ 

CM CM CM CM CM CM CM 

^O 

o o 

O —i 

o o 

o o 

o — 

o — — 

o o 

o o 

o o 

O —i 

o o o o 

o o o o 

o o —> —i 

—i — o o 

o o o o 

CM CM CM CM 

O O O —• 

O O — O 

—' —i O O 

o o 

o o 

Th co 

o Th 

CO CM 

— o 

> 3 

' — i ida 

V 

J3 

OT 

i i sici nic 

1*3 

u 

-O 

T 

o 

51 

G 

-a 

© 

- V dar 

a, 

rill 

!H 

> 

a 

3 

a rrat 

O 

2 -S 

a 3 J J. 

•2 C 

2

 w 

a 1 

s

 s 

•S -2 

.a re 

5 "S 

g. 

3 > 

<3 

o -o 2 •? £ 

fi a h 

E ~ — 

re 

.c 

6 3 

«3 OT ^ Zti 

.3 3 J3 

^T 03 
120 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Results 

Polychaetes and mollusks represented 51% of 

1960 individuals collected in 20 core samples. 

Nematodes, oligochaetes, and arthropods comprised 47% and miscellaneous other groups (predominantly sipunculids, anthozoans, and echinoderms) comprised 2% of the total macrobenthos. Numerical dominance and the fact that our 

species identifications were most complete for 

polychaetes and mollusks led us to assume that 

these taxonomic groups adequately represent the 

macrobenthos at the community level of organization. A comparison of rankings of polychaetes 

and mollusks from all five stations (Table 10), 

using a grand mean of two specimens per species 

per station as a criterion, shows that nine species 

were common to both bare sand and areas containing Thalassia; only three species {Platynereis 

dumerilii (Audouin and Milne-Edwards), Capitella 

capitata (Fabricius), and Phyllaplysia engeli Marcus) 

were found in seagrass alone. Mean densities 

(extrapolated) of 8000/m2 (polychaetes and mollusks) were sampled in bare sand and 6476/m 

(polychaetes and mollusks) in Thalassia. 

The faunal list given in Table 10 is not intended to be all inclusive. Several species found 

among grass blades from the clipped meter square 

areas did not appear in the core samples, for 

example, the mollusks Astrea phoebia Roding, Tricolia thalassicola Robertson, and Alaba incerta (Orbigny). 

Figure 73 shows numbers of individuals and 

species of polychaetes and mollusks per sample at 

each of the five stations. The number of individuals at station 3 was significantly (SNK) different 

70 

o 60 

o per Individuals 

"o 

| 3 0 

z 

20 

i 

_ 

-

i 

A 

_ 

r -

-

i II I 

5 

(-5m) 

Buoy 

(0m) 

4 3 

f*5m) (-»12m) 

Stations 

2 

H8m) 

1 

(+23m) 

FIGURE 73.—Numbers of individuals and numbers of species 

of polychaetes and mollusks per core (x + Sx) at five stations 

in the Carrie Bow Cay lagoon. 

5.0 

4.8 

4.6 

4.4 

4.2 

4.0 

1 

-

i 

A 1 II I 

» 

^^^\ K / \ 

"""""^ N. / \ 

\ / \ 

\ • 

i II I 

0.8 

0.7 

LU 

0.6 

0.5 

0.4 

10.0 

9.0 

•°8.0 

7.0 

6.0 

-

-

-

i i iii -

o-_ 

1 I 1 1 1 

1 1 1 1 1 

..A.. A 

...-•"" ' ""••- , ,-•• " 

..••"" -A-"' A" 

A 

5 

(-5m) 

Buoy 

(0m) 

4 3 

(+5m) (+12m) 

Stations 

2 

(+18m) 

1 

(+23m) 

FIGURE 74.—Species diversity (H' with solid lines and solid 

circles), species evenness (E' with dashed lines and open 

circles), and species richness (d with dotted lines and open 

triangles) of polychaetes and mollusks at five stations in the 

Carrie Bow Cay lagoon; each sample consists of four replicate cores. 
NUMBER 12 121 

TABLE 11.—Standing crop of Thalassia per square meter at three stations in the Carrie Bow 

Cay lagoon; dry weights and percentages of organic and inorganic (after ashing) fractions 

compared for live and dead blades 

Stati No. of 

shoots 

Total dry 

weight (g) 

Live blades Dead blades 

WL (g) Org. (%) Inorg. (%) Wt. (g) Org. (%) Inorg. (%) 

117 

136 

422 

104.04 

79.59 

338.07 

58.73 

57.49 

178.67 

51 

51 

52 

49 

49 

48 

45.31 

22.10 

159.40 

27 

34 

31 

73 

66 

69 

.? 0 

01 

5 

-1.0 0.0 +4.0 

40 

30 

20 

10 

n 

- Station 4 -. 

/S\ . 

S* > . 

" ^ ^ , \ " 

40 

30 

20 

10 

n 

i i i 

Station 5 

T I 1 i 

1 1 

" \ 

\ 

* 

+1.0 +2.0 +3.0 

Sediment Size (0) 

FIGURE 75.—Sediment size analyses at five stations in Carrie 

Bow Cay lagoon. Weight percentages of sediment sizes (—1.0 

to +4.0 d>) at each station. 

(95% level of confidence) from that at station 5. 

All other stations, whether containing Thalassia 

or no vegetation, were not significantly different 

from one another. Whe n numbers of species at all 

stations were compared, stations 2 and 4 were 

significantly (SNK) different (95% level of confidence) from stations 1 and 5. Station 3 was not 

significantly (SNK) different from all other stations. 

Figure 74 shows that species diversity and richness indices for all stations except station 5 reflect 

the same trend shown by numbers of species. Th e 

higher species diversity at station 5 (dense Thalassia) compared to stations 1 and 3 (bare sand) 

was largely due to the effect of high species 

evenness at station 5. Species evenness was higher 

in all seagrass stations than in bare sand stations. 

Grass blades included in core samples contained eight polychaetes that had burrowed into 

the tissue of the grass blade, in some cases leaving 

a trail of fecal pellets behind them. These represent a variety of families: three capitellids, three 

syllids, one maldanid, and one sabellid. This phenomenon is not explained. 

Although 216 additional specimens of polychaetes and mollusks were added by the 0.5 m m 

mesh from five cores (one from each station), only 

one species (the mollusk, Caecum ryssotitum Folin) 

was found that did not occur in 1.0 mm mesh 

samples. 

Gravimetric measurements (Table 11) of both 

live and dead blades of Thalassia clipped from 

one meter square areas adjacent to stations 2, 4, 

and 5 show that seagrass standing crop increased 

from the patch at station 2 to the dense growth 

at station 5. Overall, 49% of living plant material 

and 70% of dead grass were inorganic in composition, whereas 51% and 30% respectively were 

organic. 

Sediment size analyses (Figure 75) indicated 

that the percentage of fine sand (2 and 3 <£) 
122 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

relative to coarse sand (—1 and 0 <|>) increases 

from station 1 to station 5. 

An examination of 20 randomly selected Thalassia blades with bite marks revealed 5.5±0.9 {x 

+ Sx) bites per blade, with a diameter of 7.8±0.3 

mm {x + Sx). 

Discussion 

Although extrapolated densities of macrobenthos per meter square in the Carrie Bow Cay 

habitat (total fauna: 12,167/m2 in Thalassia; 

16,750/m2 in bare sand) fall within the ranges 

reported for subtropical and tropical locations 

(Table 12), this study does not show a direct 

relationship among seagrass standing crop, species densities, and numbers of species. Although 

some invertebrates may benefit from the protection and living space afforded by blades, roots, 

and rhizomes, the overall composition of seagrass 

and bare sand assemblages of macrobenthos indicates that there is no clear-cut effect of seagrass 

versus bare sand on species densities. In fact, 

species densities are significantly (SNK) higher at 

one of the bare sand stations (station 3) than at 

the dense seagrass station (station 5). Numbers of 

species from seagrass stations differ significantly 

(SNK) from those collected in bare sand stations, 

with the exception of station 5. Numbers of species at station 5, in dense Thalassia, do not significantly differ from those in bare sand stations 

(stations 1 and 3). 

The abundant species of polychaetes and mollusks are found in both sand and seagrass. Among 

the polychaetes, seven of the ten top ranked 

species are common to all stations. Only three 

species, the polychaetes Platynereis dumerilii and 

Capitella capitata and the opisthobranch Phyllaplysia engeli, have relatively high densities in seagrass 

and not in bare sand. In this habitat, all three 

species live closely associated with seagrass blades: 

P. dumerilii builds tubes on the blades, C. capitata 

tunnels within the tissue of the blades, and P. 

engeli feeds upon seagrass epiphytes. 

In other studies comparing animal assemblages 

of sand and seagrass, the water depths, tidal 

range, hydrography, and sediment characteristics 

have differed from those considered here. For 

example, along the sampling transect at Carrie 

Bow Cay the sediment interface between Thalassia beds and adjacent bare sand is imperceptible 

and not evidenced by erosional ridges (Jackson, 

1972) or sand bars (Orth, 1977). Nor is there any 

indication in the study area of erosion around the 

rhizomes of seagrass plants or exposure of the root 

system, in a pattern of "raised terraces" (Kikuchi 

and Peres, 1977). In other words, the transition 

from sand to seagrass off Carrie Bow Cay is 

marked only by the advent of vegetation and by 

a gradual lagoonward decrease in sediment grain 

size. There is no evidence of sedimentary bedload 

movement in the form of ripple marks (Santos 

and Simon, 1974). Because the bare sand areas 

studied are physically stable, no effect of sediment 

instability on densities or species composition of 

macrobenthos is evident. 

The overall small size of specimens indicates 

that few macrobenthic invertebrates survive to 

maturity in this habitat. Only a few specimens 

fall within the mature size ranges reported in the 

literature; the lack of such larger sizes off Carrie 

Bow Cay implies high and persistent predation. 

The Thalassia and adjacent bare sand environments here may be defined, according to the 

model of Connell (1975), as having benign physical conditions in which predators usually kill all 

young colonists each year, and thereby reduce 

competition for resources (for instance, food and 

space) and help to perpetuate a state of nearly 

continual recolonization. Connell's model accounts for an occasional event in time and space 

wherein certain prey species can reach invulnerable sizes. 

While seasonality may be less influential in a 

tropical than a temperate habitat, seasonal differences in densities of macrobenthos clearly 

should be expected in the habitat studied here. 

Nevertheless, our expectation that macrobenthic 

densities would correlate positively at any given 

time with seagrass was not substantiated by the 

evidence from this study. 

In Carrie Bow lagoon, a majority of Thalassia 
NUMBER 12 123 

TABLE 12.—Published data on densities of macrobenthic invertebrates per square meter in 

seagrass and bare sand from subtropical and tropical benthic studies 

Reference 

Bloom et al., 1972 

Virnstein, 1972 

Santos and Simon, 

Young et al., 1976 

McNulty, 1970 

Brook, 1975 

Brook, 1978 

Murina et al., 1973 

Young and Young, 

1974 

this study 

Seagrass 

-

33,485° 

3994-14,236 

-

Substrate 

4000 (1972) 

1895 (1973) 

292-10,644 

4508 

583 

12,167 

6476" 

Bare sand 

510 

17,151 

8795" 

-

291 (1956) 

108 (1960) 

-

-

354 

16,750 

8000* 

Sieve mesh size (mm) 

1.0 

0.5 

0.5 

1.0 

1.0 

1.0 

1.0 

1.0 

1.0 

1.0 

Location 

Tampa Bay, Florida 

Tampa Bay, Florida 

Tampa Bay, Florida 

Indian River, Florida 

Biscayne Bay, Florida 

Card Sound, Florida 

Biscayne Bay, Florida 

Florida Bay, Florida 

Cuba 

Carrie Bow Cay lagoon, 

Belize 

° Polychaetes only. 

Polychaetes and mollusks only. 

blades were thickly encrusted with epiphytic coralline algae at the time of sampling. Humm 

(1964) notes that such encrusted blades sink to 

the bottom when they are broken off the parent 

plant and are not readily carried to other areas 

by tidal currents as are nonencrusted blades. In 

addition, observations of seagrass along the transect of this study suggest that individual Thalassia 

blades are increasingly weighted down by heavy 

growths of encrusting coralline algae at their tips, 

so that dead blades are commonly retained in 

situ. Gravimetric measurements of live and dead 

seagrass clipped from meter-square plots at stations 2, 4, and 5 show that 49% of the total 

biomass of live grass and 70% of dead seagrass consisted of inorganic material. Microscopic 

examination of this inorganic residue revealed 

mainly fragments of coralline algae. It is likely 

that coralline algal epiphytes on Thalassia contribute substantial carbonate sediments to the 

lagoon at Carrie Bow in a manner similar to that 

reported in Florida (Humm, 1964) and in Jamaica (Land, 1970). Patriquin (1972) estimated 

that the annual carbonate production by encrusting coralline algal epiphytes on Thalassia in Barbados is 2800 g/m2

. 

Grazing fish help to distribute a portion of the 

epiphytic crust on seagrass blades—sometimes 

widely beyond the seagrass beds (Randall, 1967). 

Numerous reef fishes, in mixed schools (mainly 

scarids and acanthurids), were observed moving 

between the patch reefs of the back reef area and 

the seagrass of the lagoon. Although schools of 

fish were observed among the seagrasses, individual fish appeared to feed on seagrass blades and 

the sediment between. Direct feeding on seagrass 

is evident from many bite marks on the edges of 

the blades. Thomas et al. (1961) reported similar 

crescent-shaped bites on Thalassia blades, and 

Zieman (1974) attributed these bite marks to the 

scarid, Sparisoma radians. Conspicuous bands of 

bare sand between reef areas and adjacent seagrass beds in the Virgin Islands have been related 

by Randall (1965) to heavy grazing by parrot fish 

(scarids) and surgeon fish (acanthurids). Primary 

invertebrate grazers of seagrass, such as echinoids 

(Margalef and Rivero, 1958), were not observed 

in the Carrie Bow lagoon. 

Until night observations are made, we cannot 

determine whether invertebrate-feeding fishes 

such as pomadasyids (grunts) migrate nocturnally 

from the reefs to the seagrasses of the lagoon, as 
124 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

described in the West Indies by Kikuchi and 

Peres (1977). Decapod crustacean predation on 

other members of the macrobenthos may be important as a regulator of species densities, as 

suggested by Young et al. (1976) in Florida seagrasses. Caging experiments in both seagrass and 

bare sand areas, as in the Indian River lagoon, 

Florida, by Virnstein (1978), would help to determine the relative impact of fish and decapod 

predation in the Carrie Bow lagoon. 

This study provides only indirect evidence to 

suggest that "within-community" predation may 

be intense among the macrobenthic species of 

Carrie Bow. Large numbers of bored shells of 

bivalves and gastropods were found in our samples; in all cases, numbers of bored shells greatly 

exceeded numbers of live shells. The only drill 

species was Polinices lacteus (Guilding), a naticid 

reportedly responsible for heavy predation on a 

wide variety of mollusks in Thalassia beds in 

Jamaica (Jackson, 1972). Polychaetes of the family Syllidae were extremely abundant in samples 

from both bare sand and Thalassia of Carrie Bow 

lagoon; five species of syllids are ranked among 

the 10 most abundant macrobenthic species. Murina et al. (1973) also reported large numbers of 

syllids in Cuban seagrass beds. Syllids are believed to be carnivorous (Pettibone, 1963; Day, 

1967), preying on other invertebrates. As often is 

the case, ecological information is lacking for 

most of the species collected here, so that details 

of predator-prey interactions among the macrobenthos remain largely speculative. 

Conclusions 

The data from this study do not substantiate 

the generalization that seagrasses support a more 

diverse and dense invertebrate fauna than areas 

without seagrasses. There are no significant differences in species densities of macrobenthos between seagrass and bare sand off Carrie Bow Cay. 

Significant differences do not exist among numbers of species of polychaetes and mollusks from 

all seagrass stations and from bare sand stations. 

We suggest that the primary mechanism responsible for the regulation of macrobenthic species densities and diversities in the Carrie Bow 

lagoon is high and persistent predation. Intense 

predation pressure probably limits the presumed 

advantages of the more variable habitats and 

increased living space provided by seagrasses. 

Predator-prey interactions could be clarified by 

the use of field-experimental techniques such as 

caging. 

Sediment instability appears to be unimportant in the regulation of macrobenthic species 

densities and diversities in the lagoon. 

Few macrobenthic species are associated exclusively with seagrasses off Carrie Bow Cay. The 

majority of species sampled are common to both 

bare sand and seagrass stations. 

Sediment size analyses showed a progressive 

increase of fine-grained sediments from bare sand 

into seagrasses. Important contributors to these 

fine sediments are coralline algal epiphytes on 

Thalassia. 

Literature Cited 

Bloom, S. A., J. L. Simon, and V. D. Hunter 

1972. Animal-Sediment Relations and Community 

Analysis of a Florida Estuary. Marine Biology, 13: 

43-56. 

Brook, I. M. 

1975. Some Aspects of the Trophic Relationships among 

the Higher Consumers in a Seagrass Community 

(Thalassia testudinum Konig) in Card Sound, Florida. 133 pages. Ph.D. dissertation, University of 

Miami, Coral Gables, Florida. 

1978. Comparative Macrofaunal Abundance in Turtle 

Grass (Thalassia testudinum) Communities in South 

Florida Characterized by High Blade Density. 

Bulletin of Marine Science, 28:212-217. 

Buzas, M. A., and T. G. Gibson 

1969. Species Diversity: Benthic Formaminifera in Western North Atlantic. Science, 163:72-75. 

Connell, J. H. 

1975. Some Mechanisms Producing Structure in Natural Communities: A Model and Evidence from 
NUMBER 12 125 

Field Experiments. In M. L. Cody and J. M. 

Diamond, editors, Ecology and Evolution of Communities, pages 460-490. Cambridge, Massachusetts: 

Belknap Press. 

Dahl, A. L. 

1973. Surface Area in Ecological Analysis: Quantification of Benthic Coral-Reef Algae. Marine Biology, 

23:239-249. 

Day, J. H. 

1967. A Monograph on the Polychaeta of Southern 

Africa, Part I: Errantia. British Museum (Natural 

History), Publication, 656: 458 pages. London. 

Fenchell, T. 

1977. Aspects of the Decomposition of Seagrass. In C. P. 

McRoy and C. Helfferich, editors, Seagrass Ecosystems, 4:123-146. New York: Marcel Dekker. 

Folk, R. J. 

1974. Petrology of Sedimentary Rocks. 182 pages. Austin, 

Texas: Hemphill Publishing Company. 

Gaufin, A. R., E. K. Harris, and H. J. Walter 

1956. A Statistical Evaluation of Stream Bottom Sampling Data Obtained from Three Standard Samplers. Ecology. 37:643-648. 

Ginsburg, R. N., and H. A. Lowenstam 

1958. The Influence of Marine Bottom Communities on 

the Depositional Environment of Sediments. Journal of Geology, 66:310-318. 

Heck, K. L., Jr., and G. S. Wetstone 

1977. Habitat Complexity and Invertebrate Species 

Richness and Abundance in Tropical Seagrass 

Meadows. Journal of Biogeography, 4:135-142. 

Humm, H. J. 

1964. Epiphytes of the Seagrass, Thalassia testudinum, in 

Florida. Bulletin of Marine Science Gulf and Caribbean, 

14:306-341. 

Jackson, J.B.C. 

1972. The Ecology of Molluscs of Thalassia Communities, Jamaica, West Indies, II: Molluscan Population Variability along an Environmental Stress 

Gradient. Marine Biology, 14:304-337. 

Kikuchi, T., and J. M. Peres 

1977. Consumer Ecology of Seagrass Beds. In C. P. 

McRoy and C. Helfferich, editors, Seagrass Ecosystems, 4:147-194. New York: Marcel Dekker 

Kjerfve, B. 

1978. Diurnal Energy Balance of a Caribbean Barrier 

Reef Environment. Bulletin of Marine Science, 28: 

137-145. 

Land, L. S. 

1970. Carbonate Mud: Production by Epibiont Growth 

on Thalassia testudinum. Journal of Sedimentary Petrology. 40:1361-1363. 

Margalef, R. 

1958. Temporal Succession and Spatial Heterogeneity 

in Natural Phytoplankton. In A. A. Buzzati-Traverso, editor, Perspectives in Marine Biology, pages 

323-349. Berkeley and Los Angeles: University of 

California Press. 

Margalef, R., and J. A. Rivero 

1958. Succession and Composition of the Thalassia Community. In Association of Island Marine Laboratories, 

Second Meeting, Bermuda, Sep 17-21, 1958, pages 19-

21. 

McNulty, J. K. 

1970. Effects of Abatement of Domestic Sewage Pollution on the Benthos, Volumes of Zooplankton, 

and the Fouling Organisms of Biscayne Bay, Florida. Studies in Tropical Oceanography, 9: 107 pages. 

Miami, Florida: University of Miami. 

Murina, V. V., V. D. Chukhchin, O. Gomez, and G. Suarez 

1973. Quantitative Distribution of Bottom Macrofauna 

in Upper Sublittoral Zone of Northwestern Part 

of Cuba. In Investigations of the Central American Seas, 

pages 242-259. New Delhi: Indian National Scientific Documentation Centre. 

Orth, R. J. 

1977. The Importance of Sediment Stability in Seagrass 

Communities. In B. C. Coull, editor, Ecology of 

Marine Benthos, 6:281-300. Columbia, S.C: Belle 

Baruch Library in Marine Science. 

Patriquin, D. G. 

1972. Carbonate Mud Production by Epibionts on Thalassia: An Estimate Based on Leaf Growth Rate 

Data. Journal of Sedimentary Petrology, 42:687-689. 

Pettibone, M. H. 

1963. Marine Polychaete Worms of the New England 

Region, I: Families Aphroditidae through Trochochaetidae. United States National Museum Bulletin, 

227: 356 pages. 

Pielou, E. C. 

1966. The Measurement of Diversity in Different Types 

of Biological Collections. Journal of Theoretical Biology, 13:131-144. 

Randall, J. E. 

1965. Grazing Effect on Seagrasses by Herbivorous Reef 

Fishes in the West Indies. Ecology, 46:255-260. 

1967. Food Habits of Reef Fishes of the West Indies. 

Studies in Tropical Oceanography, 5:665-847. Miami, 

Florida: University of Miami. 

Reise, K. 

1977. Predation Pressure and Community Structure of 

an Intertidal Soft-Bottom Fauna. In B. K. Keegan, 

P. O. Ceidigh and P.J.S. Boaden, editors, Biology 

of Benthic Organisms, pages 513-519. New York: 

Pergamon Press. 

Santos, S. L., and J. L. Simon 

1974. Distribution and Abundance of the Polychaetous 

Annelids in a South Florida Estuary. Bulletin of 

Marine Science, 24:669-689. 

Sokal, R. R., and F. J. Rohlf 

1969. Biometry: The Principles and Practice of Statistics in 
126 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Biological Research. 776 pages. San Francisco: W.H. 

Freeman and Company. 

Thomas, L. P., D. R. Moore, and R. C. Work 

1961. Effects of Hurricane Donna on the Turtle Grass 

Beds of Biscayne Bay. Florida, Bulletin of Marine 

Science Gulf and Caribbean, 11:191-197. 

Virnstein, R. W. 

1972. Effects of Heated Effluent on Density and Diversity of Benthic Infauna at Big Bend, Tampa Bay, 

Florida. 59 pages. M.A. Thesis, University of 

South Florida, Tampa. 

1978. Predator Caging Experiments in Soft-Sediments: 

Caution Advised. In M. L. Wiley, editor, Estuarine 

Interactions, pages 261-273. New York: Academic 

Press. 

Young, D. K., M. A. Buzas, and M. W. Young 

1976. Species Densities of Macrobenthos Associated with 

Seagrass: A Field Experimental Study of Predation. Journal of Marine Research, 34:577-592. 

Young, D. K., and M. W. Young 

1978. Regulation of Species Densities of Seagrass-Associated Macrobenthos: Evidence from Field Experiments in the Indian River Estuary, Florida. Journal of Marine Research, 36:569-593. 

Zieman, J. C. 

1974. Quantitative and Dynamic Aspects of the Ecology 

of Turtle Grass, Thalassia testudinum. In Recent Advances in Estuarine Research, Proceedings of the 2nd 

International Estuarine Research Conference, Myrtle 

Beach, South Carolina, October, 1973, pages 541-562. 
A Submarine Gave near Columbus Cay, Belize: 

A Bizarre Cryptic Habitat 

Ian G. Macintyre, Klaus Riitzler, 

James N. Norris, and Kristian Fauchald 

ABSTRACT 

An unusual cryptic habitat having an extensive 

covering of serpulid worms has been discovered 

in a submerged Pleistocene cave in the Belize 

barrier-reef platform near Columbus Cay. Aggregates of serpulids, which have been named "pseudostalactites," project from the ceiling of this cave 

in the direction of a narrow opening (10 m long 

and less than 3 m wide) that breaches the roof of 

the cave at a water depth of 17 m. Apparently 

this opening has restricted the movement of water 

within the cave so that serpulid worms have 

become more abundant on the ceiling than other 

cryptobiota, which include some sponges, filamentous algae, mollusks, and bryozoans. The 

latter group, with the exception of boring bivalves, occurs only within a radius of 25 m from 

the entrance of the cave. The pseudostalactites 

are composed mainly of serpulids belonging to 

two species of the Vermiliopsis glandigera infundibulum group and extend at least 40 m from the cave 

opening, which is the limit of our observations. 

Varying amounts of submarine cement consisting 

of magnesium calcite form a coating on, or matrix 

in, the serpulid aggregations. A barren sediment 

cone (very fine sand to mud) occurs at a depth of 

30 m below the cave opening. 

and 5 cm in diameter that he had collected from 

a submarine cave near Columbus Cay. This fragment had two surprisingly different sides: one 

composed predominantly of serpulid tubes and 

the other composed of dense microcrystalline Mg 

calcite forming a knobby surface. In our subsequent search for the source of this sample, we 

discovered an unusual megacryptic environment. 

This paper presents the results of our preliminary 

investigation of the biological and geological 

characteristics of a highly unusual habitat. 

ACKNOWLEDGMENTS.—We are grateful to H. 

Bowman, Jr., and A. Usher of Dangriga for bringing this submarine cave to our attention. We 

thank K. E. Bucher, W. Gerwick, G. L. Hendler, 

P. M. Kier, W. M. Kier, R. J. Larson, K. Muzik, 

A. B. Rath, and P. E. Videtich for their field 

assistance. G. L. Hendler, W. M. Kier, and A. B. 

Rath also took underwater photographs. For help 

with identifications we thank A. H. Cheetham 

(bryozoans), I. M. Goodbody (tunicates), and R. 

S. Houbrick and T. R. Waller (mollusks). R. B. 

Burke provided the aerial photograph of the cave 

site and V. Krantz provided other photographic 

assistance. I. Jewett drafted the diagrams. 

Introduction 

In 1977 a Belizean SCUBA diver sent us a 

"stalactite" sample approximately 15 cm long 

Ian G Macintyre, Department of Paleobiology; Klaus Riitzler and 

Kristian Fauchald, Department of Invertebrate Zoology; and James 

N. Norris, Department of Botany, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C. 20560. 

Description of the Submarine Cave 

AEREAL SETTING.—The submarine cave about 

1 km northwest of Columbus Cay (17°00'N, 

88°02'W) occurs in a Thalassia-covered lagoon (5 

m deep) at a distance of about 3 km inside the 

outer edge of the Belizean barrier reef platform 

(Figure 76). The only other natural depression in 

127 
128 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

88°10'W .^Mosquito Cay 

j A Columbus Cay 

\ & Cross 

C<> p Cay<si7ffi> 

A<S?GarboHs £ f J 

Cay f/7j i/J e 

8 

Coco 

Plum 

Cay 

Blue 

Ground 

Range 

Stewart 

'•'C? 

//* Tobacco Cay 

Tobacco Cay 

Entrance 

16°50'N 

Cay k 7 /QVVfee Wfee 

South Water Cay 

Carrie Bow Cay 

Curlew Bank 

N 

5 km. 

FIGURE 76.—Map showing location of submarine cave near Columbus Cay, Belize. 

the smooth Thalassia meadow is a sand-filled 

sinkhole 200 m southeast of the cave (Figure 77). 

The sinkhole (50 m in diameter) has a smooth 

sandy bottom (10 m deep) that is ringed by a rich 

coral community established on rocky ledges and 

dominated by the corals Montastrea annularis (Ellis 

and Solander), Acropora cervicornis (Lamarck), Diploria strigosa (Dana), Porites astreoides Lamarck, 

Porites porites (Pallas), Montastrea cavernosa (Linnaeus), Agaricia agaricites (Linnaeus), along with 

the octocoral Briareum asbestium (Pallas), and the 

sponges Niphates digitalis (Lamarck), Iotrochota birotula (Higgin), and Amphimedon compressa (Duchassaing and Michelotti). 

FIGURE 77 (opposite page, top).—Aerial view of location of 

submarine cave (sc) and sinkhole (sh); regularly spaced and 

aligned patches of bare sand in the Thalassia-covered lagoon 

delineate seismic shot-hole (ssh) patterns that have not been 

overgrown since seismic work was carried out here in the 

early 1960s; diameter of sinkhole in foreground is 50 m. 

FIGURE 78 (opposite page, bottom).—Cave entrance; note 

sediment-laden algal mat (am) covering the surface sloping 

into the cave; water depth is 17 m. 
NUMBER 12 129 
130 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

5 -

10-

15-

2 0 -

Q. 

03 

Q 

"5 

2 5 -

3 0 -

3 5 -

4 0 -

4 5 -

y.y.V..y:..y,y y. 

Sediment 

11 | | Limestone 

[v v| Thalassia 

[ g j Sponge .

:

;v; 

|j|] Coral 

[ f ^| Octocoral 

Serpulid projection 

17m. 

~T~ 

25 20 25 30 40 ~~r 

20 

~T~ 

10 10 15 35 

South Horizontal Distance (m) North 

FIGURE 79.—Diagrammatic sketch of cave in cross section; note large sediment cone directly 

under cave entrance. 

Interspersed in the Thalassia testudinum Banks 

ex Konig bed are several dense and large calcified 

green algae, mainly Penicillus pyriformis f. explanatus 

Boergesen and Udotea wilsonii Gepp and Howe ex 

A. and E. Gepp, with lesser amounts of Halimeda 

simulans Howe, Rhipocephalus phoenix (Ellis and 

Solander) Kiitzing, and Penicillus pyriformis A. and 

E. Gepp. Brown algae, however, are dominant 

among the coral heads, primarily Dictyota bartayressii Lamouroux, Padina sp., and Lobophora variegata (Lamouroux) Womersley (all Dictyotales; 

Dictyotaceae), as well as Sargassum hystrix 

J. Agardh (Fucales; Sargassaceae). 

In contrast to the clear water and rich growth 

of corals at the sinkhole, the waters near the cave 

are turbid and the sea floor adjacent to the 

opening is covered by a few hardy scleractinian 

corals such as Montastrea cavernosa, M. annularis, 

Porites porites, Mussa angulosa (Pallas), and Scolymia 

sp., together with the octocorals Briareum asbestium 

and Plexaura sp., and some sponges typical of 

seagrass bottoms in the lagoon of the barrier-reef 

platform, branching Aplysina fulva (Pallas), Amphimedon compressa and Iotrochota birotula, vaseshaped Niphates digitalis and Callyspongia vaginalis 

(Lamarck) and massive Ircinia strobilina (Lamarck) . 

ENTRANCE ZONE.—An extensive sedimentladen, green algal mat up to 8 cm thick and 1-2 

m wide surrounds the entrance to the cave at a 

depth of 17 m (Figure 78). The sediment is very 

fine sand to mud size carbonate containing an 

abundance of sponge chip material. The dominant algal component is the turf/mat-forming 

Cladophoropsis membranaceae (C. Agardh) B0rgesen 

(Siphonocladales; Siphonocladaceae), a siphona-
NUMBER 12 131 

'*&•<• • 

v? a 

-'- JL 

Wv ' 

1 m A 

*~ir~ 

fci^V 

'VTi 

m • • 

a 

•W- & Ejjitr" «.^^ ^ 

A> % •' 

B ' 

HKBIKM" ^ 

- Jf" 1 

,»--'~ 

> p 

Hnd§H.\ ^ 

B^k ^ l 

^BB 

rf^ 

BBBfctBfl 

•iW 

^.^^kX" -v A.i?V H^^H •i 

^|JIV' - BK«fe*a%fK^BBl E -*y| 

PF- BBF / : /- ) 

- HP ^ m.4*#;'•• ' %r 

JfJBjh ^-^R B/' ijJJ-'^ 

§ J i-'mP'Mmto- . < 

44W , iB** J F Vxl •BF frw > ••'*-1m •' .BBr' LJBB T ' #)' . TBBB'P!^'^ -^B/ 

'i*r

 %±-^ttiw •/ wr 4BB 

«k V **T £% • !^9- • f'•••rjm-:,*fr 

UBB> ^SwBte^' V/J^s, • •' m t •' /JjB 

y

«^5BBBBMiLi._i^BF'S.-,. •••....jfflBM. TBJifc*^^...**.--.2B« B 

* M H ^ " SBB W 

: ^^S>-z^*^*'' > 

j y 

FA 

^SP 

^ a -liBBBBl B ^ ''"^ # " Bf ^ 

^' ^ ^ ^ | BW^E T • -a, -jf m 

<tit4~* 7 / _^» ~ .^MB B IB"--— KJHk&rAx. 

IJBBJ H •'<BB' "*5 MUEMBB* ' ^*B E * 

•L J~SA rJL 

^^^m jl^^^-.^JH j H r '7M 

SBBBV- BBC JBBBI ^^Ubgrad 

? ' ''^Jl > £ • 

^—• 

' BBJ 

V 

r'J£>. J JP ••<• j t 

J%- f w.£# '• 

f ;M 

- • Jr . • 

y 

m&*^Wk 

it, Jj 

^"f'-^^BBS 

1 

FIGURE 80.—Underwater views of pseudostalactites: a, club-shaped and steeply inclined projections 10 m from entrance, with conch shell fragment attached for settlement study; 6, tapering 

and almost horizontal projections 20 m from entrance. (Each picture width = 1.5 m.) 
132 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 
NUMBER 12 133 

ceous green alga (mainly unilaterally branched 

and having filaments up to 145 /xm in diameter) 

that typically occurs in the intertidal zone in calm 

water, where it can form extensive mats that 

sometimes become infiltrated with fine sand 

(Taylor, 1960; Kapraun, 1972). Because fine sediment clouds the water around the cave entrance 

at the slightest motion of water, it appears that 

the area is generally calm. Although suspended 

or gradually settling very fine sediment hampers 

visibility around the entrance, apparently enough 

light is still present to permit luxuriant algal 

growth, which may be enhanced by nutrients 

retained in the fine sediments. 

Large schools of spadefish, Chaetodipterus faber 

(Broussonet), and permits, Trachinotus falcatus 

(Linnaeus), are common around the cave opening. In addition, a few cobia, Rachycentron canadum 

(Linnaeus), appear along with reef sharks, Carcharhinus springeri (Bigelow and Schroeder), which 

are known to use submarine caves as a retreat 

where they can rest undisturbed on the sediment 

floor (Clark, 1975). 

CAVE INTERIOR, GEOLOGY AND BIOLOGY.—At a 

17 m depth, the opening of the cave itself is no 

more than a large crack 3 m wide and 10 m long 

(Figure 78). The resulting poor light conditions 

within the cave as well as the considerable depth 

of water to which the cave descends makes it 

difficult to assess the overall size of the cave. At 

a point 40 m north from the opening, the cavefloor depth recorded on a line dropped to bottom 

was 47 m, which indicates a sharp slope away 

from the crest of the sedimentary cone located in 

depths of 30 m just below the opening (Figure 

79). 

The ceiling of the cave, covered by countless 

projections that are angled toward the opening, 

is a phenomenon only recently recorded in the 

FIGURE 81.—Dried samples of pseudostalactites: a, lower 

surface of club-shaped projections (15 m from entrance) 

covered by serpulid tubes; b, upper surface of specimen in a 

showing knobby calcite cement capping; c, side view of 

tapering projection (25 m from entrance) showing thick 

calcite cement coating over serpulid tubes; d, pencil-thin 

projection collected 40 m from cave entrance (limit of observation). (Scale = 0.5 cm.) 

literature. These projections consist largely of serpulid tubes and magnesium calcite cement, and 

Macintyre and Videtich (1979) have named them 

"pseudostalactites" because they superficially resemble stalactites but are entirely of marine origin. The pseudostalactite cover begins about 3 m 

from the cave opening and extends for at least 40 

m (the limit of our diving observations) at a water 

depth of 30 m (Figure 79). They slope toward the 

opening at angles varying from 40° to 60° near 

the opening to an almost horizontal orientation 

at the inner limit of observation. 

These pseudostalactites are composed of serpulid tubes and varying amounts of microcrystalline magnesium calcite cement in and around 

tubes and infilling borings. Here and there, boring and subsequent lithification have resulted in 

extensive replacement of worm-tube framework 

by magnesium calcite cement. The largest pseudostalactites occur about 10 m from the opening 

of the cave and are distinctly club shaped, in 

some cases measuring up to 30 cm in width across 

the tip (Figures 80a, 8\a,b). Within this area the 

pseudostalactites are dominantly serpulid aggregations and form an almost continuous field of 

tightly packed projections. In contrast, deeper 

into the cave, in addition to changing their angle, 

they become more widely spaced and progressively narrower, and the innermost surfaces observed are covered with almost pencil-thin projections (Figures 806, 81 c,d). The upper surface of 

most pseudostalactites consists of a layer of sediment-rich dense to chalky microcrystalline calcite. In general these sediment-rich calcite cement 

cappings are best developed on the innermost 

pseudostalactites observed (Figure 8\c). Scanning 

electron photomicrographs show that this dominantly silt-sized sediment consists mainly of chips 

produced by boring sponges (Riitzler, 1975). 

The serpulid polychaetes partly responsible for 

these projections have been identified as members 

of two species of the Vermiliopsis glandigera infundibulum group (H. A. ten Hove, pers. comm. 1978), 

one of which is a thin-walled species commonly 

referred to in the Caribbean as Vermiliopsis annulata (Schmarda), and the other a thicker-walled 

species yet to be described (Figure 82a,b). Other 
134 

FIGURE 82.—Underwater close-up of pseudostalactite surfaces: a, Vermiliopsis spp. worm aggregation with sponge Placospongia carinata; b, live Vermiliopsis sp. with tentacle crown extended. 

(Scales = 10 mm.) 

relatively common tubes, which are long and 

narrow, closely resemble those of Metravermilia 

multicristata (Philippi). There are also a few Spirorbinae tubes and some fine tubes of Josephella 

marenzelleri Caullery and Mesnic. Although the 

relative number of living serpulids could not be 

accurately ascertained, the lack of extended 

worms as well as a scarcity of worm-filled tubes 

in collected samples suggest that only about one 

percent of the surface tubes contain living serpulids. 

In the poorly lighted to dark zone of the cave 

ceiling between 1 m and 20 m from the entrance, 

sponges are the most conspicuous associates of the 

serpulid projections. Samples from this area contained gray to white Geodia gibberosa Lamarck, 

some with drooping digitate processes (Figure 

83a), a stringy yellow Aplysina sp., whitish cushions of Chondrilla nucula Schmidt, orange-brown 

cushions and crusts of Placospongia carinata Bowerbank (Figure 82a), and whitish crusts of Spirastrella coccinea Duchassaing and Michelotti. Two 

other common whitish sponges belong to the 

Lithistida, encrusting Desmanthus incrustans (Topsent) and egg-shaped Gastrophanella implexa 

Schmidt (Figure 836). Small cryptic species, including Microciona affinis (Topsent), Stellettinopsis 

sp., and Samus anonyma Gray, coat and permeate 

dead serpulid worm tubes. The orange sclerosponge Ceratoporella nicholsoni (Hickson) (Figure 

83a) is visible on a close-up photograph of the 

vertical surface of a break in the ceiling contour, 

but only a few minute specimens could be secured 

from a nearby location. 

The dominant living mollusks found in association with the pseudostalactites for about 25 m 

from the cave entrance were the nestling arcid 

bivalve, Barbatia dominensis (Lamarck), and the 

encrusting venerids, Chama congregata Conrad and 

Chama macerophylla (Gmelin). Although boring 

bivalves are common in the pseudostalactites to 

the limit of our observations, to date none have 

been collected alive. Most of these bivalves are 

well lithified in their borings by magnesium calcite submarine cement. They include Gastrochaena 

sp., Lithophaga nigra (Orbigny), Gregariella coralliophaga (Gmelin), and Coralliophaga coralliophaga 

(Gmelin). The common occurrence of as many as 

three of these bivalves, one inside another, in the 

same bore hole, indicates that many of them 
NUMBER 12 135 

FIGURE 83.—Views of cave ceiling showing sponge population: a, Geodia gibberosa (15 cm long) 

projecting from ceiling and Ceratoporella nicholsoni attached nearby (arrow); b, Gastrophanella 

implexa (5 cm diameter). 
136 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

prefer to occupy an extant boring rather than to 

form their own. 

Other biota present in this habitat include 

trace occurrences of small and seldom freshly 

preserved bryozoan colonies belonging to eight 

species of cheilostomes: Parellisina curvirostris 

(Hincks), Cribrilaria radiata (Moll), s.l., Escharina 

pesanseris (Smitt) (?), Exechonella antillea (Osburn), 

Hippopodina feegeensis (Busk), Hippothoa flagellum 

(Manzoni) (?), Tremogasterina cf. T perplexa Cook, 

and Hippoporina pertusa (Esper). Specimens of ascidians collected belong to Pyura sp., possibly near 

P. vittata (Stimpson). 

Several epizoic algae were found growing on 

the pseudostalactites near the cave opening (17 

m depth), primarily the red algae Spermothamnion 

investiens (H. and P. Crouan) Vickers (Ceramiales; 

Ceramiaceae), and Polysiphonia macrocarpa Harvey 

in MacKay (Ceramiales; Rhodomelaceae). Some 

very small specimens of the green alga, Cladophoropsis membranaceae occur here and there, and a 

single specimen of the filamentous brown alga, 

Sphacelaria sp. (Sphacelariales; Sphacelariaceae), 

which was non-fertile and could not be identified 

to species, was also recorded. Some microscopic 

blue-green algae of Calothrix sp. (Nostocales; Rivulariaceae) were also found. 

The thickness of the entire veneer of serpulidcalcite cement on the ceiling of the cave could 

not be determined, and individual projections 

could not be measured beyond 50 cm because 

they tend to coalesce as they increase in size. In 

several recessed areas, the veneer appeared to be 

poorly developed and the ceiling became a relatively smooth surface except for some rounded 

depressions that are probably solution cavities in 

the Pleistocene host limestone. Inverse steps in 

the relief of the ceiling resembling "false walls" 

to a diver also indicate that a thin Holocene cover 

is present. These steps mark areas where the roof 

slumped off during the formation of the cave in 

subaerial Pleistocene conditions. In addition, here 

and there vertical cones that are coated with 

small inclined pseudostalactites project from the 

ceiling (Figure 84). These cones are probably true 

Pleistocene stalactites. The prominence of this 

^~" - ~-

mho*-** 

HNP-A* " 

-

PgaJsT . jgifl 

j 

j yjf 

V \ 

i \ • 

FIGURE 84.^Large (1 m) vertical projection (probably true 

stalactite) with surface covered by inclined pseudostalactites. 

Pleistocene stepped surface and the presence of 

the stalactites as well as solution cavities in recessed areas indicate that the maximum thickness 

of the serpulid-cement veneer is probably less 

than 1 m. 

The crest of a large cone of sediment located in 

a depth of 30 m directly below the cave opening 

consists mainly of very fine sand to mud-sized 

sediment (Figure 79); it has a relatively smooth 

surface and shows no signs of current ripple 

marks. Some recognizable trails leading to a few 

empty Strombus shells, indicate an unsuccessful 

search for food after these animals fell through 

the cave opening. The fresh condition of these 

trails long after the death and decay of the 
NUMBER 12 137 

trapped gastropods suggest a lack of bottom currents strong enough to disturb the sediments on 

the cave floor. The flanks of this sedimentary 

cone slope away to unknown depths. 

Discussion 

CAVE BIOTA.—Characteristics of the species 

composition of the sessile benthos of marine caves 

in the euphotic zone has long been known to 

differ considerably from that of benthos in surrounding light-exposed substrates. Early work on 

shaded communities in the Mediterranean (summarized in Riedl, 1966), was followed by studies 

of coral reef tunnels off Madagascar (Vasseur, 

1974), but little attention was given to cryptofauna of the Atlantic reef systems until the discovery of sclerosponges and of a characteristic 

brachiopod-sclerosponge community under overhangs and in caverns and tunnels on the fore-reef 

slope of Jamaica (Hartman and Goreau, 1970; 

Jackson et al., 1971). The distinctive character of 

the cave faunas is principally the result of decreased illumination—lack of light excludes fastgrowing space competitors, mainly algae, and, in 

coral reefs, hermatypic hydrocorals, scleractinians, and octocorals. The available substrate inside 

a cave is generally well suited for colonization by 

sessile filter feeders because its ceiling and vertical 

walls prevent burial in sediment. Water circulation, however, must be sufficiently strong to ensure the renewal of food supplies and the transport of metabolic waste. 

Whether complete darkness may be limiting 

for organisms that do not depend on photosynthetic processes is difficult to determine because 

low light levels in shallow-water environments 

commonly occur in association with poor water 

circulation. A few observations on dark but openended (and therefore well-ventilated) tunnels 

show that predominant sessile biota consist of 

sponges, serpulid polychaetes, and a few ascidians 

and foraminifers (Vasseur, 1974), as well as some 

ahermatypic corals, sessile or boring mollusks 

{Chama sp., Lithophaga sp.), and minute bryozoans 

(Riitzler, unpublished observation from Bermuda). These data indicate that except for 

sponges, only a few representatives of a small 

number of sessile groups have adapted to complete darkness within the shallow-water environment. Dark caves in the Mediterranean having a 

deep (10-40 m) single entrance—and therefore 

poor water circulation—are reported to be colonized mainly by serpulid polychaetes, along with 

boring bivalves {Lithodomus lithophagus (Linnaeus)), scleractinians, and a few sponges (Laborel and Vacelet, 1959). From such accounts it 

appears that serpulid polychaetes can adjust more 

easily than most cave dwellers to both complete 

darkness and stagnant water conditions. 

DISTRIBUTION AND RECRUITMENT OF SERPULIDS.—The dominant serpulids in the pseudostalactites of Belize are not typical colonial organisms, for each tube is the result of the activity of 

a single individual. The projections thus represent 

a series of discrete settlement events, but we do 

not know whether the serpulids mature in the 

cave and their successive generations are recruited 

locally, or whether the settlements represent invasions from the outside. Most of the common 

serpulids in the cave near Columbus Cay mature 

in about one year, and some reproduce the year 

round. As a result, if serpulids were recruited 

locally, distinct settlement surfaces would be difficult to identify. If recruitment was the result of 

a rare event—such as a major flushing of the 

cave—then these events should be recognizable 

in the projections. The distribution of live serpulids suggests local recruitment in that a few scattered individuals were alive when the projections 

were sampled, but there were not signs of sheets 

or groups of live animals. 

The low percentage of live serpulids and other 

biota in an area of such impressive serpulid-tube 

aggregations does suggest the episodic development of this cryptic community, which is supported by observations from "Grotte de la Triperie" near Marseille (Vacelet, 1964). Vacelet 

found that the center ceiling and the entire blindending tunnel of a large subtidal grotto were 

azoic except for a cover of empty serpulid tubes 

of Serpula concharum Langerhans, Pomatostegus 
138 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

polytrema (Philippi), and Vermiliopsis infundibulum 

(Linnaeus) on the walls and on the numerous 

stalactites present in that cave. During maneuvering at the end of August 1963, a submersible 

(Soucoupe Cousteau) that entered the grotto 

stirred up the water considerably. Subsequently, 

small but live serpulids that had been established 

in the normally azoic zones in mid-October, by 

mid-January were dead. 

Because filter-feeding serpulids are rheotactic 

rather than phototactic, the orientation of the 

pseudostalactites toward the cave entrance on 

both sides of this elongate opening is probably 

related to current conditions rather than light 

conditions. The accumulation of fine sediment 

around the entrance of the cave, the preservation 

of tracks of deceased and decayed Strombus on the 

sediment floor, and the lack of any currents during diving observations suggest minimal current 

activity around the entrance and inside the cave. 

Apparently there is, however, enough movement 

of nutrient-rich water associated with the cave 

opening to influence the direction of serpulid 

growth. The abundance of clionid sponge chips 

on the upper surfaces of pseudostalactites and the 

apparent lack of excavating sponges in the cave 

indicate that the occurrence of temporary influxes 

of outside water capable of transporting this fine 

sediment into the cave. 

DISTRIBUTION OF CAVE SPONGES.—Divers studying the ecology of shallow marine caves (0-5 m) 

in the Mediterranean have found large and diverse sponge populations whose composition differs significantly from that of sponges found on 

light-exposed rock substrates in similar depths. 

Qualitative and quantitative comparisons revealed that gradients of light, water movement, 

and inclination of substrate (exposure to sedimentation) are responsible for these zonation patterns. 

Sponges known from much greater depths (10s to 

100s m) were also found in these Shallow caves 

(Sara, 1958, 1961; Russ and Riitzler, 1959; Riitzler, 1965, 1966). Similar findings have been reported in shallow reef tunnels in the Indian 

Ocean (Vacelet and Vasseur, 1965) and in deeper 

caves (4-25 m) in the Mediterranean (Pouliquen, 

1972). Among the most interesting sponges collected from dark caves were live pharetronids 

(Vacelet, 1964; Vacelet and Vasseur, 1965) and 

sclerosponges (Hartman and Goreau, 1970). In 

addition, lithistid sponges common at bathyal 

depths elsewhere have been found in shallow 

caves in Madagascar (Vacelet and Vasseur, 1971) 

and in the Mediterranean (Pouliquen, 1972). 

The sponge fauna in Columbus Cay cave differs considerably from that of the lagoon surrounding the entrance, which is similar to the 

fauna of other Caribbean lagoon habitats at similar depths, for example, near Carrie Bow Cay. 

Of the sponges in the cave, Geodia gibberosa, Placospongia carinata, and Spirastrella coccinea are species common elsewhere under rocks or back-reef 

rubble; Chondrilla nucula is one of the few sponges 

that grow abundantly either in full light, where 

it is olive to brown from symbiotic zoocyanelles, 

or in full darkness, where it is white to cream 

colored (Wilkinson and Vacelet, 1979); and Aplysina sp. has been collected only in caves, although 

morphologically it closely resembles species from 

illuminated habitats and ecologically it resembles 

Verongia {= Aplysina) cavernicola (Vacelet, 1959). 

Little information is available on the cryptic 

species found in the cave. The absence of clinoids 

in the cave is surprising because most of the 

carbonate silt on the upper surfaces of the pseudostalactites is derived from these sponges. The 

lithistid that is present {Desmanthus incrustans) is 

known also from shallow caves in the Mediterranean and Indian Ocean; although its type specimen was dredged from the Campeche Bank, Gulf 

of Mexico (unknown depth), it has not since been 

reported from the western Atlantic (Vacelet et 

al., 1976). The other lithistid in the cave, Gastrophanella implexa, has not been found since its first 

description from west of Florida (180 m depth) 

and southwest of Cuba (230 m depth) (Schmidt, 

1879:29). Although both sclerosponges and bathyal lithistids are now documented in Caribbean 

caves, no representative of the pharetronids has 

yet been found in these waters. 

Whereas the serpulid pseudostalactites occur 

along at least 40 m of the ceiling of the cave, 
NUMBER 12 139 

sponges and the few other sessile biota present do 

not extend beyond the first step in the ceiling, 

which is less than 20 m from the entrance. 

Whether or not other openings occur in deeper 

parts of the cave and possibly admit water from 

the lagoon, the sponge population near the known 

entrance appears to receive an adequate supply 

of nutrient-rich water. This is probably introduced by waves, by the activity of resident or 

visiting fishes, or by some tidal flow through the 

small entrance. The inner zones of the ceiling 

may now and then receive renewed water containing nutrients and serpulid larvae as a result 

of upheaval during severe winter storms or even 

hurricanes. 

DISTRIBUTION OF ALGAE.—The epizooic algae 

associated with the pseudostalactites near the 

entrance of the cave (17 m depth) are all filamentous, small in size, and sparsely distributed, and 

are typical of cryptic habitats as well as algal 

turfs, although elsewhere they generally occur in 

shallower depths. 

MINERALOGY OF PSEUDOSTALACTITES.—Microprobe analyses indicate that the magnesium calcite cement, which commonly constitutes more 

than half of the pseudostalactites averages 15 

mole percent MgC03. This composition together 

with the dentate crystals, peloidal texture, 

knobby surface relief, and minor amounts of acicular aragonite present in the pseudostalactites is 

characteristic of shallow-water submarine cements (Macintyre et al., 1968; Shinn, 1969; Land 

and Goreau, 1970; Ginsburg et al., 1971; Alexandersson, 1972; James et al., 1976; Macintyre, 

1977). Oxygen and carbon isotope analyses indicate that fresh water did not influence the formation of these projections (Macintyre and Videtich, 1979). Thus, petrographic and geochemical data point to a marine origin for the cave 

cement. 

Radiocarbon dates were obtained for two pseudostalactite samples: the core of a club-shaped 

specimen 5-10 m from the cave opening gave a 

date of 820±60 years B.P., and the dominantly 

Mg calcite cement cap of a specimen 25 m from 

the opening gave a date of 650±40 years B.P. 

Because these dates are uncorrected, however, 

they may be less than true dates if bomb carbon 

is present. A slow accretion of the Holocene cave 

encrustation is confirmed by the lack of measurable encrustation over a one-year period on 

conch-shell fragments nailed to the ceiling at a 

distance 10 m from the opening. All the projections in the cave, including massive club-shaped 

formations near the entrance, appear to be accreting slowly. The date of the cement cap clearly 

indicates that even the thick crusts of cement may 

have developed over a long period of time. This 

finding is surprising in light of Goreau and Land's 

(1974) observation at a blast site in the deep fore 

reef of Jamaica. They found appreciable lithification within a period of one year. 

The extensive cover of Mg calcite submarine 

cement on the ceiling of the cave confirms earlier 

observations that submarine lithification is most 

extensive in areas where little or no sediment 

accumulation occurs and where substrates are 

exposed for long periods of time (see Macintyre, 

1977). Although similar cements have been well 

documented in a variety of marine habitats, the 

processes responsible for submarine lithification 

remain unidentified. 

Conclusion 

The submerged Pleistocene cave in a Thalassiacovered lagoon of the barrier-reef platform off 

Belize is an unusual cryptic habitat having no 

known counterparts in other marine environments. The restricted flow of water in this cave 

has been more favorable to the establishment of 

serpulid worms than to other forms of encrusting 

fauna. Sessile biota, including sponges, filamentous algae, mollusks, and bryozoans are only minor constituents sparsely distributed on the ceiling of the cave over an area generally less than 

25 m from the entrance. The serpulids, on the 

other hand, have formed massive projections superficially resembling stalactites (called pseudostalactites) that are inclined at decreasing angles 

away from the entrance and oriented toward the 

cave opening. These projections are present for 

at least 40 m from the opening, but their 
140 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

shape changes gradually from club-shaped forms 

reaching 30 cm in width near the cave entrance 

to pencil-thin forms at the inner limit of our 

observation. The pseudostalactites are extensively 

infilled and encrusted by submarine cement, 

which is a magnesium calcite cement similar to 

submarine cements reported elsewhere off Belize 

as well as in other tropical reef areas of the world. 

Literature Cited 

Alexandersson, T. 

1972. Intragranular Growth of Marine Aragonite and 

Mg-Calcite: Evidence of Precipitation from Supersaturated Seawater. Journal of Sedimentary Petrology. 42:441-460. 

Clark, E. 

1975. Into the Lairs of "Sleeping" Sharks. National Geographic Magazine, 147:570-584. 

Ginsburg, R. N., D. S. Marszalek, and N. Schneidermann 

1971. Ultrastructures of Carbonate Cements in a Holocene Algal Reef of Bermuda. Journal of Sedimentary 

Petrology. 41:472-482. 

Goreau, T. F., and L. S. Land 

1974. Fore-Reef Morphology and Depositional Processes, North Jamaica. In L. F. LaPorte, editor, 

Reefs in Time and Space. Society of Economic Paleontologists and Mineralogists, Special Publication, 18: 

77-89. Tulsa, Oklahoma. 

Hartman, W. D., and T. F. Goreau 

1970. Jamaican Coralline Sponges: Their Morphology, 

Ecology and Fossil Relatives. In W. G. Fry, editor, 

Symposia of the Zoological Society of London, 25:205-

243. London: Academic Press. 

Jackson, J.B.C., T. F. Goreau, and W. D. Hartman 

1971. Recent Brachiopod-Coralline Sponge Communities and Their Paleoecological Significance. Science, 173:623-625. 

James, N. P., R. N. Ginsburg, D. S. Marszalek, and P. W. 

Choquette 

1976. Facies and Fabril Specificity of Early Subsea Cements in Shallow Belize (British Honduras) Reefs. 

Journal of Sedimentary Petrology, 46:523-544. 

Kapraun, D. F. 

1972. Notes on the Benthic Marine Algae of San Andres, 

Colombia. Caribbean Journal of Science, 12(3/4): 199-

203. 

Laborel, J., and J. Vacelet 

1959. Les grottes, sous-marines obscures en Mediterranee. Comptes Rendus de Seances de l'Academie des Sciences (Oceanographic), 248:2619-2621. 

Land, L. S., and T. F. Goreau 

1970. Submarine Lithification of Jamaican Reefs. Journal of Sedimentary Petrology, 40:457-462. 

Macintyre, I. G. 

1977. Distribution of Submarine Cements in a Modern 

Caribbean Fringing Reef, Galeta Point, Panama. 

Journal of Sedimentary Petrology, 47:503-516. 

Macintyre, I. G., and P. E. Videtich 

1979. Pseudostalactites from Submarine Cave near Columbus Cay, Belize Barrier-Reef Complex—Evidence of Extensive Submarine Lithification [Abstract]. In Program of (he American Association of 

Petroleum Geologists, Society of Economic Paleontologists 

and Mineralogists, Houston, Texas, April 1-4, 1979, 

page 125. 

Macintyre, I. G., E. W. Mountjoy, and B. F. D'Anglejan 

1968. An Occurrence of Submarine Cementation of Carbonate Sediments off the West Coast of Barbados, 

W. I. Journal of Sedimentary Petrology, 38:660-664. 

Pouliquen, L. 

1972. Les spongiaires des grottes sous-marines de la 

region de Marseille: Ecologie et systematique. 

Tethys, 3:717-788. 

Riedl, R. 

1966. Biologie des Meereshdhlen. 636 pages. Hamburg and 

Berlin: Paul Parey. 

Russ, K., and K. Riitzler 

1959. Zur Kenntnis der Schwammfauna unterseeischer 

Hohlen. Publicazione, Stazione Zoologica di Napoli, 30: 

756-787. 

Riitzler, K. 

1965. Systematik und Okologie der Poriferen aus Litoral-Schattengebieten der Nordadria. Zeitschrift 

der Morphologie und Okologie der Tiere, 55:1-82. 

1966. Die Poriferen einer sorrentiner Hbhle. Zoologischer 

Anzeiger, 176:303-319. 

1975. The Role of Burrowing Sponges in Bioerosion. 

Oecologia, 19:203-216. 

Sara, M. 

1958. Studio sui Poriferi di una grotta di marea dell 

Golfo di Napoli. Archivio Zoologica ltalia.no, 43:203-

280. 

1961. La fauna di Porifera delle grotte delle isole Tremiti. Studio ecologico e sistematico. Archivio Zoologico 

Ilaliano, 46:1-59. 
NUMBER 12 141 

Schmidt, O. 

1879. Die Spongien des Meerbusen von Mexico. 90 pages. 

Jena: Gustav Fisher. 

Shinn, E. A. 

1969. Submarine Lithification of Holocene Carbonate 

Sediments in the Persian Gulf. Sedimentology, 12: 

109-144. 

Taylor, W. R. 

1960. Marine Algae of the Eastern Tropical and Subtropical 

Coasts of the Americas. 870 pages. Ann Arbor: University of Michigan, Press. 

Vacelet, J. 

1959. Repartition generale des Eponges et systematique 

des Eponges cornees de la region de Marseille et 

de quelques stations mediterraneennes. Recuil des 

Travaux de la Station Marine d'Endoume, 26:39-101. 

1964. Etude monographique de l'Eponge Calcaire Pharetronide de Mediteranee, Petrobiona massiliana Vacelet & Levi: Les Pharetronides actuelles et fossiles. 

Recuil des Travaux de la Station Marine d'Endoume, 50: 

1-125. 

Vacelet, J., and P. Vasseur 

1965. Spongiaires des grottes et surplombs des recifs de 

Tulear (Madagascar). Recuil des Traveaux de la Station Marine d'Endoume, supplement 4:71-123. 

1971. Eponges des recifs coralliens de Tulear (Madagascar). Tethys, supplement 1:51-126. 

Vacelet, J., P. Vasseur, and C. Levi 

1976. Spongaires de la pente externe des recifs coralliens 

de Tulear (sud-ouest de Madagascar). Memoires du 

Museum National d'Histoire Naturelle, 49:1-116. 

Vasseur, P. 

1974. The Overhangs, Tunnels and Dark Reef Galleries 

of Tulear (Madagascar) and Their Sessile Invertebrate Communities. In A. M. Cameron et al., 

editors, Proceedings of the Second International Coral 

Reef Symposium, 2:143-159. Brisbane, Australia: 

Great Barrier Reef Committee. 

Wilkinson, C , and J. Vacelet 

1979. Transplantation of Marine Sponges to Different 

Conditions of Light and Current. Journal, Experimental Marine Biology and Ecology, 37:91-104. 
Surface Zooplankton at Carrie Bow Cay, Belize 

Joan D. Ferraris 

ABSTRACT 

Zooplankton at Carrie Bow Cay, Belize, was 

sampled by conventional plankton net tows in 

the open ocean immediately seaward of the reef 

and in the lagoon in order to obtain plankton 

both before and after it had passed over the reef. 

Tows were taken at the surface and in depths of 

3 m at each location, at approximately 0400 h, 

0800 h, 1600 h, and 2000 h. The composition of 

the samples indicates primarily oceanic plankton 

at both locations, and elements of a presumed 

resident reef fauna at the shallower lagoon station. Results demonstrate a net import of zooplankton to the reef and substantiate observations 

by other workers that plankton abundance increases at night. 

Introduction 

Investigations of zooplankton diversity, dynamics, and biomass provide essential information about breeding patterns (Moore and Sander, 

1976, 1977), behavioral strategies (Emery, 1968; 

Alldredge and King, 1977), emergence patterns 

(Glynn, 1973; Porter, 1974; Porter et al., 1977), 

predation (Johannes et al., 1970; Goreau et al., 

1971; Stevenson, 1972; Davis and Birdsong, 1973; 

Gerber and Marshall, 1974; Hobson, 1974; Porter, 1974), and the energetic impact of planktonic 

organisms on coral reef communities (Yonge, 

1930; Sargent and Austin, 1954; Odum and 

Odum, 1955; Quasim and Sankaranarayanan, 

1970; Tranter and George, 1972; Glynn, 1973; 

Johannes, 1974; Johannes and Gerber, 1974; reJoan D. Ferraris, Mount Desert Island Biological Laboratory, Salsbury Cove, Maine 04672. 

view by Lewis, 1977). The present report is concerned with zooplankton from the surface water 

adjacent to the coral reef at Carrie Bow Cay, 

Belize, and its possible import to that coral reef 

community. 

ACKNOWLEDGMENTS.—I thank B. W. Spracklin 

for invaluable aid in the field and R. J. Larson 

for assistance in the laboratory. 

Materials and Methods 

Plankton was sampled by towing a 29 cm 

diameter plankton net (100 jum mesh net, 250 /tm 

mesh basket) at a fixed distance behind a small 

boat during April and May 1976. In order to 

sample plankton both before and after it had 

passed over the barrier reef, the net was towed 

parallel to the reef at two locations. Site 1 

("ocean") is immediately seaward of the fore-reef 

slope. Water depth in this area is greater than 

200 m. Site 2 ("lagoon") is approximately 0.5 km 

shoreward of the reef at Carrie Bow Cay. Water 

depth at site 2 ranges from 5 to 8 m. The predominant wind direction at the time was ENE, so 

that trade winds were blowing from the open 

ocean in the direction of the lagoon. 

Horizontal tows at each site were taken consecutively at two depths, 0-0.5 and 2.5-3.0 m. Approximate times of tows were 0400 h (approximately 1 hour before sunrise), 0800 h, 1600 h and 

2000 h (approximately 1.5 hours after sunset) 

during the same 24-hour period, whenever 

weather conditions permitted. 

The net was equipped with a calibrated flow 

meter and all tows were timed (5-10 min) with a 

stopwatch, so that volume of water filtered could 

be calculated. Samples were fixed in 10% for143 
144 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

malin in sea water in the boat immediately after 

each tow. In addition, live samples were returned 

to Carrie Bow Cay for observation. 

The settled volume of the plankton in entire 

samples was determined by straining samples 

through 100 jum mesh plankton netting and rinsing them into a 25 ml graduated cylinder. Volume was recorded until constant readings were 

obtained (24 to 36 h). Large organisms such as 

Scyphozoa and Leptocephalus larvae (Pisces) as 

well as plant debris (for instance, Thalassia testudinum Banks ex Konig and Turbinaria sp.) were 

removed prior to volume determination. Any embryos suspended in mucus, however, could not be 

removed without interfering with other organisms 

in the sample. 

All samples were then subdivided using an 

eight-chambered plankton splitter. The volume 

of each sample was adjusted to 100 ml by the 

addition of 10% formalin in filtered sea water 

prior to splitting. One-eighth of each sample was 

removed and stained with 0.1% rose bengal to 

facilitate identification and enumeration of organisms. The highly abundant copepods and 

non-cirripede nauplii (mostly copepod) were 

counted by adjusting the one-eighth subsample 

to a volume of 20 ml and further subsampling it 

by removing 1 or 2 ml with a Hensen Stemple 

pipette. All counts were made under a Wild M 5 

stereomicroscope having a plankton counting 

wheel mounted on the base. The field of view at 

X 25 magnification spanned the width of the 

plankton wheel groove and permitted continuous 

counting while the wheel was rotated. 

The remaining seven-eighths subsample was 

strained through 100 /xm mesh netting, dried at 

60°C for 24 h and weighed to the nearest 1 mg 

after cooling in a desiccator. Samples were then 

ashed at 510°C and ash-free dry weights calculated. 

The Wilcoxon signed-ranks test (Sokal and 

Rohlf, 1969) was applied to means generated 

exclusively from paired groups of samples. 

Results 

Copepods and nauplii comprise much of the 

plankton in both lagoon (n = 40) and ocean (n 

= 32) samples (Table 13). In the lagoon, copepods 

account for 53.0% and nauplii (mostly copepod) 

account for 31.5% of the total number of individuals (combined value = 84.5%). Similarly, in 

ocean samples the mean percentage of copepods 

was 52.7 and of nauplii 29.7 (combined value = 

82.4%). At the taxonomic level at which the 

organisms were counted, more than 82% of the 

taxa were common to both lagoon and ocean 

samples. These values are comparable to those 

reported by Moore and Sander (1976) for the 

inshore zooplankton overlying two other Caribbean coral reef areas, where copepods comprised 

76.0% (Jamaica) and 84.4% (Barbados) of the 

plankton. 

Significant quantitative differences appear 

when samples are compared on the basis of the 

TABLE 13.—Relative abundance (percentage of individuals) 

of major plankton categories (> 0.5%) on either side of the 

barrier reef at Carrie Bow Cay (number of samples in 

parentheses) 

Copepoda 

Nauplii 

Gastropoda larvae 

Fish ova 

Larvacea 

Polychaeta larvae 

Shrimp* larvae 

Brachyura larvae 

Pteropoda 

Cirripeda larvae 

Chaetognatha 

Lagoon 

(40) 

53.0 

31.5 

3.0 

2.4 

2.2 

1.9 

1.1 

1.1 

0.9 

0.8 

0.7 

Ocean 

(32) 

52.7 

29.7 

2.0 

1.2 

8.3 

3.9 

0.3 

0.3 

1.0 

0.7 

0.5 

* Euphausiacea, Penaeidea, Caridea. 

TABLE 14.—Results of Wilcoxon signed-ranks test applied to 

abundance of organisms in different sample groups compared on basis of mean number of individuals/m3

 (data 

from Table 15; + = significant at P< 0.05; 0 = no significant 

difference) 

Sample groups compared 

Ocean : Lagoon 

Night : Day 

0-0.5m : 2.5-3.0 m depth 

Total 

plankton 

Copepods 

and 

nauplii Others 
NUMBER 12 145 

B 3 

3 J= 

S 

°7 II 

CM" 

g _ 

^ K 

B 

°7 II 

<? II 

IT) ^ 

C\i 

O to 

°7 II 

IT) N C_ 

csi 

CO 

— I 

r~ 

<£> 

+1 

co 

en 

co 

— i 

Ol 

L O 

+1 

tO 

CO 

^ 

01 

o 

+1 CM 

, i 

"*< 

— ' — i C M 

+1 +1 +1 

CM t O I O 

—> m 

+1 

o 

IO 

+1 

o 

CO 

+1 

-f 

+1 

CO 

CO 

+1 

to 

L.O 

8±6 

co 

9+8 o 

0±9 

• * 

• * 

r~» ~^ 

in —i — 

0+ o 8 

CM 0+6 

oS 

^ 

+i 

co 

o 

o 

+1 

o 

CO 

o 

+1 

r-~ 

LT) 

^-» 

+1 

<o 

o 

o 

C+~ 1 

ns 

a 

d !S S is fl p 2 s 

- • '' p.;- h fr a 

l O —i 

o 

o 

+1 

L O 

o 

+1 

> E- Z 2 $ 

fi o -C 

°7 II 

IT) ^ 

S 

°n II 

g ^ 

° 7 II 

irj ^ 

c\i 

g 

cs t ^ 

°7 II 

c\i 

CPi 

to 

CM 

Th 

+1 

ra> 

tO 

to 

CO 

+1 

o 

^ H 

to in 

C M — i 

+1 +1 

r-. o 

- ; to 

01 

o 

CO 

r -

+1 

o 

30.0 

0±1 

to 

o 

C O — 

^-> C O 

+1 +1 

CM i-i —i 

d +1 +1 

CO LO O 

C M - H 

— ' — i C M 

O l - H —, _ 

r-: +1 +1 +1 

CM CO CO o 

LO —I —i 

CO — 

Ol 

+1 

co 

to 

+1 

co 

LO 

+ 1 

o 

C M — I 

as J2 

-a 3 

o ci -a 

s 

O CO 

d d 

+1 +1 

co — 

C M — < 

— I C O 

E '> 

o .a R a 

> 

aj « C 

. 3 j -

•- o 

2 

2 O -C CL 3 <L> -3 

C n O U C fi -5-

< 
146 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

6 £ 

o 

o 

tan 11a 

at 

o 

(A 

s-

<u 

al 

* 

C 

uO 

O 

V 

3 

X* 

4_l 

c 

u 

<u 

n 

h + 

n 

h 

u 

TJ 

al 

T3 

C 

+1 eans 

E 

03 

3 

TI ivi 

TJ 

C 

t— 

O 

IH 

V 

i) 

B 

c 

-o 

c es, 

o 

tan 

V 

a 

u 

c 

o 

-Mi 

c 03 

OH 

1 

tf> 

Ul J 

CO 

< 

D. 

fc' (0 

o 

1) 

X 

fc 3 

C 

TJ 

C 

as 

C 

a o; TI 

U 

3 

O 

JO 

>^ 

XI 

-D 

OJ 

ts 

c 

V 

TJ 

U 

as 

rt 

X> 

oS 

X 

o 

£ 

V 

X 

fc 

tip _ 

as 

oS 

TJ 

<U 

,ti 

o 

a 

CS t<-) 

C M II 

CS \J -

°7 II 

IT) s=^ 

c\j 

B 

C S tr-j 

° 7 II 

CM 

CS to 

°7 ll 

IT) ^ 

C\3 

0+6 0+Z. —c o 

0+9 o 

o o o 

+1 +1 +1 

in r- o 

d o — 

+1 

+ «? + 

o 

as o 

r~ co 

d d 

+1 +1 

+ OS Ol 

d d 

+ + 

o o o 

+1 +1 +1 

CO OS O 

- o n 

d 

+1 

+ ~l 

+ + 

r- o co 

oo co co 6.8±0 0.7±0 0.7±0 

+ 

tO CM 

Tf OS CM 

•^ ca n-i 0.9±0 54.6±3 94.0±3 

+ + 

co r~ 

in — i 0.9±0 0.7±0 

CO 

CM 

+i 

CO 

CO 

+ 

7.2±4 

H 

co 

d 

+1 CO 

LO 

CO 

r~ 

iri 

+1 co 

to 1 ' 

+ 

o CO 

d 

i

+n 1 

d 

+ 

+ 

o 

c\i 

+1 

+ 1 CO 

r-~ 

CO 

CO 

+1 

co 

OS 

o CO 

d 

+1 in 

d 

m 

LO 

d 

+1 rd 

o 

co 

d 

+1 to 

LO 

o 

d 

+1 L O 

co 

+ 

+ 

CO 

Th 

d 

+1 r~-

d 

+ 

in 

L O 

OS 

+1 as 

5 250 

CM 

i — l 

+1 (O 

d 

CM 

Th 

d 

LO 

+1 

"*! 

8 133 

CO 

"f' 

L O 

+1 os 

• * ' 

Th 

CM 

rd 

+1 

p 

r» 

r~-

d 

+1 

— CM 

+ 

to 

d 

+1 co 

d 

<o IO 

d 

C+M 1 

+ 

CM 

CM 

CM 

+1 os 

c\i 

+ 

Oi 

CM 

d 

i

+n 1 

d 

+ 

to 

to 

d 

+1 to 

CM 

o to 

CM 

+1 to 

to 

+ 

+ 

"* in 

— 

+1 

—• • * 

CO 

co 

d 

C

+O 1 

c\i 

+ + + 

r-~ 

CM 

L+O 1 

CO 

r-« to 

o 

+1 LO 

• — ' 

CO 

CO 

»-( 

+1 CO 

CO + + 

CO 

OS 

— c 

• * +> 1 

Ol 

Th 

e-> 

LO 

L+O 1 

CM 

to 

to 

O 

+1 

o OJ + + 

-• — * ' 

-* 

+1 CO 

m 

co 

r~ 

O 

+1 

o 

— 

Th 

co 

CO 

+1 LO 

CM 

OS 

co 

o 

+1 

"-" CM 

Th 

Th 0+0 —' 

CM 2±0 

- • 

** 0+9 

+ 

o 

r»-0+9 m 

+ 

OS 

co 0±1 

CO 

os •* 

CO •* 

CM — 

+1 +1 

j- os co 

tO CM 

— "* CO 

r-' TH ^ 

CM CO O 

+1 +1 +1 

—i OS to 

CO CM O 

OS -* 

+ + + + 

• * 

Th 

+ 1 

o 

LO 

L O 

CO 

+1 

+ <? 

to 

r- O 

CM tO 

— o +1 +1 

r- r~-

CM O + 

Tjl 1-1 

CM CO CO 

co d °J 

CM co m 

+1 +1 +1 

Th (O CM 

to co oo 

+ + 

o 

+1 

r~~ 

o 

+1 Th 

o 

+1 

oi + + + 

Th 

en • — i 

+1 

— CO 

rCM « 

o 

+1 CO 

o 

Th 

co 

o 

+1 CO 

CO + 

Th 

CO 

o 

+1 

m 

o 

CM 

r^ 

_; Th 

C+M 1 

CO 

CM 

. — i 

to 

CM 

+1 in 109 

Th 

m 

O 

+1 

—' 

"~' 

+ + 

+1 +1 

— T"**h! o "1 + 

to 

CM 

+1 to 

os 

OJ 

CO 

CS 

+1 

ro 

Th 

I--

+1 os 

CM 

Th 

in 

f s 

+1 

o + 

« ^ ^o 

O—i 0 CO0 O *>. 

+1 +1 +1 

m in to 

O CM o 

CM CM 

CO —l 

+ + + + 

-: + 

+ + 

Os 

CM 0+6 o 

to 0+6 o 

en 

to 0+8 o 

+ + 

- H as t o 

Th O co 

to ^ d 

+1 +1 +1 

CM cq r^- _|_ 

co CM — 

^ r~ m 

CM d d 

+1 +1 +1 

r- -H TJH o 

+1 +1 +1 +1 

co o — to 

^ t\i N O 

+ + + 

m 

O 7±1 

Th 

CM 8±1 

— 

CM 

to 4±4 

r-» 

o 

CO 5±0 

o 

CM - H CM — 

Q Th CO CM 

- d co -

+1 +1 +1 +1 

Th o o —' Th 

co — os — 

m co o 

o to CM 

CM d co 

+1 +1 +1 

_|_ CO —I Tf to CM 

O CO CO 

—i CO OS 

— d o 

+1 +1 +1 

— to OS 

iri CM CM 

+ 

tD 

r^ 

O 

+1 CM 

^ H 

(TM 

rH 

+1 OS 

to 

to 

_ 

+1 to 

^ 2 

?i 

u 

O ei 

O -C , 

o 2 > 

« b H > oS 

^ u y y T5 oS as 3 oj 

JS XI " p 

3 

O o T3 

o. a o 

2 ^ 2-

fi o o 

T3 

8,3 y 

._, T3 

3 ' 

TJ 

O oj 

CX W 

2 y 

* - > a s 

a b "J —' 

T3 J2 

a l l 

as 

S u aiijS-'o' o e-^^'C^-2 / g oJ.bo^ B g_^ 

3 T3 

>- o 

-c c 

OS 1 5 

u O 

y 2 

—i <u 

X 

03 

c b/s as 

O ' 

^ 

o J^ a;2 

: o 

w Q 

•JS y 2 ? 3 

o co IT! ° —' 

PH X 
NUMBER 12 147 

O 

cean 

O 

-«; 

CS 

CS 

TjCS 

-c 

CS 

cs 

cs CM 

-*Ca : 

Cs 

to 

-c Cs 

Cs 

CO 

Cs 

S f Plankt 

£ 

C S CTj 

c? II 

CNi 

^ Sj, 

Cs ^ 

s 

Csco 

°7 II 

^C\i e 

c"s I? I 

1 JS 

Cs —' 

s Cs >. 

°7 II >A>S. c\i 

S II 

Cs ^ 

B ^ 

Cs <o 

C7 II 

ir, J£ 

CM -0.5 m 

(n = 6) 

^ 

s 

V) 

I 

r i 

CM 

d 

+1 CO 

—• 

CO 

— 

+1 in 

OJ 

OJ 

CO 

d 

+1 to 

d 

in 

0 

d 

+1 r^ 

d 

Th 

CM 

O 

+1 

r» 

—' 

L O 

L O 

d 

+1 to 

0 

0 

oi 

r+^ 1 

CO 

Th 

Th 

CO 

+1 to 

L O 

y Cnidaria larva 

+ 

r^ 

r-

— 

+ 1 

co 

CM 

+ 

+ 

OJ 

d 

C+O 1 

d 

+ 

+ Medusae 

r^ 

to 

d 

+1 

0 

in 

.—1 

+1 CM -|- r~-

• — 1 

Ol 

p» 

d 

T+h 1 

04 

OS 

d 

+1 OS 

-^ 

L O 

O 

d 

+1 

p 

—• 

r~ 

CO 

d 

+1 

— 

Th 

CO 

d 

C+O 1 

L O 

CO 

d 

+1 co 

d 

+ 

L O 

LO 

d 

+1 

r~ _i_ r~ 

~^ 

fl>» 

d 

+1 to 

oi 

L O 

L O 

d 

+1 Th 

c\i 

O 

OS 

d 

+1 CM 

CM 

0 

LO 

to 

0 

C+O 1 

os CM 

CM 

+1 00 

CO 

0 

CO 

0 

+1 CO 

X 

va 

V 

Siphonophora 

Ctenophora 

Platyhelminth 

larvae 

r~-

CO 

Th 

+1 

r~ 

os 

to 

Th 

CM 

T+h i 

r-* 

co 

O 

CM 

+1 co 

r^ 

O 

O 

X 

— 

+1 

r-» 

co 

L O 

cs 

r~ 

+1 01 

r^ 

OJ 

co 

X 

OS 

+1 

r» 

oi 

CO 

00 

CO 

CM 

O+S 1 

iri 

CM 

co 

L O 

d 

C+M i 

t~ 

CM vae Polychaeta lar 

+ 

+ 

CO 

Th 

d 

+1 

x 

d 

+ 

+ 

+ 

+ ults Polychaeta ad 

+ 

+ 

co L O 

d 

+1 

— 

f» 

os 

X 

+1 O 

oi 

to 

Th 

d 

+1 OS 

d rvae Sipunculida la 

+ 

+ 

0 

r~~ 

d 

+1 

rd 

+ 

+ 

r~-

co 

d 

+1 

p 

X Bryozoa larvae 

LO 

Th 

oi 

+1 os 

iri 

co 

•n 

— < +1 co 

CO 

CO 

m 

CO 

+1 CM 

Th 

0 

r~ 

OS 

C+O 1 

co 

Ol 

CO 

0 

co 

+1 

X 

d 

OS 

to 

+1 

os 

Th 

LO 

L O 

oi 

r+^ 1 

OS 

Th 

CM 

oi 

+1 

— 

oi rvae Gastropoda la; 

_ 

n 

d 

+1 

p - 1 

m 

d 

+1 

p 

m 0 

d 

+1 OJ 

— 

+ 

X 

X 

d 

+1 0 

oi 

O 

L O 

d 

O+S 1 

t— 

os 

d 

—+• 1 

oi 

0 

CM 

d 

+1 in 

d •vae Pelecypoda lai 

CO 

0 

oi 

+1 X 

CO 

m 

X 

oS 

+1 co 

os 

0 

X 

d 

+1 

co 

oi 

L O 

—.—; 1 

i+n 1 

ri 

0 

Th 

r~ 

+1 O 

CM 

OS 

X 

CM 

0+4 1 

OS 

OS 

CO 

X 

O+J 1 

oi 

X 

—; r^ 

+1 CO 

co Pteropoda 

m 

n 

1—1 

+1 to 

— 

CO 

I—H 

fl 

to 

co CO 

d 

+1 

m 

d 

co 

L O 

d 

+1 

p 

— 

L O 

L O 

d 

T+h 1 

OJ 

CO 

X 

+1 CO 

oi 

— iri 

+1 OS 

ri 

CO 

if) 

1^ 

+1 OJ 

co 

Os 

X 

X 

+1 

r^ 

r^ 

Th 

d 

+1 CO 

d 

+ 

+ 

+ 

r-» 

CO 

d 

+1 m 

d 

m x 12.1 40.7 

.62 

CM — O 

+1 +1 +1 

CM r- os 

•^ d — 

Tf — 

CO CM 

OS 

O CO 

^ OS CM 

— d "I — Th CM 

+1 +1 +1 

cq CM in 

d os iri 

CO — 

CM — 

CM CO 

CO r~ 

co r* 

OS Th r^ m 

+1 -H C O Tf< -) -

X OS 

c—o C OJ O 

•n 0 

m Th 

co" ri 2 

os os P 

r» co co 

+1 +1 +1 

O CO O 

r^ CM co 

O r» 

co in 

r-- in 0 

p Th in 

C M iri iri 

Oa Th — 

+1 +1 +1 

CM CM m 

N ri us 

CTMh C CM O — 

in 

— CO d P £ x co ^ 

— Oa m 

+1 +1 +1 

co co os 

oi — d 

os — 

CO CO 

CM 

Oa to ri «? 

CM r-~ 

— Th 

+1 +1 , 

m 0 + 

CO °\ 

r-~ to 

CM — 

— 

CM X 

CM CO Th 

CO CO ^ 

Tf — O 

+1 +1 +1 

00 X 00 

oi co d 

0 —1 

CM — 

V 

Cladocera Ostracoda 

Copepoda 

Nauplii Cirripeda larv 

+ 

in 

0 

d 

r-

+1 s 

d 

+ 

co 

Th 

d 

+1 m 

d larvae Stomatopoda 

in 

m 

d 

+1 X 

d Mysidacea 

r^ 

r^ 

d 

+1 

co 

d 

+ 

+ 

L O 

CO 

d 

+1 

°°. + 0 

+ 

+ + 

+ 

Cumacea Amphipoda 

_ 

—1 

X 

+1 CO 

X 

co Ol 

X 

+1 in 

oi 

01 

rd 

+1 Th 

rt 

in 

Th 

X 

C+O 1 

CO 

Ol 

+ 

co 

OJ 

d 

+1 X 

d 

+ 

OJ 

OJ 

d 

+1 in 

d 

V Shrimp* larva 

0 

Ol 

d 

+1 co 

d 

X 

m 

d 

+1 X 

d 

+ 

+ 

pi Anomura larv 

,_, 

OJ 

d 

+1 Th 

X 

as OS 

d 

+1 OS 

Th 

X 

d 

+1 OJ 

—' 

0 

00 

Th 

O+J i 

oi 

OJ 

r-» 

r-« 

d 

+1 

p 

—

c 

CO 

CO 

d 

+1 

X 

d 

OJ 

Oa 

d 

C+O 1 

X 

+ 

oS 

s> Brachyura lar 

CO 

d 

+1 CO 

X 

CO 

CO 

d 

+1 CO 

X 

CO 

d 

+1 X 

d 

m 

0 

d 

+1 r^ 

d 

CO 

CM 

d 

+1 X 

d 

+ 

CO 

CO 

d 

+1 CO 

d 

+ 

a Echinodermat 

larvae 

OJ 

X 

d 

+1 -1- 00 

0 

+ + 

CO 

X 

+1 

—' 

co 

Th 

0 

+1 

"1 + 0 

0 

OJ 

+1 OJ 

+ + 

V 

. larva 

arvae 

Hemichordata 

Urochordata 1 

Oa 

Th 0 x CM x — 

to x y-; CM m CM 

d d co — — d 

+1 +1 +1 +1 +1 +1 

to i~~ •>• x — r^ 

d d d co co d 

m 

00 

00 — in 0 in 

co QJ t>- <x> CM 

0 CM 0 r^ d 

+1 +1 +1 +1 +1 

-j. in C M in 00 in 

d os CM os d 

CO 

Th 

— in x 

CO ^ P 

-H (N -

+1 +1 +1 

-|- _|- Th r-; CM 

CO t-^ CO 

CM 

0 

0 in C M in 

Th co d co 

0 (o — d 

+1 +1 +1 +1 

+ + r-- r-. p —; 

r^ os co — 

— OJ 

Th 

CO 00 CM 

^ X X 

CM —< — 

+1 +1 +1 

-(--(- — CO CO 

r^ co CM 

Tf 

Tf — 

CO — O OS 

* CO 1 -

O CO — CO 

+1 +1 +1 +1 

X -)- Tf CO X -}-

d 06 Th oi 

Th CO 

x os Th — 

< ^ ri <=? <=>. O CM — — 

+1 , +1 +1 +1 , 

x + CM in p + 

d x CM oi in 

X 

co Th 0 os 

"* Th ^ ^ 

O CM O — 

+1 +1 +1 +1 

m CM oa co -|-

d d —< Tf 

m 

Salpa Doliolida Larvacea 

Chaetognatha 

Fish ova Fish larvae 
148 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

mean abundance of organisms in different habitats and light periods (Table 14). In order to 

eliminate the influence of the dominant copepods 

and nauplii, these groups were tested as a separate 

category. Abundance of organisms in ocean samples is greater than in lagoon samples with respect 

to all three counts. Night tows (0400 h and 2000 

h) yield a larger combined number of individuals 

than daytime samples (0800 h and 1600 h), as 

would be expected from the bimodal pattern of 

plankton emergence (near dawn and dusk) indicated by Glynn (1973). This difference does not 

hold, however, for organisms other than copepods 

and nauplii which show no significant quantitative difference between night and day. On the 

other hand, surface tows (0-0.5 m) differ significantly from 2.5-3.0 m tows only with respect to 

plankton other than copepods and nauplii. 

These differences are not entirely reflected by 

settled volume or ash-free dry weights (Table 15). 

When mean values for settled volume generated 

by paired groups of samples are compared by the 

Wilcoxon signed-ranks test, significantly more 

plankton volume/m3 is found in ocean samples 

than lagoon samples and, contrary to individual 

numbers, in day tows than in night tows {P < 

0.05). The latter result is probably due to the 

presence of significantly more siphonophores in 

day samples (see below). Depth of tow did not 

have bearing on volume of plankton. Ash-free dry 

weight, which is a more reliable estimate of biomass than settled volume, does not differ significantly when locations or times of day are compared (Table 17) using means generated by 

paired groups of samples (Table 15). Significantly 

more plankton biomass (mg/m3), however, is 

found at 0-0.5 m than in 2.5-3.0 m, regardless of 

location or time of day. The lack of agreement 

between these measurements of numbers of individuals and biomass probably reflects differences 

in the composition of the samples (Tables 13, 16) 

as well as patchiness or unevenness in zooplankton distribution. Patchiness is indicated by the 

large standard errors of means calculated for all 

counts and biomass measurements in the present 

study (Tables 15, 16). It should be noted, however, that mean ash-free dry weight (mg/m ) of 

ocean samples is greater than that of lagoon 

samples, that of night samples exceeds that of day 

samples, and that of 0-0.5 m samples is greater 

than that of 2.5-3.0 m samples (Table 17). 

The dominant organisms other than copepods 

and nauplii represented at both locations indicate 

a primarily oceanic plankton: larval polychaetes, 

gastropods, cirripedes, and shrimps (Euphausiacea, Penaeidea, Caridea) as well as pteropods, 

larvaceans, chaetognaths, and fish ova (Tables 

13, 16). Of these groups, polychaete larvae, gastropod larvae, larvaceans, and chaetognaths are 

significantly more numerous in ocean than in 

lagoon samples {P < 0.05). Similarly, larval cnidarians, platyhelminthes, bryozoans, pelecypods, 

and echinoderms, as well as siphonophores, ostracods, copepods, nauplii, salps, and doliolids are 

more numerous in ocean samples. Also, although 

the blue-green alga Oscillatoria {= Trichodesmium) 

sp. was not counted, it was observed to be a major 

component of the ocean but not the lagoon plankton. These results may indicate a net import of 

organisms to the reef because oceanic plankton is 

removed from the water column by the reef community. Similar findings are reported by Tranter 

and George (1972) at Kavaratti and Kalpeni 

atolls in the Laccadives, by Glynn (1973) at 

Puerto Rico, and by Johannes and Gerber (1974) 

at Eniwetok Atoll. In contrast, Motoda (1940) 

and Johnson (1949, 1954) found zooplankton to 

be less abundant in ocean waters than in lagoon 

waters. However, as pointed out by several auTABLE 17.—Carrie Bow Cay plankton biomass, expressed as 

ash-free dry weights (mg/m3 ± standard error) for the six 

major sample groups, with results of Wilcoxon signed-ranks 

test for paired groups (+ = significant at P < 0.05; 0 = no 

significant difference) 

Sample 

group 

Ocean 

Lagoon 

Night 

Day 

0-0.5 m 

2.5-3.0 m 

mg/m3 

2.35±0.41 

1.85±0.30 

2.48±0.41 

1.71±0.25 

2.44±0.41 

1.75±0.28 

Wilcoxon test 

0 

0 

+ 
NUMBER 12 149 

thors (Alldredge and King, 1977; Porter and 

Porter, 1977; Porter et al., 1978; Riitzler et al., 

1980), plankton samples from the surface waters 

associated with coral reefs do not necessarily reflect the amount of plankton actually available 

to reef organisms. In the present study, certain 

groups of organisms (mysids, cumaceans, brachyuran larvae) are present in significantly {P < 

0.05) greater numbers per m3 in the lagoon. Also, 

the percentage of the plankton represented by 

these groups, together with adult polychaetes and 

shrimp larvae, is greater in the lagoon than outside the reef (based on data in Table 13, with the 

addition of lagoon versus ocean sample percentages for adult polychaetes, 0.2:0.04; mysids, 0.08: 

0.004; and cumaceans, 0.4:0.005). Similarly, the 

foraminiferan Rosalina (= Tretomphalus) sp. was 

recorded as abundant in lagoon but not in ocean 

samples. These results may indicate a significant 

difference in the composition of the plankton 

between the two localities. Hence, this difference 

may also reflect the presence of a resident reef 

fauna in the lagoon, represented by organisms 

that are at times considered demersal, such as 

adult polychaetes, mysids, cumaceans, shrimp 

larvae, and brachyuran larvae (Tranter and 

George, 1972; Sale et al., 1976; Alldredge and 

King, 1977; Porter and Porter, 1977). Such a 

resident reef fauna would be sampled only partially by the methods used in the present study 

and more likely would appear in samples from 

the lagoon, where water depth is only 5-8 m and 

small patch reefs are common. Similarly, differences in composition but not necessarily in biomass of lagoon and offshore plankton were found 

by Johnson (1954), Odum and Odum (1955), 

Bakus (1964), Tranter and George (1972), and 

Sale et al. (1976). This difference would indicate 

removal of oceanic plankton by reef organisms 

and replacement by a resident lagoon plankton 

(Emery, 1968). 

Tows taken at 0-0.5 m and 2.5-3.0 m appear 

to have similar composition. Mysids, cumaceans, 

and fish ova were more abundant {P < 0.05) at 

0-0.5 m whereas pelecypod and anomuran larvae 

were more common in 2.5-3.0 m tows. 

Comparing day and night tows, significantly 

more cnidarian larvae, siphonophores, platyhelminth larvae, sipunculid larvae and cladocerans 

appear in daytime (0800 h and 1600 h) tows. 

Crustacean groups (ostracods, mysids, cumaceans, amphipods, shrimp larvae, and anomuran 

larvae) as well as doliolids, however, are more 

abundant at night (0400 h and 2000 h). Results 

of this study agree, in general, with Emery (1968), 

Johannes et al. (1970), Glynn (1973), Porter 

(1974), and Renon (1977), who have noted both 

quantitative and qualitative differences between 

day and night samples of zooplankton over a 

variety of coral reefs. More specifically, Alldredge 

and King (1977) also report more ostracods, mysids, cumaceans, amphipods, shrimps, and decapod larvae in night tows than in daylight tows at 

Lizard Island on the Great Barrier Reef. 

Conclusions 

Copepods and nauplii (mostly copepod) combined account for 84.5% and 82.4% of the zooplankton in lagoon and ocean samples, respectively, from Carrie Bow Cay. Other major groups 

of organisms represented at both locations (larval 

polychaetes, gastropods, cirripedes, and shrimp, 

as well as pteropods, larvaceans, chaetognaths, 

and fish ova) indicate a primarily oceanic plankton. 

The relative abundance of mysids, cumaceans, 

adult polychaetes, and brachyuran and shrimp 

larvae in the lagoon may indicate partial sampling of a resident reef fauna in this habitat. 

Samples from the open ocean contain a significantly greater number of organisms (copepods 

and nauplii, as well as all others) per m water 

than those from the lagoon, and therefore suggest 

a net import of zooplankton to the reef. Plankton 

biomass, expressed by ash-free dry weight was 

greater in ocean, night, and 0-0.5 m samples than 

in lagoon, day, and 2.5-3.0 m tows. 

Resident reef fauna will have to be sampled 

directly before final conclusions can be drawn 

about the impact of zooplankton on the energy 

budget of the coral reef community. 
150 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Literature Cited 

Alldredge, A. L., and J. M. King 

1977. Distribution, Abundance, Substrate Preferences of 

Demersal Reef Zooplankton at Lizard Island Lagoon, Great Barrier Reef. Marine Biology, 41:317-

333. 

Bakus, G. J. 

1964. The Effects of Fish Grazing on Invertebrate Evolution in Shallow Tropical Waters. Allan Hancock 

Foundation Publications Occasional Paper, 27:1-29. 

Davis, W. P., and R. S. Birdsong 

1973. Coral Reef Fishes Which Forage in the Water 

Column. Helgb'lander Wissenschaftliche Meeresuntersuchungen, 24:292-306. 

Emery, A. R. 

1968. Preliminary Observations on Coral Reef Plankton. 

Limnology and Oceanography, 13:293-303. 

Gerber, R., and N. Marshall 

1974. Reef Pseudoplankton in Lagoon Trophic Systems. 

In A. M. Cameron, B. M. Campbell, A. B. Cribb, 

R. Endean, J. S. Jell, O. A. Jones, P. Mather, and 

F. H. Talbot, editors, Proceedings of the Second International Coral Reef Symposium, 1:105-107. Brisbane, 

Australia: The Great Barrier Reef Committee. 

Glynn, P. W. 

1973. Ecology of a Caribbean Coral Reef. The Porites 

Reef-Flat Biotope, Part II: Plankton Community 

with Evidence for Depletion. Marine Biology, 22:1-

21. 

Goreau, T. F., N. I. Goreau, and C. M. Yonge 

1971. Reef Corals: Autotrophs or Heterotrophs? Biological Bulletin, 141:247-260. 

Hobson, E. S. 

1974. Feeding Relationships of Teleostean Fishes on 

Coral Reefs in Kona, Hawaii. Fishery Bulletin, 72: 

915-1031. 

Johannes, R. E. 

1974. Sources of Nutritional Energy for Corals. In A. M. 

Cameron, B. M. Campbell, A. B. Cribb, R. Endean, J. S. Jell, O. A. Jones, P. Mather, and F. H. 

Talbot, editors, Proceedings of the Second International 

Coral Reef Symposium, 1:133-137. Brisbane, Australia: The Great Barrier Reef Committee. 

Johannes, R. E., S. L. Coles, and N. T. Kuenzel 

1970. The Role of Zooplankton in the Nutrition of Some 

Scleractinian Corals. Limnology and Oceanography, 

15:579-586. 

Johannes, R. E., and R. Gerber 

1974. Import and Export of Net Plankton by an Eniwetok Coral Reef Community. In A. M. Cameron, 

B. M. Campbell, A. B. Cribb, R. Endean, J. S. 

Jell, O. A. Jones, P. Mather, and F. H. Talbot, 

editors, Proceedings of the Second International Coral 

Reef Symposium, 1:97-104. Brisbane, Australia: The 

Great Barrier Reef Committee. 

Johnson, M. W. 

1949. Zooplankton as an Index of Water Exchange between Bikini Lagoon and the Open Sea. Transactions of the American Geophysical Union, 30:238-244. 

1954. Plankton of Northern Marshall Islands, Bikini and 

Nearby Atolls. United States Geological Survey Professional Papers, 260-F:301-304. 

Lewis, J. B. 

1977. Processes of Organic Production on Coral Reefs. 

Biological Reviews, 52:305-347. 

Moore, E., and F. Sander 

1976. Quantitative and Qualitative Aspects of the Zooplankton and Breeding Patterns of Copepods at 

Two Caribbean Coral Reef Stations. Estuarine and 

Coastal Marine Science, 4:589-607. 

1977. A Study of the Offshore Zooplankton of the Tropical Western Atlantic Near Barbados. Ophelia, 16: 

77-96. 

Motoda, S. 

1940. Comparison of the Conditions of Water in Bay, 

Lagoon and Open Sea in Palao. Palao Tropical 

Biological Station Studies, 2:41-48. 

Odum, H. T., and E. P. Odum 

1955. Trophic Structure and Productivity of a Windward Coral Reef Community on Eniwetok Atoll. 

Ecological Monographs, 25:291-320. 

Porter, J. W. 

1974. Zooplankton Feeding by the Caribbean Reefbuilding Coral Montastrea cavernosa. In A. M. Cameron, B. M. Campbell, A. B. Cribb, R. Endean, J. 

S. Jell, O A. Jones, P. Mather and F. H. Talbot, 

editors, Proceedings of the Second International Coral 

Reef Symposium, 1:111-126. Brisbane, Australia: 

The Great Barrier Reef Committee. 

Porter, J. W., and K. G. Porter 

1977. Quantitative Sampling of Demersal Plankton Migrating from Different Coral Reef Substrates. Limnology and Oceanography, 22:553-556. 

Porter, J. W., K. G. Porter, and Z. Batac-Catalan 

1977. Quantitative Sampling of Indo-Pacific Demersal 

Reef Plankton. In D. L. Taylor, editor, Proceedings 

of the Third International Coral Reef Symposium, 1:105-

112. Miami, Florida: Rosenstiel School of Marine 

and Atmospheric Science. 

Porter, K. G , J. W. Porter, and S. L. Ohlhorst 

1978. Resident Reef Plankton. In D. R. Stoddart and R. 

E. Johannes, editors, Coral Reefs: Research Methods, 

Monographs on Oceanographic Methodology, 5:499-514. 

Paris: Unesco. 
NUMBER 12 151 

Qasim, S. Z., and V. N. Sankaranarayanan 

1970. Production of Particulate Organic Matter by the 

Reef on Kavaratti Atoll (Laccadives). Limnology 

and Oceanography, 15:574-578. 

Renon, J. P. 

1977. Zooplankton du lagon de I'atoll de Takapoto (Polynesie Francaise). Annates de I'Instilul de la Oceanographie (Paris), 53:217-236. 

Riitzler, K., J. D. Ferraris, and R. J. Larson 

1980. A New Plankton Sampler for Coral Reefs. Marine 

Ecology, 1:65-71. 

Sale, P. F., P. S. McWilliam, and D. T. Anderson 

1976. Composition of the Near-Reef Zooplankton at 

Heron Reef, Great Barrier Reef. Marine Biology, 

34:59-66. 

Sargent, M. C , and T. S. Austin 

1954. Biologic Economy of Coral Reefs. United States 

Geological Survey Professional Papers, 260-E:293-300. 

Sokal, R. R., and F. J. Rohlf 

1969. Biometry. 776 pages. San Francisco: W. H. Freeman 

and Co. 

Stevenson, R. A., Jr. 

1972. Regulation of Feeding Behavior of the Bicolor 

Damsel-Fish (Eupomacentrus parlitus Poey) by Environmental Factors. In H. E. Winn and B. L. 

Olla, editors, Behavior of Marine Animals, 2:278-302. 

New York: Plenum Press. 

Tranter, D. J., and J. George 

1972. Zooplankton Abundance at Kavaratti and Kalpeni Atolls in the Laccadives. In C. Mukundan 

and C. S. Gopinadha Pillai, editors, Proceedings of 

the Symposium on Corals and Coral Reefs, pages 239-

256. Cochin: Marine Biological Association of 

India. 

Yonge, C. M. 

1930. Studies of the Physiology of Corals, I: Feeding 

Mechanisms and Food. Scientific Reports of the Great 

Barrier Reef Expedition, 1:13-57. 
Plankton Diatoms (Bacillariophyceae) 

from Carrie Bow Cay, Belize 

Paul E. Hargraves 

ABSTRACT 

Seventy-three diatom taxa from the coastal 

waters around Carrie Bow Cay, Belize (Central 

America) were found in samples collected April/ 

May 1977 and January 1978. Chaetoceros, Rhizosolenia, and Coscinodiscus were the most common 

genera. Six species are new records for the Caribbean region. Distinct differences exist between 

spring and winter diatom flora. Although numerous spore-forming species were found, resting 

spores were absent. Cultivation of more tropical 

diatoms is seen as an aid in solving problems in 

the systematics of tropical phytoplankton. 

Introduction 

The species composition of plankton diatoms 

along the Carribean coast, unlike that of coastal 

areas in temperate regions, is poorly known. For 

oceanic areas of the Caribbean more information 

is available because a considerable number of 

oceanographic cruises have included sampling for 

phytoplankton (for instance, Hargraves et al., 

1970; Hulbert, 1968; Marshall, 1973; Takano, 

1960). Coastal floristic works are few and widely 

scattered geographically (Buchanan, 1971; Margalef, 1957, 1968; Sander, 1976). Apparently no 

publications deal with coastal plankton diatoms 

of Belize, although planktonic coccolithophorids 

have been examined (Kling, 1975). 

In terms of turnover rate and primary production in tropical waters, the nanoplankton (larger 

Paul E. Hargraves, Graduate School of Oceanography, University of 

Rhode Island, Kingston, R. I. 02881. 

than 1-10 jiim but smaller than 50-70 jum, depending on authors' definition) is often more 

important than the net plankton (Malone, 1971; 

Texeira, 1963); nanoplankton may occasionally 

dominate the biomass as well (Garrison, 1975). 

Owing to my limited sampling, the annotated list 

below does not completely represent plankton 

diatoms of Carrie Bow Cay. Also, primarily 

benthic species are excluded from this report, 

except for the few whose consistent presence indicates a pelagic as well as benthic existence. 

Generally, diatom species have been defined on 

the basis of morphological characteristics without 

a proven genetic basis. Variability within a species is often considerable, not only morphologically (for example, Van Landingham, 1967, lists 

over 100 synonyms for Actinocyclus ehrenbergii that 

were formerly considered species, differentiated 

on morphological bases), but physiologically as 

well (for example, Carpenter and Guillard, 1971; 

Hargraves and Guillard, 1974). Although impractical for routine purposes, genetic analysis by 

means of enzyme electrophoresis (Gallagher, 

1979; Murphy and Guillard, 1976) shows considerable promise for separating diatom species. 

Nevertheless, morphological features are the main 

criteria for current classification schemes. 

The grouping of species into genera, of genera 

into families, and of families into orders is in a 

state of flux, primarily because of recent discoveries about fine structure of the siliceous diatom 

frustule (see reviews in Werner, 1977). Organization of these hierarchies is subjective, depending on individual assessment of the relative importance of given structures, and diatomists have 

153 
154 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

probably not yet attained a wholly natural classification system. For this reason, the list below is 

arranged in alphabetical order rather than in a 

particular phylogenetic scheme. Some recent attempts at an incisive phylogenetic arrangement 

have been reviewed by Hendey (1974) and Simonsen (1974). 

ACKNOWLEDGMENTS.—The assistance of P. 

Boyd, D. Hargraves, and K. Zimmerly is gratefully acknowledged. Partial support came from 

the National Science Foundation, Grant No. 

OCE 76-82280. 

Methods 

Samples were collected with 30 cm diameter 

plankton nets, having mesh sizes of 65 jum or 10 

ju,m and were preserved in hexamine-buffered 

formalin or Lugol's iodine, without acetic acid. 

Aliquots were washed with distilled water to remove salts. To remove organic matter, cells were 

oxidized by either boiling for one hour in 30% 

hydrogen peroxide, or by ashing samples for three 

hours on cover slips on a hot plate. Permanent 

slides of the oxidized material were prepared 

using Hyrax (Custom Research & Development, 

Inc., Auburn, California) as a mounting medium. 

These permanent mounts and the liquid-preserved material were examined in a Zeiss Photomicroscope II using brightfield, phase contrast, 

and Nomarski interference contrast optics. When 

necessary, identifications were confirmed using a 

Cambridge Stereoscan electron microscope. 

Slides are maintained in the author's collection. 

Collection sites were in the vicinity of Carrie 

Bow Cay, in the central portion of the Belize 

barrier reef. Surface tows were made over Thalassia, near the mangrove forests of Twin Cays, in 

the channel between Carrie Bow Cay and South 

Water Cay, in deeper water (over 200 m) about 

0.5 km east of Carrie Bow Cay, and in the "Southern Shelf of the lagoon, at approximately 

16°55'N, 88°12'W in over 20 m depth. The samples were collected in April and May 1977 and 

January 1978 at the same localities, and thus they 

cover the late spring and mid-winter periods. 

Annotated List of Diatoms 

(Species new to the Caribbean flora are designated by *) 

Actinocyclus ehrenbergii Ralfs in Pritchard (Figure 

87a) 

[= A. octonarius Ehrenberg] 

Found only in lagoon samples in spring; never 

common. Previously reported from the Caribbean by Hagelstein (1938) and Margalef 

(1968). Characteristic of both benthic and 

pelagic environments. Reference: Hendey 

(1964). 

Actinocyclus subtilis (Gregory) Ralfs in Pritchard 

Found only in lagoon samples in spring; never 

common. Previously reported from the Caribbean by Hagelstein (1938) and Margalef 

(1968). Characteristic of both benthic and 

pelagic environments. Reference: Hendey 

(1964). 

Actinoptychus senarius (Ehrenberg) Ehrenberg 

[= A. undulatus (Bailey) Ralfs in Pritchard] 

Found only rarely in samples from the lagoon 

side of Carrie Bow Cay in winter. A cosmopolitan species, it is widespread in temperate 

coastal waters in all seasons, but apparently 

rarer in the tropics. Reported from the Caribbean by Hagelstein (1938) and Margalef 

(1968) among others. Reference: Hendey 

(1964). 

Asterionella notata (Grunow) Grunow in Van 

Heurck (Figure 8bd) 

Uncommon to common in all samples; more 

abundant on the ocean side of Carrie Bow 

Cay. A tropical coastal species straying rarely 

into temperate waters. Widespread in the Caribbean, but apparently misidentified occasionally as A. kariana, an Arctic species. Reference: Hustedt (1931-1959). 

* Asteromphalus hookeri Ehrenberg (Figure 85c) 

Present rarely and only in spring from the 

ocean side. Normally oceanic in cooler waters. 

Not previously reported from the Caribbean. 

Reference: Lebour (1930). 

Bacteriastrum comosum Pavillard 

Occasionally found in winter only in lagoon 
NUMBER 12 155 

and ocean-side samples. A predominantly 

tropical species, previously reported from the 

Caribbean by Margalef (1968). Reference: 

Hustedt (1927-1930). 

Bacteriastrum delicatulum Cleve 

Found in lagoon and ocean-side samples in 

spring and winter. An oceanic species, reaching its maximum abundance in temperate 

waters. Widespread and occasionally abundant throughout the Caribbean. Reference: 

Hendey (1964). 

Bacteriastrum elongatum Cleve (Figure 86b) 

Found rarely in ocean-side tows and only in 

spring. Commonly found in tropical waters, 

and widely reported from the Caribbean. Reference: Hendey (1964). 

Bacteriastrum cf. furcatum Shadbolt (Figure 86a) 

This cell was seen in one sample from the 

ocean side in spring. It resembles B. furcatum 

Shadbolt, sensu Boalch (1975), but insufficient material was found to make an identification. 

Bacteriastrum hyalinum Lauder (Figure 85a) 

Occasionally found in winter samples, from 

both lagoon and ocean side. Common in temperate and tropical coastal waters. Previously 

reported from the Caribbean by Hargraves et 

al. (1970) and Margalef (1968). Reference: 

Hendey (1964). 

Bacteriastrum varians Lauder 

Found once in a spring sample from the lagoon side of Carrie Bow Cay. Generally considered a tropical oceanic species, it was reported in the Caribbean by Takano (1960). 

Boalch (1975) considers B. varians a synonym 

of B. furcatum Shadbolt. Reference: Boalch 

(1975). 

Cerataulina pelagica (Cleve) Hendey 

[= Cerataulina bergonii (Peragallo) Schutt] 

Present in small numbers in winter samples. 

Widely distributed in coastal temperate and 

tropical regions. Found throughout the Caribbean region. Reference: Hendey (1964). 

Chaetoceros affine Lauder 

Occasionally present in ocean-side samples in 

winter and spring. Widely distributed in temperate and tropical coastal waters. Commonly 

reported from the Caribbean. The morphology of this species is highly variable. Some 

chains appear transitional to C. diversum and 

C. laciniosum. Single cells are also seen rarely. 

Reference: Hustedt (1927-1930). 

Chaetoceros atlanticum Cleve 

Very rare in ocean samples in both winter and 

spring. A widespread oceanic species in temperate and tropical waters. Reported from the 

Caribbean by Margalef (1968), Roukiyainen 

et al. (1973), and Sander (1976). The variety 

neapolitana seems more common to tropical 

waters. The morphology of some chains resembles C. laciniosum, from which it can be 

distinguished by the multiple chloroplasts and 

central tubule of the valve. Reference: Hustedt (1927-1930). 

* Chaetoceros boreale Bailey 

Rarely found in a spring sample on the ocean 

side of Carrie Bow Cay. This rather distinctive 

species is normally found in temperate or cold 

waters. It has not been reported previously 

from the Caribbean. Reference: Hustedt 

(1927-1930). 

Chaetoceros coarctatum Lauder 

A rarely found species from ocean-side samples in winter and spring. Generally found in 

the tropics, occasionally in temperate waters. 

Previously reported from the Caribbean region by Hargraves et al. (1970), Margalef 

(1968), and Takano (1960). Rarely vorticellid 

protozoans are attached to several cells in a 

chain. Reference: Hustedt (1927-1930). 

Chaetoceros compressum Lauder 

Only seen once in a winter sample. This species is normally considered a coastal species 

from temperate or boreal waters. Nevertheless, 

it is widely reported from the Caribbean, 

sometimes as a dominant species (Sander, 

1976). Reference: Hustedt (1927-1930). 

Chaetoceros curvisetum Cleve 

Occasionally seen in spring, but only in samples from the ocean side of Carrie Bow Cay. 

Widely distributed in coastal waters in temperate and tropical regions. Reported by most 
156 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 85.—Light photomicrographs of representative diatoms: a, Bacteriastrum hyalinum, valve 

view of cell with 17 setae (Hyrax mount); b, Chaetoceros peruvianum, girdle view of entire cell 

(only bases of setae are shown; Hyrax mount); c, Asteromphalus hookeri, valve view of cell (Hyrax 

mount); d, Astenonella notata, isolated cell in valve view (Hyrax mount); e, Coscinodiscusjonesianus, 

valve view (Hyrax mount);/, Thalassionema nitzschioides, eight-celled colony in girdle view (living 

material). (Scale = 10 /xm.) 
NUMBER 12 157 

FIGURE 86.—Scanning electron photomicrographs of representative diatoms: a, Bacteriastrum sp., 

probably B. furcatum (sensu Boalch, 1975), valve view of terminal cell; b, B. elongatum, terminal 

and interior cells of a chain; c, Chaetoceros lorenzianum, girdle view; d, Hemialus hauckii, girdle view 

of an isolated valve. (Scale = 10 jum.). 
158 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 
NUMBER 12 159 

authors throughout the Caribbean. Reference: Hustedt (1927-1930). 

Chaetoceros danicum Cleve 

Occasionally found in lagoon samples in winter. Normally characterized as a temperate 

species favoring somewhat reduced salinity. 

Previously reported from the Caribbean by 

Margalef (1968) and Sander (1976). Reference: Hustedt (1927-1930). 

Chaetoceros decipiens Cleve 

Rarely observed in winter samples from the 

lagoon and ocean side of Carrie Bow Cay. 

Nominally an oceanic species from temperate 

and boreal waters, it has been previously reported from the Caribbean by most authors. 

The morphology of the species is variable; the 

distinctive features distinguishing it from C. 

lorenzianum are the nature of fusion of the setae 

and the presence of resting spores in C. lorenzianum. Fusion of the setae is a somewhat 

unreliable character in nature, and spores are 

rarely seen; it is possible that misidentifications have occurred, particularly in the case 

of narrow cells and short chains. Minor differences between the two are apparent at the 

ultrastructural level (Evenson and Hasle, 

1976). Reference: Cupp (1943). 

Chaetoceros didymum Ehrenbert 

Fairly common in ocean-side samples in both 

spring and winter. Widely distributed in temperate and warm waters; also widely distributed in the Caribbean. Reference: Hustedt 

(1927-1930). 

Chaetoceros diversum Cleve 

Found in winter and spring in most samples. 

Considered to be a tropical coastal species, it 

is occasionally reported also in temperate waters. Previously reported from the Caribbean 

by Sander (1976) and Takano (1960). This 

species is easily confused with C. laeve LeuFIGURE 87.—Scanning electron photomicrographs of representative diatoms: a, Actinocyclus ehrenbergii, interior side of a 

small specimen; b, Trigonium formosum, external side of a fivesided form; C, Rhizosolenia calcaravis, side view of the valve; 

d, Triceratium pentacrinus, external side; e, Isthmia enervis, girdle 

view of the entire cell. (Scale = 10 ftm.) 

duger-Fort morel and perhaps the two should 

be synonymous. Sander and Takano include 

both these species in their lists. Short chains 

may resemble C. affine. Reference: Cupp 

(1943). 

Chaetoceros laciniosum Schutt 

[= C. distans Cleve] 

Occasionally present in ocean-side samples in 

spring. This species is common and widely 

distributed in temperate waters but also occurs in the tropics. Previously reported from 

the Caribbean by Hulburt (1968) and Sander 

(1976), the latter as "dominant." Chaetoceros 

laciniosum is another highly variable species, 

and in its growth forms can resemble C. pelagicum Cleve and C. breve Schutt. Superficially 

some chains also resemble C affine and narrow 

forms of C. atlanticum. Reference: Hustedt 

(1927-1930). 

Chaetoceros lorenzianum Grunow (Figure 86c) 

Found in most samples in winter and spring. 

A common coastal form from temperate waters, it also is widely distributed in the tropics. 

It is widely reported, sometimes in abundance, 

from the Caribbean. Narrow chains particularly can be confused with C. decipiens (see 

comment for that species). Reference: Hustedt 

(1927-1930). 

Chaetoceros pendulum Karsten 

Seen once in a spring tow from the ocean side. 

A fairly rare oceanic species characteristic of 

tropical waters. Reports from the Caribbean 

by Sander (1976) and Takano (1960). Reference: Cupp (1943). 

Chaetoceros peruvianum Brightwell (Figure 85b) 

Found in a spring tow from the ocean side. A 

very common and widely distributed species 

in temperate and tropical waters, although 

rarely abundant. Present in both coastal and 

oceanic areas. Widely reported from the Caribbean (for instance, Sanders, 1976; Takano, 

1960), as short chains and single cells. Reference: Cupp (1943). 

Chaetoceros pseudocurvisetum Mangin 

Occasionally in winter samples. A coastal species widely distributed in temperate and trop-
160 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ical areas, previously reported from the Caribbean by Hagelstein (1938), Margalef (1968), 

and Takan o (1960). Reference: Cup p (1943). 

Chaetoceros tortissimum Gran 

Seen only once in a winter tow on the ocean 

side of Carrie Bow Cay. This is a characteristically temperat e coastal species only occasionally seen and rarely of any abundance . It 

has been reported from the Caribbean by 

Margalef (1968) and Takan o (1960). Reference: Hendey (1964). 

Climacodium biconcavum Cleve 

Rare ; observed in only one tow from the ocean 

side in spring. A widely distributed tropical 

oceanic species. Previously reported from the 

Caribbean by Sander (1976) an d Takan o 

(1960). Reference: Hustedt (1927-1930). 

Climacodium frauenfeldianum Grunow 

Fairly common in the spring in both lagoon 

and ocean-side samples. Like C biconcavum, 

this species is found only rarely outside the 

tropics, but it is apparently more common 

that C. biconcavum. Th e shape of the intercellular apertur e is apparently the only distinguishing characteristic between the two species: elliptical or quadrat e in C frauenfeldianum, 

lanceolate or diamond-shaped in C biconcavum. 

Sande r (1976) and Takan o (1960) have reported it in Caribbean waters. Reference: 

Cup p (1943). 

Coscinodiscus asteromphalus Ehrenberg 

Occasionally seen in ocean tows in the spring 

and lagoon samples in the winter. This is one 

of the most common Coscinodiscus species in 

temperat e coastal waters. It is morphologically variable and may be confused with other 

species, and has rarely been reported from the 

tropics. Th e only previous Caribbean record 

is Hargraves et al. (1970). Reference: Hustedt 

(1927-1930). 

Coscinodiscus centralis Ehrenberg 

Present in one sample in the spring from the 

ocean side. Mor e common in oceanic waters, 

this species is also found in coastal temperat e 

areas. Reported from the Caribbean by Bucanan (1971), Hargraves et al. (1970), Marshall (1973), and Takan o (1960). Reference: 

Hustedt (1927-1930). 

Coscinodiscus concinniformis Simonsen 

Occasionally seen in ocean-side samples in 

both spring and winter. This species was established by Simonsen (1974) and includes in 

part C. concinnus in the sense of Hustedt (1927-

1930). Th e relationship of C. concinniformis to 

C concinnus, C. concinnoides, C granii, C. nobilis 

and C. wailesii is discussed by Simonsen (1974: 

14-16). At least some of the tropical records 

of C. concinnus are probably C. concinniformis, 

since the latter appears to be a tropical species, 

whereas the former is found in "higher latitudes. " Assuming this to be true, C. concinniformis is widespread in the Caribbean. Reference: Cup p (1943); Simonsen (1974). 

Coscinodiscus jonesianus (Greville) Ostenfeld (Figure 85<?) 

Found once in a winter sample from the lagoon side. Widespread in tropical and temperate coastal areas. Previously reported from 

the Caribbean by Hargraves et al. (1970), 

Margalef (1968), and Takan o (1960). Reference: Hustedt (1927-1930). 

Coscinodiscus nitidus Gregory 

A distinctive species occasionally found in 

lagoon samples in both spring and winter. 

Commonly a benthic species. A widespread 

species of coastal regions. Commonly reported 

throughout the Caribbean region. Reference: 

Hustedt (1927-1930). 

Coscinodiscus rothii (Ehrenberg) Grunow 

Seen rarely in a lagoon sample in winter. A 

widespread species often seen in waters of 

reduced salinity. Previously reported from the 

Caribbean by Hargraves et al. (1970) and 

Hagelstein (1938). This is a morphologically 

variable species, and the taxonomy of the 

group Fasciculati, to which this species belongs, is somewhat confused. Reference: Hustedt (1927-1930). 

Coscinodiscus sp. 

Rarely seen in winter in lagoon samples, this 

species has features that resemble both C. 

perforatus Ehrenberg and C. apiculatus Ehren-
NUMBER 12 161 

berg. The former species has been reported 

several times from the Caribbean, whereas the 

latter has not. 

Cylindrotheca closterium (Ehrenberg) Reimann and 

Lewin 

[= Nitzschia closterium (Ehrenberg) Smith] 

Rare to common in all samples, but more 

common in samples from shallow water. This 

species is ubiquitous in the plankton and benthos from boreal to tropical regions. Previously reported from the Caribbean by Margalef (1957) and Hagelstein (1938). Reference: Hendey (1964; as N. closterium). 

Eucampia zoodiacus Ehrenberg 

Seen once in a lagoon sample in winter. A 

widespread coastal temperate species, uncommon in the tropics. Previously reported from 

the Caribbean by Buchanan (1971) and 

Sander (1976). Reference: Cupp (1943). 

Guinardia flaccida (Castracane) Peragallo 

Fairly common in winter from both lagoon 

and ocean-side samples. Widespread and common in coastal and oceanic regions; it is predominantly temperate but also found in tropical areas. Commonly reported in the Caribbean region (for instance, Sander, 1976). Reference: Hustedt (1927-1930). 

Hemiaulus hauckii Grunow in Van Heurck (Figure 86d) 

Found to be fairly common in ocean-side 

samples in both spring and winter. This species occurs worldwide in tropical waters, occasionally abundant. Commonly reported 

from the Caribbean (for instance, Hargraves 

et al., 1970; Sander, 1976). Reference: Hustedt (1927-1930). 

Hemiaulus membranaceus Cleve 

Distribution similar to that of H hauckii in 

Belize waters. Normally this species is a rare 

member of tropical plankton. The polymorphism of this species has been discussed by 

Sournia (1968). Previously recorded from the 

Caribbean by Hargraves et al. (1970) and 

Marshall (1973). Reference: Cupp (1943). 

Hemiaulus sinensis Greville 

Found rarely in lagoon samples in spring and 

ocean-side samples in winter. A widespread 

but uncommon species of tropical areas. Previously reported from the Caribbean by several investigators. Reference: Hustedt (1927-

1930). 

Isthmia enervis Ehrenberg (Figure 81e) 

Commonly found in most samples, winter and 

spring. This distinctive species is normally 

epiphytic on algae and seagrasses in tropical 

and temperate waters; more common in the 

tropics. Several investigators have reported it 

from the Caribbean. Reference: Hustedt 

(1927-1930). 

Nitzschia bicapitata Cleve 

This species was rarely seen in ocean-side 

samples in both seasons. It is a widespread 

oceanic species from polar regions to the tropics. It has previously been reported from the 

Caribbean by Hargraves et al. (1970). Reference: Hasle (1960). 

Nitzschia marina Grunow 

Found occasionally in ocean-side samples in 

the spring. A widespread species in tropical 

waters. Previously reported from the Caribbean by Hargraves et al. (1970). Reference: 

Simonsen (1974). 

Nitzschia pseudodelicatissima Hasle 

Seen once in a winter sample from the ocean 

side of Carrie Bow Cay. This species is difficult 

to distinguish from N delicatissima Cleve in the 

light microscope. The distribution of this species in tropical waters is highly uncertain. 

Several authors have reported N. seriata in the 

tropical Atlantic and Caribbean but Hasle 

(1971) has shown that N. seriata does not occur 

south of 45°N; several other seriate species are 

morphologically similar. Some of the tropical 

and Caribbean records are probably N. pseudodelicatissima or a closely related species. The 

only listing positively noted as N. pseudodelicatissima is by Buchanan (1971). Reference: Hasle (1971). 

Odontella aurita (Lyngbye) Agardh 

[= Biddulphia aurita (Lyngbye) Brebisson] 

Present in a sample from the lagoon side in 

winter. This is a benthic species, often epiphy-
162 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

tic but occasionally found in the plankton. It 

is widespread in temperate coastal waters, less 

common in the tropics. Previously reported 

from the Caribbean by Hagelstein (1938) and 

Sander (1976). The distinction between Biddulphia and Odontella is reviewed by Simonsen 

(1974). Reference: Hustedt (1927-1930). 

Odontella mobiliensis (Bailey) Grunow 

[= Biddulphia mobiliensis Grunow in Van 

Heurck] 

Occasionally seen in lagoon samples of both 

seasons. A widespread species characteristic of 

temperate and tropical coastal plankton. 

Widely reported from the Caribbean (for instance, Hargraves et al., 1970; Sander, 1976; 

Margalef, 1968). Reference: Hustedt (1927-

1930). 

Odontella sinensis (Greville) Grunow 

[= Biddulphia sinensis Greville] 

Comments for O. mobiliensis apply here as well. 

Paralia sulcata (Ehrenberg) Cleve 

[= Melosira Sulcata (Ehrenberg) Kutzing] 

A widely distributed coastal form, found during winter and spring, apparently equally at 

home in the plankton and benthos. More 

common in temperate than tropical areas. 

Found in the Caribbean by Hagelstein (1938), 

Marshall (1973), and Buchanan (1971). Reference: Hendey (1964). 

*Pleurosigma chilensis Krasske 

Very rare in an ocean-side sample in spring, 

this species is rare and its distribution poorly 

known. It is one of the very few planktonic 

species of this genus. Not previously reported 

from the Caribbean, although the P. pelagicum 

reported by Takano (1960) could be this species. Reference: Krasske (1941). 

Rhizosolenia acuminata (Peragallo) Gran 

Found rarely in spring in samples from the 

ocean side of Carrie Bow Cay. An oceanic 

species commonly found in tropical waters. 

Reported previously by several investigators 

from the Caribbean (Sander, 1976; Takano, 

1960). Reference: Hustedt (1927-1930). 

Rhizosolenia alata Brightwell 

Occasional in samples from the ocean side in 

spring. A widespread and often abundant species from temperate and tropical coastal waters. This species occurs in several morphological forms, of which the form indica (Peragallo) 

Ostenfeld was also occasionally seen. Often 

observed in the Caribbean (Hargraves et al., 

1970; Marshall, 1973; and others). Reference: 

Hustedt (1927r1930). 

Rhizosolenia calcaravis Schultze (Figure 81c) 

Common in all samples from both seasons. 

This species is nominally an oceanic one, but 

it is also in coastal tropical waters. Widespread 

and occasionally abundant throughout the 

Caribbean. Reference: Hustedt (1927-1930). 

Rhizosolenia castracanei Peragallo 

Found once in an ocean-side sample in winter. 

Nearly always found in tropical waters. Previously reported from the Caribbean by Hargraves et al. (1970), Margalef (1968), and 

Marshall (1973). Reference: Hustedt (1927-

1930). 

Rhizosolenia delicatula Cleve 

Seen once in a winter sample from the ocean 

side. This species is usually found in boreal or 

temperate coastal waters. Since it seems to 

differ from R. stolterfothii mainly in the range 

of cell diameter, number of chloroplasts, and 

degree of bending of the cell chains, this record may be merely a narrow specimen of R. 

stolterfothii with a minimum bending of the 

chain. Rhizosolenia delicatula has been reported 

from the Caribbean by Margalef (1968) and 

Takano (1960). Reference: Hustedt (1927-

1930). 

Rhizosolenia hebetata Bailey forma semispina (Hensen) Gran 

Rarely seen in a winter sample from the ocean 

side. An oceanic species common to temperate 

and tropical waters. In north temperate waters 

the spine is much reduced and the taxon is 

called R. hebetata forma hiemalis Gran. Widely 

distributed in the Caribbean (Marshall, 1973, 

and others). Reference: Hustedt (1927-1930). 

Rhizosolenia imbricata Brightwell 

Found once in a winter sample from the ocean 

side. Widely distributed in tropical and temperate oceanic and coastal waters. Several 

authors differentiate R. shrubsolei from this 
NUMBER 12 163 

species, based on the former's narrower diameter and occurrence in cooler waters, but 

probably they should not be separated. Widespread throughout the Caribbean. The variety 

shrubsolei has been reported by Hagelstein 

(1938) and Takano (1960). Reference: Hustedt (1927-1930). 

Rhizosolenia robusta Norman 

Occasionally seen in ocean-side samples in 

winter. This species is found in oceanic and 

coastal waters from tropical and occasionally 

from temperate areas. Previously found in the 

Caribbean by Hargraves et al. (1970), Margalef (1968), and Takano (1960). Reference: 

Hustedt (1927-1930). 

Rhizosolenia stolterfothii Peragallo 

Rare in winter samples from the ocean side of 

Carrie Bow Cay. See comments for R. delicatula. Widespread in the Caribbean (for instance, Hargraves et al., 1970; Sander, 1976). 

Reference: Hustedt (1927-1930). 

Rhizosolenia styliformis Brightwell 

Found in both winter and spring in samples 

from the ocean side. Widely distributed in 

coastal and oceanic areas of tropical and temperate waters. Widespread in the Caribbean 

(Hargraves et al., 1970; Sander, 1976; among 

others). Reference: Hustedt (1927-1930). 

Skeletonema costatum (Greville) Cleve 

Rare in winter samples. Widespread throughout the world's oceans. Many authors separate 

this species from & tropicum Cleve on the basis 

of multiple chloroplasts in the latter. The 

Belize specimens all had two chloroplasts so 

that they are unquestionably S. costatum. 

Widely reported from the Caribbean, often 

perhaps as S. tropicum. Reference: Cupp 

(1943). 

Stephanopyxis palmeriana (Greville) Grunow 

Seen once in a lagoon sample in winter. This 

is a coastal species common to tropics, occasionally ranging into temperate waters. Previously reported from the Caribbean by Margalef (1968) and Takano (1960). Reference: 

Cupp (1943). 

Thalassionema nitzschioides Grunow (Figure 85/) 

Rare in winter samples from both the lagoon 

and ocean sides. Widespread in oceanic and 

coastal areas in temperate and tropical waters, 

and common in the Caribbean. Reference: 

Cupp (1943). 

Thalassiosira eccentrica (Ehrenberg) Cleve 

[= Coscinodiscus excentricus Ehrenberg] 

Uncommon but usually present in all samples 

in both spring and winter. A widely distributed tropical and temperate species in both 

coastal and oceanic waters. Several other species resemble T eccentrica closely and electron 

microscopy is often useful in differentiating 

similar species. Found throughout the Caribbean, and often listed as C. excentricus; the 

possibility of occasional misidentifications 

cannot be excluded. Reference: Cupp (1943; 

as C. excentricus). 

* Thalassiosira lineata Jouse 

Seen once in a spring sample from the ocean 

side. Apparently widely distributed in warmer 

waters. This species resembles T leptopus (Grunow) Hasle and Fryxell, and electron microscopy is often necessary to distinguish the two. 

Not previously reported from the Caribbean, 

although several researchers report Coscinodiscus lineatus Ehrenberg, which is synonymous 

with T. leptopus. Reference: Hasle and Fryxell 

(1977). 

Thalassiothrix frauenfeldii Grunow 

Seen occasionally in ocean-side samples in 

winter. A widely distributed oceanic and 

coastal species in tropical and temperate waters. Several investigators have previously 

found it in the Caribbean (for example, 

Sanders, 1976). Reference: Cupp (1943). 

* Thalassiothrix vanhoeffenii Heiden 

Found rarely in ocean-side samples in spring 

and winter. A tropical oceanic species, T. 

vanhoeffenii is similar to T. longissima Cleve and 

Grunow from which it differs in the lack of 

marginal spines. Not previously reported from 

the Caribbean, although Hagelstein (1938) 

and Margalef (1968) list T. longissima. Reference: Simonsen (1974). 

* Triceratium antediluvianum (Ehrenberg) Grunow 

Occasionally present in winter samples from 

the lagoon side. This is a coastal species in 
164 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

temperate and tropical waters. Both quadraand pentavalves were seen. Not previously 

reported from the Caribbean, but has been 

seen in the Gulf of Mexico (pers. obs.). Reference: Hustedt (1927-1930). 

Triceratium pentacrinus (Ehrenberg) Wallach (Figure 81 d) 

[= Triceratium balearicum Cleve and Grunow] 

Seen once in a lagoon sample in spring. 

Widely distributed in benthic and coastal 

samples in tropical waters. Previously reported from the Caribbean by Hagelstein 

(1938; as Biddulphia pentacrinus). Reference: 

Hustedt (1927-1930; as T balearicum); Simonsen (1974). 

Triceratium shadboltianum Greville 

Found in most samples (although uncommon) 

in spring and winter. The cells generally resemble the variety of elongata Grunow, which, 

considering species variability of this genus, is 

of questionable taxonomic status. The species 

is benthic or planktonic in tropical coastal 

waters. It has been reported previously from 

the Caribbean by Hagelstein (1938; as Buddulphia orbiculata (Shadbolt) Boy), Hargraves 

et al. (1970), and Margalef (1968). 

Trigonium formosum (Brightwell) Hendey (Figure 

81b) 

[= Triceratium formosum Brightwell] 

Occasionally found in samples of both seasons 

from the lagoon side of Carrie Bow Cay. 

Three-, four-, and five-sided valves were seen, 

all of which, according to some authors, have 

distinct taxonomic status. This species is primarily benthic, occasionally planktonic, in 

tropical coastal waters. Hargraves et al. (1970) 

have previously reported it from the Caribbean. Reference: Hustedt (1927-1930). 

Discussion 

Seventy-three diatom taxa were identified in 

this study. For most species the observed morphological variation might justify further differentiation of varieties and forms, but recent studies of 

diatoms in culture (Hargraves and Guillard, 

1974, among others) have demonstrated a broad 

morphological plasticity, which makes many 

names in the literature open to question. This 

problem will remain unresolved until more species are cultivated and examined. To date, few 

tropical species have been cultivated. 

The most diverse genus is Chaetoceros (16 species), followed by Rhizosolenia (10 species) and 

Coscinodiscus (1 species). The dominance of representatives of these genera in tropical waters has 

been noted before; Takano (1960), and Roukiyainen et al. (1973) list 42, 14, and 16 species for 

these genera, respectively, from the Caribbean. 

These are impressive numbers, even allowing for 

misidentifications. In the material studied for this 

report, several additional species of Chaetoceros, 

Coscinodiscus, and Bacteriastrum were noted which 

could not be identified with certainty. 

A valuable characteristic in the identification 

of plankton diatoms is the presence of resting 

spores, particularly within the genus Chaetoceros. 

For unknown reasons spores are rare in tropical 

waters. No spores were seen among the species 

listed here, although at least seven form spores in 

temperate waters. Margalef (1968) illustrates several species with resting spores but fails to indicate 

whether they were, in fact, observed in his samples. Likewise, Takano (1960) shows resting spores 

in C. affine, but indicates that the material used 

as an illustration came from Aomori Bay, Japan. 

Species of Bacteriastrum, Cerataulina, Rhizosolenia, 

Stephanopyxis, and Thalassiosira are also known to 

form spores, but none were seen. The almost total 

lack of resting spores in species presumably capable of spore formation presents a problem for 

those interested in speciation and biogeographical 

ecology of phytoplankton. 

It is often assumed that extensive season fluctuations of phytoplankton do not occur in tropical 

waters as compared to temperate regions (Parsons 

et al., 1977). Near Carrie Bow Cay 26 species of 

diatoms were found in spring and winter, 27 

species only in winter, and 20 species only in 

spring samples. That is, 64% of the species listed 

here appear to be seasonal in occurrence. Most 

data on annual cycles of tropical phytoplankton, 

however, are based on measurements of chlorophyll or isotopic carbon uptake and not species 
NUMBER 12 165 

succession. Thus, what appears to be a contradic- significance to these samples, which were coition may not be. Tropical waters could have an 

intricate species succession pattern and yet relatively minor fluctuations in annual production 

rates. It is perhaps premature to attach major 

lected during two seasons. Clearly needed is a 

complete analysis of the annual cycle of species 

succession and diversity coupled with primary 

productivity measurements. 

Literature Cited 

Boalch, G. 

1975. The Lauder Species of the Diatom Genus Bacteriastrum Shadbolt. Nova Hedwigia Beiheft, 53:185-

191. 

Buchanan, R. J. 

1971. Studies at Oyster Bay in Jamaica, West Indies, V: 

Qualitative Observations on the Planktonic Algae 

and Protozoa. Bulletin of Marine Science, 21:914-

937. 

Carpenter, D. J., and R.R.L. Guillard 

1971. Intraspecific Differences in Nitrate Half-Saturation Constants for Three Species of Marine Phytoplankton. Ecology, 52:183-185. 

Cupp, E. E. 

1943. Marine Plankton Diatoms of the West Coast of 

North America. Bulletin of the Scripps Institute of 

Oceanography, 5:1-238. 

Evensen, D. L., and G. R. Hasle 

1976. The Morphology of Some Chaetoceros (Bacillariophyceae) Species as Seen in the Electron Microscopes. Nova Hedwigia Beiheft, 53:153-174. 

Gallagher, J. 

1979. Population Genetics of Skeletonema costatum (Grev.) 

CI. in Narragansett Bay. Ph.D. dissertation, University of Rhode Island. 

Garrison, D. L. 

1975. Contribution of the Net Plankton and Nannoplankton to the Standing Stocks and Primary 

Productivity in Monterey Bay, California during 

the Upwelling Season. Fisheries Bulletin, 74:183-

194. 

Hagelstein, R. 

1938. Diatomaceae. In Scientific Survey of Porto Rico and the 

Virgin Islands, 8(3):313-450. New York: New York 

Academy of Sciences. 

Hargraves, P. E., R. W. Brody, and P. R. Burkholder 

1970. A Study of Phytoplankton in the Lesser Antilles 

Region. Bulletin of Marine Science, 20:331-349. 

Hargraves, P. E., and R.R.L. Guillard 

1970. Structural and Physiological Observations on 

Some Small Marine Diatoms. Phycologia, 13:163-

172. 

Hasle, G. R. 

1960. Phytoplankton and Ciliate Species from the Tropical Pacific. Skrifter det Norske Videnskaps—Akademi 

i Oslo (1960), 2:1-50. 

1971. The Distribution oi Nitzschia seriata CI. and Allied 

Species. Nova Hedwigia Beiheft, 39:171-190. 

Hasle, G. R., and G. A. Fryxell 

1977. The Genus Thalassiosira: Some Species with a Linear Areola Array. Nova Hedwigia Beiheft, 54:15-47. 

Hendey, N. I. 

1964. An Introductory Account of the Smaller Algae of British 

Coastal Waters, V: Bacillariophyceae. 318 pages. London: Her Majesty's Stationery Office. 

1974. A Revised Check-List of British Marine Diatoms. 

Journal of the Marine Biological Association of the United 

Kingdom, 54:277-300. 

Hulburt, E. M. 

1968. Phytoplankton Observations in the Western Caribbean Sea. Bulletin of Marine Science, 18:388-399. 

Hustedt, F. 

1927-1930. Die Kieselalgen Deutschlands . . . Teil 1. In 

L. Rabenhorst, editor, Kryptogamen- Flora von 

Deutschland, Osterreich und der Schweiz, 7(1): 920 pp. 

Leipzig: Akademische Verlagsgesellschaft. 

1931-1959. Die Kieselalgen Deutschlands . . . Teil 2. In L. 

Rabenhorst, editor, Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz, 7(2): 845 pp. Leipzig: Akademische Verlagsgesellschaft. 

Kling, S. A. 

1975. A Lagoonal Coccolithophore Flora from Belize 

(British Honduras). Micropaleontology, 21:1-3. 

Krasske, G 

1941. Die Kieselalgen des chilenischen Kiistenplanktons. 

Archiv fiir Hydrobiologie, 38:260-287. 

Lebour, M. V. 

1930. Planktonic Diatoms of Northern Seas. 244 pages London: Royal Society. 

Malone, T C. 

1971. The Relative Importance of Nannoplankton and 

Netplankton as Primary Producers in Tropical 

Oceanic and Neritic Phytoplankton Communities. Limnology and Oceanography, 6:633-639. 
166 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Margalef, R. 

1957. Fitoplancton de las costas de Puerto Rico. Investigaci'on Pesquera, 6:39-52. 

1968. Estudios sobre el ecosistema pelagico del NE de 

Venezuela: Composicion y distribucion del fitoplancton. Memoria de la Sociedad de Ciencias Naturales 

La Salle, 70-72:139-205. 

Marshall, H. G. 

1973. Phytoplankton Observations in the Eastern Caribbean Sea. Hydrobiologia, 41:45-55. 

Murphy, L., and R.R.L. Guillard 

1976. Biochemical Taxonomy of Marine Phytoplankton 

by Enzyme Electrophoresis, I: The Centric Diatoms Thalassiosira pseudonana Hasle et Heimdal, 

Thalassiosiva fluviatilis Hustedt. Journal of Phycology, 

12:9-13. 

Parsons, T. R., M. Takahashi, and B. Hargrave 

1977. Biological Oceanographic Processes. 2nd edition, 332 

pages. New York: Pergamon Press. 

Roukiyainen, M. I., L. G. Senichkina, and L. V. Georgiyeva 

1973. Review of the Taxonomic Composition of Phytoplankton in the Central-American Seas. Fisheries 

Research Board of Canada, Translation Series, 2666: 47 

pages. Vancouver. 

Sander, F. 

1976. Quantitative and Qualitative Aspects of Inshore 

Phytoplankton off Barbados. Canadian Journal of 

Botany, 54:2306-2314. 

Simonsen, R. 

1974. The Diatom Plankton of the Indian Ocean Expedition of R.V. "Meteor" 1964-65. "Meteor" Forschungsergebnisse, series D, 19:1-66. 

Sournia, A. 

1968. Diatomees planctoniques du Canal de Mozambique et de File Maurice. Memoirs ORSTOM, 31: 

1-120. Paris. 

Takano, H. 

1960. Plankton diatoms in the Eastern Caribbean Sea. 

Journal of the Oceanographical Society of Japan, 16:180— 

184. 

Texeira, C. 

1963. Relative Rates of Photosynthesis and Standing 

Stock of the Netphytoplankton and Nanoplankton. Boletim do Instituto Oceanografico Sao Paulo, 13: 

53-60. 

Van Landingham, S. L. 

1967. Catalog of the Fossil and Recent Genera and Species of 

Diatoms and their Synonyms, Part I: Acanthoceras 

through Bacillaria. 493 pages. Lehre, Germany: J. 

Cramer. 

Werner, D., editor. 

1977. The Biology of Diatoms. 498 pages. Berkeley: University of California Press. 
Marine Algae and Seagrasses 

from Carrie Bow Cay, Belize 

James N. Norris 

and Katina E. Bucher 

ABSTRACT 

A total of 165 taxa of benthic marine algae and 

three of seagrasses (angiosperms) has been found 

in the intertidal and subtidal habitats of Carrie 

Bow Cay, South Water Cay, and Twin Cays on 

the central barrier reef, Belize. Of the algae, 52 

are Chlorophyta, 23 Phaeophyta, and 90 Rhodophyta; 34 genera, 77 species, six varieties and 

five forms represent new records for Belize. The 

annotated list includes for each taxon the date, 

page and place of valid publication of its name, 

selected references to descriptions and illustrations, collecting stations with notes on habitats, 

and records of distribution within the Caribbean. 

The nomenclature, morphology, and taxonomy 

of several species are critically reviewed. 

Introduction 

Despite the diverse marine habitats of the 

mainland coast, barrier reef, atolls, and numerous 

cays of Belize, Central America, little has been 

recorded about its marine plants. To date, only 

two floristic lists have dealt with the marine algae 

of Belize, the first of which included algal collections of C. L. Lundell and W. C. Schipp (Taylor, 

1935). Eighty-four marine algal species included 

in this paper were later incorporated into Taylor's 

comprehensive book on algae of the eastern tropical and subtropical coasts of the Americas (1960). 

James N. Norris and Katina E. Bucher, Department of Botany, 

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 

Recently, Tsuda and Dawes (1974) published a 

list of marine plants collected at Glover's Reef 

Belize. In their treatment of Polysiphonia from 

Carrie Bow Cay, Kapraun and Norris (herein: 

225) report seven taxa new to Belize. Seagrasses 

were reported in Belize by den Hartog (1970). 

These studies brought the number of benthic 

algae and seagrasses recorded from Belize to 159 

taxa. 

ACKNOWLEDGMENTS.—We are grateful to I. A. 

Abbot for studying the Helminthocladiaceae and 

Antithamnion, D. Cheney for examination and discussion of Eucheuma echinocarpum and E. schrammii, 

S. Earle for the list of species she found at Carrie 

Bow Cay (April 1974), and W. Adey and J. L. 

Connor for discussion on tropical algal species. 

W. Fenical, O. McConnell, I. Macintyre, F. R. 

Fosberg, M. Dante, B. Spracklin, T. Rath, W. 

Gerwick, and A. Cohen assisted as diving partners 

and shared their algal collections. Our thanks to 

S. Rechen and R. Sims for processing some of the 

specimens, and especially to C. Feller for the 

illustrations of algae. A. R. Tangerini skillfully 

prepared the maps and M. E. Hale printed their 

legends. In particular, we thank I. A. Abbott, C. 

J. Dawes, and R. B. Searles for reading the 

manuscript and their helpful comments, and D. 

H. Nicolson for critically reading and discussing 

our remarks on nomenclature. 

Study Area and Methods 

During three ten-day expeditions to the Belizean barrier reef in the spring (March-May) of 

167 
168 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

1976, 1977, and 1979, benthic marine algae and 

seagrasses were collected in the vicinity of Carrie 

Bow Cay (16°48'N, 88°05'W), and nearby South 

Water and Twin cays (Figures 88, 89). This area 

and the characteristics of its habitats are discussed 

by Riitzler and Macintyre (herein: 9). 

In the following presentation higher taxa are 

arranged phylogenetically, species alphabetically. 

Taxa reported for the first time in Belize are 

marked with an asterisk. The author, date, and 

page of publication of the name of each taxon are 

given, and wherever possible the original source 

was examined by us. In place of a typical synonymy we include one or more references to published descriptions and illustrations suitable for 

identification in the hope of facilitating and encouraging further studies on the algae of Belize. 

Each collecting locality (field station) is numbered and shown on the maps (Figures 88b, 89). 

Habitat data of each collecting locality are listed 

below. For Caribbean distribution, we include 

published records, from references presently 

known to us, on the Central American coast from 

Mexico's Yucatan Peninsula to Venezuela, the 

Greater and Lesser Antilles, and the Bahama 

Islands. Specimens were collected by the authors 

TWIN CAYS 

SOUTH WATER CAY 

South Water Cut 

CARRIE BOW CAY 

(20) Carrie Bow Cut 

FIGURE 88.—Belize, Central America: a, 

location of Carrie Bow Cay on the barrier reef; b, Carrie Bow Cay and vicinity, 

showing some collecting localities (stations 17-26). 

CURLEW BANK 

Curlew Cut 
NUMBER 12 169 

?^M Coral 

"d^l,C^ Coral covered 

hard bottom 

Thalassia 

Sand 

i i 

FIGURE 89.—Habitats and simplified substrate types in the vicinity of Carrie Bow Cay, with 

some collecting localities (stations 1-16). 

unless otherwise noted; the numbers with prefix 

JN refer to the authors' field notebooks. Symbols 

for reproductive condition are as follows: bisporangia (O); cruicate or tetrahedral tetrasporangia 

(©); zonate tetrasporangia (Z); carposporangia 

or cystocarps ($); spermatangia {6); and monoecious thallus {<?). Material studied will be deposited in the United States National Herbarium, 

Smithsonian Institution (US), with duplicates 

sent to the herbaria of the University of California, Berkeley (UC), the University of Michigan 

(MICH), and to the Fisheries Unit, Belize City. 
170 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Station List 

(With habitat and depth data; station numbers keyed to 

Figures 88 b, 89) 

CARRIE Bow CAY 

1. South Water Cut, along south side of channel 

between South Water and Carrie Bow cays; 

on sands, pavement rock, and coral rubble, 

and among corals and gorgonians, 3-9 m. 

2. North of cay, near 0 m to 100 m section of 

IMSWE transect; sand and coral rubble, 2-

3 m. 

3. West of cay, protected shallows of inner lagoon; sand, 1.5 m. 

4. West of cay, in Thalassia bed; sand and coral 

rubble, 2-5 m. 

5. South of cay, on reef; among corals and 

gorgonians, 2-5 m. 

6. East of cay, reef flat on lee side of reef crest; 

coral and rubble-covered hard bottom, intertidal to 1 m. 

7. East of cay, on reef crest; coral rubble and 

rock, intertidal to 1 m. 

8. Northeast of cay, high-relief spur and groove 

zone, coral and sand-filled channels, 4.5-

6.5 m. 

9. Northeast of cay, area between high- and 

low-relief spur and groove zones, corals, 7.5-

9.0 m. 

10. Northeast of cay, low-relief spur and groove 

zone, corals, 9-12 m. 

11. Northeast of cay, inner trough, vicinity of 

IMSWE transect; corals, 12.0-13.5 m. 

12. Northeast of cay, sand trough on lee side of 

outer reef; sand and coral rubble, 18-20 m. 

13. Northeast of cay, outer trough and slope of 

reef; corals and coral rubble, some sand, 18-

23 m. 

14. Northeast of cay, outer ridge of reef; coral 

rubble, 15 m. 

15. Northeast of cay, fore-reef slope, vicinity of 

IMSWE transect; corals, 18-40 m. 

16. Southeast of cay, outer ridge of reef; corals, 

15-30 m. 

PATCH REEFS AND Thalassia BEDS, SOUTH OF CARRIE Bow CAY 

17. Southwest of cay, coral patch reef; 6.5-8.0 m. 

18. Southwest of cay, coral patch reef; 7.5 m. 

19. Southwest of cay, coral patch reef; 7.5-9.0 m. 

20. Southwest of cay, in Thalassia bed; sand, 6.5-

7.5 m. 

SOUTH WATER CAY 

21. East of cay, inside of reef crest; coral-covered 

hard bottom, 0.5-1.0 m. 

22. West of cay, sheltered sand-bottom lagoon; 

on rocks, coral rubble and wood pier pilings, 

0.5-1.0 m. 

23. North of cay, lee side of reef crest; sand and 

coral rubble, 1-2 m. 

TWIN CAYS 

24. East cay of Twin Cays, among mangroves; 

roots of Rhizophora mangle Linnaeus (red mangrove) and sediments held by the roots. Intertidal to 1 m. 

25. Southeast end of east cay of Twin Cays; sand 

and coral rubble bottom, intertidal to 1.0 m. 

26. West cay of Twin Cays, channel between 

Cays; mangrove roots, and sand among Thalassia, intertidal to 2 m. 

Division CHLOROPHYTA 

Order CHAETOPHORALES 

Family CHAETOPHORACEAE 

* Entocladia vagans (B0rgesen) Taylor 

Entocladia vagans (B0rgesen) Taylor, 1960:49.—B0rgesen, 

1920:418, fig. 400 [as Endoderma vagans]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

growing within the cell walls of Griffithsia globulifera, 22 Apr 1977, JN-6991b. 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(B0rgesen, 1920). 

REMARKS.—This minute epiphyte, representing a genus new to Belize, was found within the 

same host from which it was originally described 

(B0rgesen, 1920). 

* Ulvella lens P. and H. Crouan 

Ulvella lens P. and H. Crouan, 1859:288.—Taylor, 1960:52, 

pl. 2: fig. 7. 

LOCAL DISTRIBUTION.—Carrie Bow Cay; sta 7, 

epiphytic on Cladophoropsis membranacea, 22 Apr 
NUMBER 12 171 

1977,JN-6949b. 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(Taylor, 1960); Puerto Rico (Diaz-Piferrer, 1963); 

Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—Another microscopic epiphyte that 

is a new Belizean record for the genus. 

Order ULVALES 

Family ULVACEAE 

* Enteromorpha chaetomorphoides B0rgesen 

Enteromorpha chaetomorphoides Btfrgesen, 1911:149; 1913:5, 

fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 10, 

epiphytic on Slypopodium zonale, 23 Apr 1977, JN6924 (fertile). 

CARIBBEAN DISTRIBUTION.—Jamaica, Puerto 

Rico, Virgin Islands, Barbados (Taylor, 1960); 

Cuba (Diaz-Piferrer, 1964a); Dominican Republic (Almodovar and Bonnelly, 1977); Costa Rica 

(Dawson, 1962b); Curacao (Diaz-Piferrer, 

1964b); Trinidad (Richardson, 1975); Venezuela 

(Diaz-Piferrer, 1970b). 

REMARKS.—Our specimen is in general agreement with Enteromorpha chaetomorphoides as described by B0rgesen (1911, 1913) and Taylor 

(1960); with cells mostly 16-18 jum diameter, with 

slightly larger and sometimes elongate cells (to 40 

jiim long) in the broader portions of the thallus. 

* Enteromorpha clathrata var. crinita (Roth) 

Hauck 

Enteromorpha clathrata var. crinita (Roth) Hauck, 1884:429.— 

Scagel, 1966:48, pl. 25: figs, A-E [as Enteromorpha crinita]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977,JN-7127. 

CARIBBEAN DISTRIBUTION.—Cuba, Hispaniola, 

Puerto Rico, Virgin Islands, Guadeloupe, Grenada (Taylor, 1960); Nevis, Dominica (Taylor, 

1969); Isla San Andres (Kapraun, 1972); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—The above distribution is of records for the species, Enteromorpha clathrata. These 

should be re-examined as some may possibly be 

var. crinita. The Carrie Bow Cay specimen superficially resembles E. chaetomorphoides but has cells 

much larger throughout the thallus, 18-30 /xm 

wide and (18-) 30-45 jum long, and thus is closer 

to E. clathrata var. crinita (see Norris, 1976:73-74). 

* Enteromorpha flexuosa (Wulfen ex Roth) 

J. Agardh 

Enteromorpha flexuosa (Wulfen ex Roth) J. Agardh, 1883: 

126.—Bliding, 1963:73-79, figs. 38-40. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976,JN-6287. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, St. 

Barthelemy, Barbuda, Guadeloupe, Barbados, 

Grenada, Costa Rica, Netherlands Antilles, Venezuela, Tobago (Taylor, 1960); St. Kitts, Antigua, 

Montserrat, Dominica, St. Vincent, Bequia (Taylor, 1969); Isla San Andres (Kapraun, 1972); 

Trinidad (Richardson, 1975). 

REMARKS.—This species is widespread in the 

Pacific and Atlantic Oceans (see Abbot and Hollenberg, 1976; Bliding, 1963). 

* Ulva rigida C. Agardh 

Ulva ngida C. Agardh, 1823:410.—Bliding, 1968:546, figs. 

6A-E, 7A-F, 8A-E, 9A-I, and 10A-B. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6285a,b. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Puerto Rico, 

Guadeloupe, Martinique, St. Vincent, Barbados, 

Grenada, Costa Rica, Panama, Colombia, Venezuela, Trinidad (Taylor, 1960, as U. lactuca var. 

rigida); Grand Cayman, Antigua, Dominica 

(Taylor, 1969, as "var. lactuca or near var. rigida"). 

REMARKS.—The Twin Cays specimens possess 

the microscopic dentation along the blade margins, characteristic of this species (Bliding, 1968). 

Order CLADOPHORALES 

Family CLADOPHORACEAE 

Cladophora cf. fuliginosa Kiitzing 

Cladophora fuliginosa Kiitzing, 1849:415.—Taylor, 1960:83, 

pl. 2: fig. 3, pl. 3: fig. 4. 
172 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

LOCAL DISTRIBUTION.—South Water Cay: sta 

23, 30 Apr 1979, JN-7329. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Turks Islands, Cuba, Cayman Islands, 

Jamaica, Hispaniola, Virgin Islands, St. 

Barthelemy, Guadeloupe, Martinique, Belize, 

Netherlands Antilles (Taylor, 1960); Nevis, Antigua, Dominica, St. Lucia (Taylor, 1969). 

REMARKS.—Our material resembles this taxon, 

but differs in cell size. The cells of the axes and 

branches are much larger and of similar diameter 

throughout, mostly 590 /am diameter and (2-) 3 -

4 times as long; upper cells of the branches were 

580 /xm diameter, slightly tapered towards apices, 

and up to 12 times as long. These measurements 

are almost 1.6 times the diameter reported for 

Cladophora fuliginosa (Taylor, 1960), which has 

main axes cells to 380 /xm diameter and 3-6 times 

as long, with branch cells 150-160 (-290) /xm 

diameter and 10-13 times as long as wide. 

We are aware that the earliest available name 

for this taxon is Cladophora catenata (Linnaeus) 

Kiitzing, 1843 (van den Hoek, 1963:123). However, the name "C. catenata (C. Agardh) Ardissone 

in Rabenhorst" is widely used in a different taxonomic sense (see for instance, Collins, 1909:347; 

Taylor, 1960:83), and does not include the actual 

Linnaean type. Thus, it seems, C. catenata may be 

a good candidate for rejection under Article 69 

(Stafleu et al., 1978), an action we are considering. 

Orde r SlPHONOCLADIALES 

Family SIPHONOCLADACEAE 

* Cladophoropsis macromeres Taylor 

Cladophoropsis macromeres Taylor, 1928:64; 1960:118, pl. 2: 

fig. 2. 

LOCAL DISTRIBUTION.—Twin Cays: sta 25, epiphytic on Digenia simplex, 26 Apr 1977, JN-6894a. 

CARIBBEAN DISTRIBUTION.—Cuba (Sosa, 1977); 

Puerto Rico (Diaz-Piferrer, 1963); Jamaica (Taylor, 1960; Chapman, 1961); Barbados (Taylor, 

1969). 

Cladophoropsis membranacea (C. Agardh) 

B0rgesen 

Cladophoropsis membranacea (C. Agardh) B0rgesen, 1905:275.— 

Taylor, 1960:117, pl. 2: fig. 1, pl. 3: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6949a. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Puerto Rico, 

Virgin Islands, St. Barthelemy, Guadeloupe, 

Martinique, Barbados, Grenada, Belize, Panama, 

Netherlands Antilles, Venezuela, Tobago (Taylor, 1960); Grand Cayman, St. Kitts, Antiqua, 

Dominica, St. Lucia, St. Vincent, Bequia (Taylor, 

1969); Costa Rica (Dawson, 1962b); Panama 

(Earle, 1972); Isla San Andres (Kapraun, 1972). 

Family VALONIACEAE 

Dictyosphaeria cavernosa (Forsskal) B0rgesen 

Dictyosphaeria cavernosa (Forsskal) B0rgesen, 1932:2.—Taylor, 

1960:116, pl. 7: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6401; sta 7, 21 Apr 1977, JN6977; sta 10, 23 Apr 1977, JN-6915; sta 14, 27 

Apr 1977, JN-6797; sta 18, 25 Apr 1977, JN-6741. 

South Water Cay: sta 21, 28 Apr 1977, JN-6715; 

sta 23, 30 Apr 1979, JN-7328. Twin Cays: sta 25, 

26 Apr 1977, JN-6862. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Turks Islands, Cuba, Cayman Islands, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

St. Barthelemy, Nevis, Guadeloupe, Martinique, 

Barbados, Netherlands Antilles, Panama, Isla las 

Aves (Taylor, 1960); Antiqua, Dominica, St. Lucia, Bequia, Grenada (Taylor, 1969); Belize 

(Tsuda and Dawes, 1974); Costa Rica (Dawson, 

1962b). 

* Valonia macrophysa Kiitzing 

Valonia macrophysa Kiitzing, 1843:307.—Taylor, 1960:110, pl. 

2: fig. 6, pl. 7: fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

27 Apr 1977, JN-6405; sta 9, 29 Apr 1977, JN6819b. 
NUMBER 12 173 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Cayman Islands, Jamaica, Hispaniola, Virgin Islands (Taylor, 1960); Nevis, Antiqua, Barbados, 

St. Vincent, Bequia (Taylor, 1969); Puerto Rico 

(Diaz-Piferrer, 1963); Curacao (van den Hoek, 

1969); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.— Valonia macrophysa is often covered 

with small epiphytic filamentous and thin crustose coralline red algae. 

Valonia ventricosa J. Agardh 

Valonia ventricosa J. Agardh, 1885:96.—Taylor, 1960:110, pl. 

9: figs. 4, 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

30 Mar 1976, JN-7257; sta 7, 22 Apr 1977, JN6963; sta 9, 27 Apr 1977, JN-6406; sta 14, 17 Apr 

1977, JN-6798. South Water Cay: sta 23, 30 Apr 

1979, JN-7318. Twin Cays: sta 25, 26 Apr 1977, 

JN-6864; sta 26, 29 Apr 1979, JN-7296. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Cayman Islands, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Barbados, Grenadines, Grenada, Old 

Providence Island, Panama, Netherlands Antilles, Tobago (Taylor, 1960); St. Kitts, Nevis, 

Antigua, St. Lucia, St. Vincent, Bequia (Taylor, 

1969); Belize (Tsuda and Dawes, 1974); Costa 

Rica (Dawson, 1962b). 

Family BOODLEACEAE 

Struvea anastomosans (Harvey) 

Piccone and Grunow ex Piccone 

Struvea anastomosans (Harvey) Piccone and Grunow ex Piccone, 1884:20.—B0rgesen, 1913:54, fig. 39a-f. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

28 Apr 1979, coll. J. Norris and I. Macintyre, JN7492. 

CARIBBEAN DISTRIBUTION.—Jamaica, Virgin Islands, Guadeloupe, Barbados, Venezuela, Trinidad, Tobago (Taylor, 1960); Cuba (Diaz-Piferrer, 

1964a); Puerto Rico (Almodovar and Blomquist, 

1965); Dominica (Taylor, 1969); Costa Rica 

(Dawson, 1962b); Belize (Tsuda and Dawes, 

1974); Curacao, Bonaire (Diaz-Piferrer, 1964b). 

Family ANADYOMENACEAE 

Anadyomene stellata (Wulfen) C. Agardh 

Anadyomene stellata (Wulfen) C. Agardh, 1823:400.-Taylor, 

1960:125, pl. 7: fig. 2, pl. 8: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6819a; sta 10, 23 Apr 1977, JN6921; sta 14, 27 Apr 1977, JN-6799. Twin Cays: 

sta 24, on mangrove roots, 31 Mar 1976, JN-6282 

and JN-6294; sta 25, 26 Apr 1977, JN-6865; sta 

26, 22 Apr 1979, JN-7446. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin 

Islands, Guadeloupe, Martinique, Barbados, Belize, Panama (Taylor, 1960); Nevis, Antigua, 

Dominica, St. Lucia (Taylor, 1969); Costa Rica 

(Dawson, 1962b); Venezuela (Diaz-Piferrer, 

1970b); Cura?ao (Diaz-Piferrer, 1964b). 

* Microdictyon hoergesenii Setchell 

Microdictyon boergesenri Setchell, 1925:106.—Taylor, 1960:120, 

pl. 8: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

27 Apr 1977,JN-6685. 

CARIBBEAN DISTRIBUTION.—Cuba (Sosa, 1977); 

Jamaica, Guadeloupe, Virgin Islands (Taylor, 

1960); Puerto Rico (Diaz-Piferrer, 1963); Barbados (Taylor, 1969); Curasao (Diaz-Piferrer, 

1964b). 

REMARKS.—Reported from moderately deep 

water, this species (a new Belizean record of the 

genus) was found at 22 m depth in Belize. 

* Valoniopsis pachynema (Martens) B0rgesen 

Valonwpsis pachynema (Martens) B0rgesen, 1934a: 10, figs. 

1-2. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

23, 30 Apr 1979, JN-7326 (fertile?). 

CARIBBEAN DISTRIBUTION.—Cuba (Suarez, 

1973); Hispaniola (Taylor, 1960). 

REMARKS.—This is a new record of the genus 

in Belize. 
174 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Order DASYCLADALES 

Family ACETABULARIACEAE 

Acetabularia crenulata Lamouroux 

Acetabularia crenulata Lamouroux, 1816:249.—Taylor, 1960: 

105, pl. 4: fig. 5, pl. 6: fig. 12. 

LOCAL DISTRIBUTION.—Twin Cay: sta 26, on 

mangrove roots, 29 Apr 1979, JN-7303. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

St. Martin, Guadeloupe, Barbados, Colombia, 

Netherlands Antilles, Venezuela (Taylor, 1960); 

Grand Cayman (Taylor, 1969); Panama (Earle, 

1972). 

* Acetabularia myriospora 

Joly and Cordeiro-Marino 

FIGURE 90 

Acetabularia myriospora Joly and Cordeiro-Marino in Joly et 

al., 1965:80.—Howe, 1909:92, pl. 6: fig. 21, pl. 8: fig. 10 

[as Acetabulum polyphysoides f. deltoideum].—Taylor, 1960: 

105 [as Acetabularia polyphysoides f. deltoideum].—Valet, 

1969:623, pl. 44: figs. 17-20. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

growing on dead coral, in turf, 21 Apr 1977, JN7051b. 

1 mm 

FIGURE 90.—Acetabularia myriospora, habit of a thallus (JN7051b). 

CARIBBEAN DISTRIBUTION.—Cuba (Sosa, 1977); 

Bahamas (Howe, 1909; Valet, 1969). 

REMARKS.—Growing on the same coral with 

the following taxon, Acetabularia polyphysoides; it is 

distinguished by its smaller size, and fewer (5-7), 

inflated rays. Described from Bahfa, Brazil, by 

Joly and Cordeiro-Marino (Joly et al., 1965); 

Valet (1969) later considered the earlier described 

taxon, Acetabularia polyphysoides f. deltoideum 

(Howe) Collins (1909), to be a synonym of Acetabularia myriospora. 

* Acetabularia polyphysoides P. and H. Crouan 

Acetabularia polyphysoides P. and H. Crouan in Schramm and 

Maze, 1866:101.—Howe, 1909:92, pl. 6: figs. 16-20, pl. 7: 

figs. 5-9. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

growing on dead coral, in short turf, 21 Apr 1977, 

JN-7051a. 

CARIBBEAN DISTRIBUTION.—Jamaica, Guadeloupe, Barbados (Taylor, 1960). 

REMARKS.—Plants were 2-5 mm tall; 3-4 mm 

disc diameter, aplanosporangia (55-) 60-75 (-79) 

in a ray, 86-94 /xm diameter; 16-20 rays, heavily 

calcified between rays; corona superior with 6-10 

hairs. The Carrie Bow Cay specimens, though 

superficially resembling Acetabularia pusilla 

(Howe) Collins, are closer in taxonomic features 

to A. polyphysoides. 

Family DASYCLADACEAE 

* Neomeris annulata Dickie 

FIGURE 91 

Neomeris annulata Dickie, 1874:198.—Taylor, 1960:101, pl. 5: 

fig. 5, pl. 6: figs. 4-6. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, on 

mangrove bark, 26 Apr 1977, JN-6863, and 29 

Apr 1979,JN-7398. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, Nevis, 

Guadeloupe, Martinique, Barbados (Taylor, 

1960); Cuba (Diaz-Piferrer, 1964a); St. Thomas 

(Valet, 1969); Antigua, Dominica (Taylor, 1969); 
NUMBER 12 175 

vWJw^ Asm 

& ;X, i v' "A n 6 * ©a: 

FIGURE 91.—Neomeris annulata, habit of a thallus (JN-7398). 

Costa Rica (Dawson, 1962b); Panama (Earle, 

1972); Curacao (Diaz-Piferrer, 1964b). 

REMARKS.—The report of this and the following species constitutes a new record for the genus 

in Belizean waters. 

* Neomeris mucosa Howe 

Neomeris mucosa Howe, 1909:84, pl. 5: figs. 1-14. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977, JN-6933; sta 7, 22 Apr 1977, JN6959; sta 9, 29 Apr 1977, JN-6820; sta 10, 23 Apr 

1977, JN-6920. South Water Cay: sta 21, 28 Apr 

1977,JN-6717. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Netherlands Antilles (Taylor, 1960); Puerto Rico 

(Almodovar, 1970); Antigua (Taylor, 1969). 

REMARKS.—This report represents a new record 

for this species along the Atlantic coast of Central 

America. Neomeris annulata has calcified sporangia 

that laterally cohere in regular bands about the 

thallus. Neomeris mucosa differs in having calcified 

sporangia free and not arranged in bands. 

Order CAULERPALES 

Family CAULERPACEAE 

* Caulerpa ambigua Okamura 

Caulerpa ambigua Okamura, 1897:4.—Eubank, 1946:410.— 

Egerod, 1952:368.—Taylor, 1960:137, pl. 10: figs. 2-9. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

growing with Bryopsis, 28 Apr 1979, JN-7414. 

CARIBBEAN DISTRIBUTION.—Jamaica, Virgin Islands, Barbados, Costa Rica (Taylor, 1960); Antigua, St. Lucia (Taylor, 1969); Cuba (Diaz-Piferrer, 1964a); Puerto Rico (Diaz-Piferrer, 1963); 

Curacao (van den Hoek, 1969); Venezuela (DiazPiferrer, 1970b). 

Caulerpa cupressoides (Vahl) C. Agardh 

Caulerpa cupressoides (Vahl) C. Agardh, 1823:441.—Taylor, 

1960:146, pl. 14: figs. 3, 4, 6, pl. 15: figs. 1-4, pl. 18: figs. 

11-13. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6393; sta 18, 25 Apr 1977, JN6743. South Water Cay: sta 23, 30 Apr 1979, JN7334 and JN-7353. Twin Cays: sta 25, 26 Apr 

1977,JN-6857. 
176 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Hispaniola, Virgin Islands, St. Barthelemy, St. Eustatius, Guadeloupe, Martinique, St. Vincent, Barbados, Grenada, Belize, Panama, Colombia, Tobago (Taylor, 

1960); Grand Cayman, Nevis, Antigua, Dominica 

(Taylor, 1969); Puerto Rico (Almod6var, 1964); 

Costa Rica (Wellington, 1973); Isla San Andres 

(Kapraun, 1972); Curacao (van den Hoek, 1969). 

Caulerpa fastigiata Montagne 

Caulerpa fastigiata Montagne, 1842:19.—Taylor, 1960:136, pl. 

10: fig. 12. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21,28 Apr 1977, JN-6729. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Hispaniola, Virgin Islands, Guadeloupe, Barbados, Belize, Panama (Taylor, 1960); Puerto Rico 

(Diaz-Piferrer, 1963); Grand Cayman (Taylor, 

1969); Trinidad (Richardson, 1975). 

Caulerpa mexicana (Sonder) Kiitzing 

Caulerpa mexicana (Sonder) Kiitzing, 1849:496.—Taylor, 

1960:141, pl. 12: figs. 2-5. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 26 

Apr 1977, JN-6860. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, Guadeloupe, Barbados, Belize, Colombia, Venezuela, 

Tobago (Taylor, 1960); Antigua, Grenada (Taylor, 1969); Costa Rica (Wellington, 1973); Curacao (Diaz-Piferrer, 1964b; van den Hoek, 1969). 

* Caulerpa paspaloides (Bory) Greville 

Caulerpa paspaloides (Bory) Greville, 1830: lxiv.—Taylor, 

1960:149, pl. 16: figs. 1-4, pl. 18: figs. 8, 14, 15. 

LOCAL DISTRIBUTION.—Twin Cays: sta 26, 29 

Apr 1979, JN-7285. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica (Taylor, 1960). 

REMARKS.—This report represents a new record 

for this species along the Caribbean coast of 

Central America. 

* Caulerpa peltata Lamouroux 

Caulerpa peltata Lamouroux, 1809d:145.—Taylor, 1960:155, 

pl. 17: fig. 2, pl. 18: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977,JN-6962. 

CARIBBEAN DISTRIBUTION.—Barbuda (Taylor, 

1960); Cuba (Diaz-Piferrer, 1964a); Puerto Rico 

(Diaz-Piferrer, 1963); Jamaica (Chapman, 1961); 

Panama (Earle, 1972); Trinidad (Richardson, 

1975); Curacao (Diaz-Piferrer, 1964b). 

Caulerpa racemosa (Forsskal) J. Agardh 

Caulerpa racemosa (Forsskal) J. Agardh, 1873:35.—Taylor, 

1960:151, pl. 17: figs. 1, 3, 4, 6, 7, pl. 18: figs. 2-5, 7. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6950, JN-6952, and 21 Apr 1977, 

JN-6975 and JN-6980. Twin Cays: sta 24, 31 Mar 

1976, JN-6288; sta 25, 26 Apr 1977, JN-6858; sta 

26, 29 Apr 1979, JN-7282 and JN-7304. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, St. Barthelemy, St. 

Eustatius, Antigua, Guadeloupe, Martinique, 

Barbados, Grenadines, Grenada, Belize, Costa 

Rica, Panama, Colombia, Netherlands Antilles, 

Isla las Aves, Venezuela, Trinidad, Tobago (Taylor, 1960); Grand Cayman, St. Kitts, Dominica, 

St. Lucia, St. Vincent, Bequia (Taylor, 1969). 

REMARKS.—The most common of the species of 

Caulerpa in the vicinity of Carrie Bow Cay. It is 

usually abundant where found. 

* Caulerpa serrulata (Forsskal) J. Agardh 

FIGURE 92 

Caulerpa serrulata (Forsskal) J. Agardh, 1823:446.—B0rgesen, 

1932:5.—Taylor, 1960:145, pl. 14: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

22 Apr 1979, JN-7402; sta 9, 1 May 1979, JN7518. 

CARIBBEAN DISTRIBUTION.—Bahamas, St. Barthelemy, Guadeloupe, Barbados, Grenada, Venezuela (Taylor, 1960). 
NUMBER 12 177 

2 mm 

FIGURE 92.—Caulerpa serrulata, habit of a portion of the 

thallus (JN-7402). 

Caulerpa sertularioides (Gmelin) Howe 

Caulerpa sertularioides (Gmelin) Howe, 1905:576.—Taylor, 

1960:144, pl. 13: figs. 1-7. 

LOCAL DISTRIBUTION.—Twin Cays: sta 25, 26 

Apr 1977, JN-6861; sta 26, 29 Apr 1979, JN-7292. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, St. Barthelemy, 

Guadeloupe, Dominica, Martinique, Barbados, 

Grenada, Belize, Old Providence Island, Costa 

Rica, Panama, Colombia, Netherlands Antilles, 

Venezuela, Trinidad, Tobago (Taylor, 1960); 

Grand Cayman, Antigua, St. Lucia, Bequia (Taylor, 1969); Isla San Andres (Kapraun, 1972). 

* Caulerpa sertularioides f. farlowii 

(Weber-van Bosse) B0rgesen 

Caulerpa sertularioides f. farlowii (Weber-van Bosse) B0rgesen, 

1907:365; 1913:133, fig. 106.—Taylor, 1960:144, pl. 13: 

figs. 4-5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

23 Apr 1979, JN-7654; sta 7, 21 Apr 1977, JN6985. 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(B0rgesen, 1907); Aruba Island (Taylor, 1942). 

* Caulerpa verticillata J. Agardh 

Caulerpa verticillata]. Agardh, 1847:6.—Taylor, 1960:138, pl. 

10: figs. 1, 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

27 Apr 1977, JN-6412, and 29 Apr 1977, JN6831; sta 10, 23 Apr 1977, JN-6913, and 25 Apr 

1977, JN-6965. Twin Cays: sta 24, 31 Mar 1976, 

JN-6279; sta 25, 26 Apr 1977, JN-6859; sta 26, 

29 Apr 1979,JN-7294. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Puerto Rico, 

Virgin Islands, Antigua, Guadeloupe, Martinique, Grenada, Netherlands Antilles, Venezuela 

(Taylor, 1960); Grand Cayman (Taylor, 1969); 

Costa Rica (Dawson, 1962b); Trinidad (Richardson, 1975). 

REMARKS.—Though encountered in several 

habitats, intertidal and subtidal (to 12 m depth), 

this species was particularly abundant under the 

mangroves at Twin Cays where it formed extensive green carpets. 

Family BRYOPSIDACEAE 

Bryopsis pennata Lamouroux 

Bryopsis pennata Lamouroux, 1809c: 134.—Taylor, 1960:132, 

pl. 9: fig. 12. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-6984 and JN-7052. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin 

Islands, Guadeloupe, Martinique, Barbados, Belize, Colombia, Netherlands Antilles, Trinidad 

(Taylor, 1960); Antigua (Taylor, 1969). 

Order CODIALES 

Family CODIACEAE 

Avrainvillea longicaulis (Kiitzing) Murray and 

Boodle 

Avrainvillea longicaulis (Kiitzing) Murray and Boodle, 1889: 

70.—Taylor, 1960:160, pl. 19: fig. 1. 
178 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

30 Mar 1976, JN-7249. 

CARIBBEAN DISTRIBUTION.—Bahamas, Salt Key 

Bank, Jamaica, Virgin Islands, Antigua, Guadeloupe, Barbados, Grenada, Belize, Isla las Aves 

(Taylor, 1960); Cuba (Diaz-Piferrer, 1964a); 

Grand Cayman (Taylor, 1969); Costa Rica (Dawson, 1962b). 

REMARKS.—Our plants had blades composed 

of cylindrical and torulose filaments similar to 

Avrainvillea longicaulis (Taylor, 1960). However, 

there is difficulty in distinguishing A. longicaulis 

from A. nigricans Decaisne solely on the basis of 

the morphology of the blade filaments. Chapman 

(1961) suggests A. longicaulis (as "A. mazei") may 

prove to be a variety of A. nigricans. Until more 

collections and detailed studies have been made 

on these taxa it seems best to continue to recognize the two as separate species. 

There has been some historical confusion in 

application of the epithet longicaulis involving 

both taxonomy and nomenclature. The basionym, Rhipilia longicaulis Kiitzing (1858:13), pertains to material with a blade composed of torulose filaments. Unfortunately, this epithet was 

transferred to Avrainvillea by Murray and Boodle 

(1889:70), in the sense of moniliform filaments. 

Howe (1905:586), on examining portions of the 

type of Rhipilia longicaulis Kiitzing (Sonder Herbarium, MEL), concluded that it was torulose 

"the same as the more recently published A. mazei 

[Murray and Boodle, 1889:70]", and later he 

(1907:508-510) recognized them as A. longicaulis, 

reducing A. mazei to synonymy. 

B0rgesen (1908), while accepting this taxonomy, rejected the name Avrainvillea longicaulis as a 

nomen confusum and called the taxon A. mazei, 

an opinion more recently followed by Chapman 

(1961). 

Gepp and Gepp (1911:40-42), in reviewing the 

situation, questioned the identification of the 

type, Rhipilia longicaulis (based on their analysis 

of Kiitzing's published illustration), and suggested it be identified as "Avrainvillea sordida Murray and Boodle" [= A. levis Howe, 1905] a later 

homonym of A. sordida (Montagne) P. and H. 

Crouan ex Maze and Schramm (1878). 

We accept the taxonomy and nomenclature 

used by Howe (1905, 1907), and note that this 

was also accepted by Taylor (1960) without discussion. 

Codium intertextum Collins and Hervey 

Codium intertextum Collins and Hervey, 1917:54.—Silva, 1960: 

500, pl. 107: figs, a-e, pl. 108: fig. a. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977,JN-6976. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Kitts, 

Guadeloupe, Barbados, Netherlands Antilles, 

Venezuela (Taylor, 1960); Cuba (Diaz-Piferrer, 

1964a); Belize (Tsuda and Dawes, 1974); Costa 

Rica (Dawson, 1962b). 

Family UDOTEACEAE 

Halimeda copiosa Goreau and Graham 

Halimeda copiosa Goreau and Graham, 1967:432-441.—Colin vaux, 1968:30, fig. 1 [as Halimeda hederacea]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977, JN-6930; sta 15, 5 Apr 1976, JN7650. 

CARIBBEAN DISTRIBUTION.—Jamaica, Curacao 

(Colinvaux, 1968); Belize (Tsuda and Dawes, 

1974); Venezuela (Diaz-Piferrer, 1970b). 

Halimeda discoidea Decaisne 

Halimeda discoidea Decaisne, 1842:112.—Taylor, 1960:179, pl. 

24: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7010, and 21 Apr 1977, JN6935; sta 8, 29 Mar 1976, JN-6157a, and 22 Apr 

1979, JN-7403; sta 10, 23 Apr 1977, JN-6900; sta 

11, 29 Apr 1977, JN-6833; sta 14, 27 Apr 1977, 

JN-6784, and 1 May 1979, JN-7362; sta 15, 27 

Apr 1977, JN-6684; sta 17, 25 Apr 1977, JN-7017. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Panama, Colombia, Venezuela (Taylor, 1960); Antigua, St. Lucia (Taylor, 

1969); Belize (Tsuda and Dawes, 1974). 
NUMBER 12 179 

Halimeda goreauii Taylor 

Halimeda goreauii Taylor, 1962a: 173, figs. 1-7. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

30 Apr 1979, JN-7345; sta 10, 25 Apr 1977, JN6966; sta 14, 27 Apr 1977, JN-6792, and 1 May 

1979, JN-7363; sta 15, 5 Apr 1976, JN-6333, and 

27 Apr 1977,JN-6681. 

CARIBBEAN DISTRIBUTION.—Cuba (Suarez, 

1973); Jamaica (Taylor, 1962a); Puerto Rico 

(Almodovar and Blomquist, 1965); Belize (Tsuda 

and Dawes, 1974). 

Halimeda incrassata 

(Ellis and Solander) Lamouroux 

Halimeda incrassata (Ellis and Solander) Lamouroux, 1812: 

186.—Taylor, 1960:181, pl. 23: figs. 1, 4. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, 20 Apr 1977, JN-6726. Twin Cays: sta 24, 31 

Mar 1976, JN-6281, and 26 Apr 1977, JN-6851, 

and 29 Apr 1979, JN-7381; sta 26, 29 Apr 1979, 

JN-7280. 

CARIBBEAN DISTRIBUTION.—Cuba, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Martin, St. Barthelemy, Guadeloupe, Dominica, Martinique, Barbados, Grenada, Belize, Panama, 

Netherlands Antilles (Taylor, 1960); Grand Cayman (Taylor, 1969); Venezuela (Diaz-Piferrer, 

1970b). 

Halimeda opuntia (Linnaeus) Lamouroux 

Halimeda opuntia (Linnaeus) Lamouroux, 1812:186.—Taylor, 

1960:176, pl. 23: fig. 3, pl. 24: fig. 1. 

LOCAL DISTRIBUTION.— Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-6979; sta 14, 27 Apr 1977, JN6796; sta 15, 27 Apr 1977, JN-6680. Twin Cays: 

sta 24, 31 Mar 1976, JN-6289; sta 26, 29 Apr 

1979, JN-7291 and JN-7299. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Anguila Islands, Cuba, Cayman 

Islands, Jamaica, Hispaniola, Puerto Rico, Virgin 

Islands, Tortola Island, St. Barthelemy, Guadeloupe, Martinique, Barbados, Grenada, Belize, 

Old Providence Island, Costa Rica, Panama, Colombia, Netherlands Antilles, Isla las Aves, Venezuela, Trinidad, Tobago (Taylor, 1960); St. 

Kitts, Nevis, Antigua, St. Lucia, St. Vincent, 

Bequia (Taylor, 1969); Isla San Andres (Kapraun, 1972). 

Halimeda tuna (Ellis and Solander) Lamouroux 

Halimeda tuna (Ellis and Solander) Lamouroux, 1812:186.— 

Taylor, 1960:178, pl. 24: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7009; sta 8, 29 Mar 1976, JN6157b; sta 9, 29 Apr 1977, JN-6832; sta 10, 23 

Apr 1977, JN-6906 and JN-6916, and 25 Apr 

1977, JN-6969; sta 14, 27 Apr 1977, JN-6785 and 

JN-6794; sta 18, 25 Apr 1977, JN-6738. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Anguila Islands, Cuba, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Martin, Saba Bank, Guadeloupe, Barbados, Belize, 

Panama, Colombia (Taylor, 1960); St. Kitts, 

Nevis (Taylor, 1969); Costa Rica (Dawson, 

1962b); Isla San Andres (Kapraun, 1972); Venezuela (Diaz-Piferrer, 1970b). 

Penicillus capitatus Lamarck 

Penicillus capitatus Lamarck, 1813:299.—Taylor, 1960:171, pl. 

21: fig. 2, pl. 25: fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6390a. South Water Cay: sta 21, 

28 Apr 1977, JN-6727. Twin Cays: sta 25, 28 Apr 

1977, JN-6852; sta 26, 29 Apr 1979, JN-7298. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Martin, St. Barthelemy, St. Eustatius, Guadeloupe, 

Dominica, Martinique, Grenada, Belize, Old 

Providence Island, Panama, Colombia, Netherlands Antilles, Isla las Aves (Taylor, 1960); St. 

Kitts, Antigua (Taylor, 1969); Costa Rica (Dawson, 1962b); Isla San Andres (Kapraun, 1972). 

Penicillus dumentosus (Lamouroux) Blainville 

Penicillus dumentosus (Lamouroux) Blainville, 1834:553.— 

Taylor, 1960:172, pl. 21:fig. 4, pl. 25: fig. 15. 
180 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

LOCAL DISTRIBUTION.—Twin Cays: sta 26, 29 

Apr 1979, JN-7306. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Cayman Islands, Jamaica, Hispaniola, Puerto 

Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Grenada, Belize, Old Providence Island, 

Colombia (Taylor, 1960); St. Kitts, Antigua 

(Taylor, 1969). 

Penicillus lamourouxii Decaisne 

Penicillus lamourouxii Decaisne, 1842:109.—Taylor, 1960:172, 

pl. 21: fig. 1, pl. 25: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6390b. Twin Cays: sta 24, 31 

Mar 1976,JN-6280. 

CARIBBEAN DISTRIBUTION.-—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Puerto Rico, Virgin Islands, St. Eustatius, Guadeloupe, Panama 

(Taylor, 1960); Grand Cayman (Taylor, 1969); 

Belize (Tsuda and Dawes, 1974). 

Penicillus pyriformis A. and E. S. Gepp 

Penicillus pyriformis Gepp and Gepp, 1905:1.—Taylor, 1960: 

170, pl. 21: figs. 3, 5, pl. 25: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

30 Mar 1976, JN-7254, and 30 Apr 1979, JN7348; sta 10, 23 Apr 1977, JN-6901, and 25 Apr 

1977, JN-6881; sta 14, 27 Apr 1977, JN-6786. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Virgin Islands, St. Eustatius, Guadeloupe, Panama (Taylor, 1960); Puerto Rico (Almodovar and Blomquist, 1965); Belize (Tsuda and Dawes, 1974); 

Curasao (Diaz-Piferrer, 1964b). 

* Penicillus pyriformis f. expianatus B0rgesen 

Penicillus pyriformis f. expianatus B0rgesen, 1913:99 [as f.

 i

explanata'], fig. 82. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

5 Apr 1976, JN-6329; sta 18, 25 Apr 1977, JN6739. 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(B0rgesen, 1913). 

Rhipocephalus phoenix (Ellis and Solander) 

Kiitzing 

Rhipocephalus phoenix (Ellis and Solander) Kiitzing, 1849: 

506.—Taylor, 1960:174, pl. 22: figs. 2, 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

27 Mar 1976, JN-7048; sta 4, growing among 

Thalassia, 20 Apr 1977, JN-7006; sta 9, 27 Apr 

1977, JN-6411; sta 10, 29 Apr 1977, JN-6818, 25 

Apr 1977, JN-6882, and 23 Apr 1977, JN-6902; 

sta 14, 27 Apr 1977, JN-6786. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, Guadeloupe, 

Belize, Old Providence Island, Panama, Colombia (Taylor, 1960); Antigua (Taylor, 1969). 

* Rhipocephalus phoenix f. brevifolius 

A. and E. S. Gepp 

Rhipocephalus phoenix f. brevifolius Gepp and Gepp, 1905:4.— 

Taylor, 1960:174, pl. 22: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6392, and 27 Mar 1976, JN7049; sta 18, 25 Apr 1977, JN-6742; sta 19, 25 

Apr 1977, JN-7016. South Water Cay: sta 23, 30 

Apr 1979,JN-7316. 

CARIBBEAN DISTRIBUTION.—Bahamas, Guadeloupe (Gepp and Gepp, 1911); Jamaica (Collins, 

1909). 

* Rhipocephalus phoenix f. longifolius 

A. and E. S. Gepp 

Rhipocephalus phoenix f. longifolius Gepp and Gepp, 1905:4.— 

Taylor, 1960:174, pl. 22: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977, JN-6936, and 30 Apr 1979, JN7339; sta 4, 27 Mar 1976, JN-7050; sta 15, 5 Apr 

1976, JN-6330, and 27 Apr 1977, JN-6679; sta 

19, 25 Apr 1977,JN-7015. 

CARIBBEAN DISTRIBUTION.—Florida (Gepp and 

Gepp, 1911). 

Udotea cyathiformis Decaisne 

Udotea cyathiformis Decaisne, 1842:106.—Taylor, 1960:166, 

pl. 22: fig. 4. 
NUMBER 12 181 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 28, 

29 Mar 1976, JN-6155; sta 9, 29 Apr 1977, JN6817; sta 10, 25 Apr 1977, JN-6883. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Virgin Islands, Guadeloupe, Barbados, 

Panama (Taylor, 1960); Dominican Republic 

(Almodovar and Bonnelly, 1977); St. Vincent 

(Taylor, 1969); Belize (Tsuda and Dawes, 1974); 

Curacao (Diaz-Piferrer, 1964b); Venezuela 

(Diaz-Piferrer, 1970b). 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

epiphytic on Avrainvillea longicaulis, 30 Mar 1976, 

JN-7249b. 

CARIBBEAN DISTRIBUTION.—St. Thomas (B0rgesen, 1914); Tobago (Taylor, 1960); Puerto Rico 

(Almodovar and Blomquist, 1965); Venezuela 

(Diaz-Piferrer, 1970b). 

REMARKS.—Representing a new record of the 

genus in Belize, it was found on a blade of 

Avrainvillea. 

Udotea flabellum (Ellis and Solander) Howe 

Udotea flabeHum (Ellis and Solander) Howe, 1904:94.—Taylor, 1960:168, pl. 20: figs. 4, 5, pl. 25: fig. 3. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977, JN-6934; sta 10, 23 Apr 1977, JN6919; sta 14, 5 Apr 1976, JN-6332; sta 18, 25 Apr 

1977, JN-6735. Twin Cays: sta 25, 26 Apr 1977, 

JN-6853, JN-6854, JN-6855, and JN-6877. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Salt Key Bank, Cuba, Cayman Islands, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

St. Barthelemy, Guadeloupe, Martinique, Grenada, Belize, Panama, Colombia (Taylor, 1960); 

St. Kitts, Antigua, Dominica, St. Lucia, Barbados, St. Vincent (Taylor, 1969); Curacao, Bonaire 

(Diaz-Piferrer, 1964a). 

* Udotea cf. occidentalis A. and E. S. Gepp 

Udotea occidentalis Gepp and Gepp, 1911:127-128, figs. 18, 

22a-b, 63-65. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977,JN-6391. 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(Taylor, 1960). 

Division PHAEOPHYTA 

Order ECTOCARPALES 

Family ECTOCARPACEAE 

* Ectocarpus elachistaeformis Heydrich 

Ectocarpus elachislaeformis Heydrich, 1892:470.—Taylor, 1960: 

202, pl. 29: fig. 9.—Earle, 1969:133, fig. 28. 

Order CHORDARIALES 

Family CHORDARIACEAE 

* Cladosiphon occidentalis Kylin 

FIGURE 93 

Cladosiphon occidentalis Kylin, 1940:27.—Earle, 1969:180, figs. 

72, 75, 76. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 2, 

epiphytic on Thalassia, 27 Apr 1979, JN-7648 

(plurilocular sporangia). South Water Cay: sta 

23, epiphytic on Thalassia testudinum, 30 Apr 1979, 

JN-7325 (plurilocular and unilocular sporangia 

present). 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(Taylor, 1960); Grand Bahamas (Taylor, 1969); 

Panama (Earle, 1972). 

REMARKS.—This is a new record of the genus 

in Belize. 

Order SPHACELARIALES 

Family SPHACELARIACEAE 

* Sphacelaria tribuloides Meneghini 

Sphacelaria tribuloides Meneghini, 1840:2.—Taylor, 1960:211, 

pl. 29: fig. 6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

epiphytic on Turbinaria, 30 Mar 1976, JN-7270 

(fertile). 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

Guadeloupe, Martinique, Barbados (Taylor, 
182 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 93.—Cladosiphon occidentalis (JN-7325): a, plurilocular 

sporangia (p), assimilatory filament and hairs; b, unilocular 

sporangium (u), and assimilatory filaments (plurilocular and 

unilocular sporangia were from same thallus. 

I960); Cuba (Suarez, 1973); Antigua, Dominica 

(Taylor, 1969); Curacao, Bonaire (Diaz-Piferrer, 

1964b); Trinidad (Richardson, 1975); Venezuela 

(Diaz-Piferrer, 1970b). 

Order DICTYOTALES 

Family DICTYOTACEAE 

Dictyota bartayresii Lamouroux 

Dictyota bartayresii Lamouroux, 1809b: 331.—Taylor, 1960: 

219, pl. 30: fig. .2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7011; sta 7, 23 Apr 1977, JN6995; sta 14, 27 Apr 1977, JN-6789. South Water 

Cay: sta 21, 28 Apr 1977, JN-6724. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Virgin Islands, Antigua, Guadeloupe, Dominica, Martinique, Barbados, Grenada, Panama, Colombia (Taylor, 

1960); Belize (Tsuda and Dawes, 1974); Costa 

Rica (Dawson, 1962b); Venezuela (Taylor, 1976); 

Curasao, Bonaire (Diaz-Piferrer, 1964b); Isla San 

Andres (Kapraun, 1972). 

Dictyota cervicornis Kiitzing 

Dictyota cervicornis Kiitzing, 1859:11.—Taylor, 1960:222, pl. 

31: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6825; sta 10, 23 Apr 1977, JN6899, and 25 Apr 1977, JN-6970; sta 14, 27 Apr 

1977, JN-6788, and 5 Apr 1976, JN-6331; sta 18, 

25 Apr 1977, JN-6750. Twin Cays: sta 24, 31 Mar 

1976, JN-6273; sta 25, 26 Apr 1977, JN-6870. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Virgin Islands, St. Barthelemy, Barbuda, Nevis, Antigua, Guadeloupe, Martinique, 

Barbados, Grenadines, Belize, Panama, Colombia, Netherlands Antilles, Venezuela, Tobago 

(Taylor, 1960); Dominica (Taylor, 1969); Trinidad (Richardson, 1975). 
NUMBER 12 183 

Dictyota dichotoma (Hudson) Lamouroux 

Dictyota dichotoma (Hudson) Lamouroux, 1809b:331.—Taylor, 1960:218, pl. 31: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

5 Apr 1976, JN-6339; sta 11, 25 Apr 1977, JN6972; sta 19, 25 Apr 1977, JN-7018. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Puerto 

Rico, Guadeloupe, Martinique, Barbados, Grenada, Colombia, Netherlands Antilles, Venezuela 

(Taylor, 1960); Nevis, Antigua, St. Vincent (Taylor, 1969); Dominican Republic (Almodovar and 

Bonnelly, 1977); Belize (Tsuda and Dawes, 1974); 

Costa Rica (Dawson, 1962b); Trinidad (Richardson, 1975). 

Dictyota divaricata Lamouroux 

Dictyota divaricata Lamouroux, 1809a:43.—Taylor, 1960:221, 

pl. 31: figs. 3, 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-6981; sta 10, 23 Apr 1977, JN6918; sta 14, 27 Apr 1977, JN-6793. South Water 

Cay: sta 21, 28 Apr 1977, JN-6725. Twin Cays: 

sta 25, 26 Apr 1977, JN-6889. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Belize, Old Providence Island, Colombia, 

Netherlands Antilles, Trinidad (Taylor, 1960); 

St. Kitts, Nevis, Antigua, Dominica, St. Lucia, 

St. Vincent, Bequia (Taylor, 1969); Barbados 

(Almodovar and Pagan, 1967); Costa Rica (Dawson, 1962b); Panama (Earle, 1972); Venezuela 

(Taylor, 1976); Trinidad (Richardson, 1975). 

* Dictyota linearis (C. Agardh) Greville 

FIGURE 94 

Dictyota linearis (C. Agardh) Greville, 1830:xliii.—Earle, 

1969:161, fig. 52. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6276. 

CARIBBEAN DISTRIBUTION.—Caicos Islands, 

Cayman Islands, Jamaica, Hispaniola, Virgin Is2 mm 

FIGURE 94.—Dictyota linearis, portion of the dichotomously 

branched frond (JN-6276), showing uneven dichotomies of 

the narrow frond. 

lands, Guadeloupe, Netherlands Antilles (Taylor, 

1960); Antigua, St. Vincent (Taylor, 1969); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—This taxon is distinguished primarily by its slender, dichotomous branches. Earle 

(1969) noted that it has forms resembling Dictyota 

dichotoma, D. divaricata, and D. cervicornis and questioned its validity as a species. Further collections 

in Belize are necessary before we can ascertain 

that it is a distinct species. 

* Dilophus alternans J. Agardh 

FIGURE 95 

Dilophus alternans J. Agardh, 1882:108.—Taylor, 1960:216, 

pl. 30: fig. 3. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6822; sta 14, 27 Apr 1977, JN6791. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Virgin Islands, Barbados (Taylor, 1960); 

Antigua, Dominica, St. Lucia (Taylor, 1969); 

Curasao (van den Hoek, 1969). 

REMARKS.—Dilophus is similar in morphological appearance to Dictyota. The two genera are 

separated on anatomical differences; Dilophus has 
184 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

2 mm 

' 000 QQQQUOOQOppSOp^n i 

aaur 

50 pm 

FIGURE 95.—Dilophus alternans (JN-6822): a, portion of the 

thallus; b, transection showing a medulla more than a single 

cell thick along the margin, and a single layer of cortical 

cells. 

a medulla two or more cells thick (particularly in 

the margins or near the base) and Dictyota a 

medulla of a single layer of cells. 

Dictyopteris delicatula Lamouroux 

Dictyopteris delicatula Lamouroux, 1809b:332.—Taylor, 1960: 

227, pl. 33: fig. 3. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, 28 Apr 1977,JN-6730. 

CARIBBEAN DISTRIBUTION.—Mexico, Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

Redonda Island, Guadeloupe, Isla las Aves, Dominica, Martinique, Barbados, Grenadines, Grenada, Costa Rica, Panama, Colombia, Netherlands Antilles, Venezuela, Trinidad, Tobago 

(Taylor, 1960); St. Kitts, Nevis, Antigua, St. Vincent, Bequia (Taylor, 1969); Belize (Tsuda and 

Dawes, 1974); Isla San Andres (Kapraun, 1972). 

Lobophora variegata (Lamouroux) Womersley 

Lobophora variegata (Lamouroux) Womersley, 1967:221.— 

Taylor, 1960:231, pl. 33: fig. 4 [as Pocockiella variegata]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7004; sta 9, 29 Apr 1977, JN6823, and 25 Apr 1977, JN-6974; sta 10, 23 Apr 

1977, JN-6905; sta 14, 27 Apr 1977, JN-6795; sta 

15, 5 Apr 1976, JN-6338, and 27 Apr 1977, JN6696b; sta 18, 25 Apr 1977, JN-6740. Twin Cays: 

sta 24, 31 Mar 1976, JN-6296 and JN-6272; sta 

25, 26 Apr 1977, JN-6868, and JN-6876. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Virgin Islands, St. Barthelemy, St. Eustatius, Nevis, Guadeloupe, Martinique, Barbados, Old Providence 

Island, Panama, Colombia (Taylor, 1960); Puerto 

Rico (Almodovar, 1964); Grand Cayman, St. 

Kitts, Antigua (Taylor, 1969); Belize (Tsuda and 

Dawes, 1974); Costa Rica (Dawson, 1962b); Venezuela (Taylor, 1976); Curasao (Diaz-Piferrer, 

1964b); Trinidad (Richardson, 1975). 

REMARKS.—Two forms differing in thickness, 

color and habitat, were encountered. Dark 

brown, thin flattened forms, 105 to 120 /xm wide, 

were collected from 12 m to 14 m depths throughout the Carrie Bow Cay area (for example, JN6974); golden brown, thicker plants with ruffled 

margins (120 to 180 /xm thick) were found usually 

in clumps in the intertidal to 1 m depths along 

the eastern shore of the east cay of Twin Cays 

(for example, JN-6876). 

Padina jamaicensis (Collins) Papenfuss 

Padina jamaicensis (Collins) Papenfuss, 1977:272.—Taylor, 

1960:237, pl. 34: fig. 2 [as P. sanctae-cruas]. 
NUMBER 12 185 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-6982. Twin Cays: sta 24, 31 Mar 

1976,JN-6269. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Anguila Islands, Cuba, Cayman Islands, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

St. Barthelemy, Guadeloupe, Grenada, Belize, 

Netherlands Antilles (Taylor, 1960); Nevis, Antigua, Dominica, Barbados, St. Vincent (Taylor, 

1969); Venezuela (Diaz-Piferrer, 1970b). 

Padina vickersiae Hoyt ex Howe 

Padina vickersiae Hoyt ex Howe, 1920b:595.—Taylor, 1960: 

236, pl. 34: fig. 1. 

LOCAL DISTRIBUTION.—Twin Cays: sta 25, 26 

Apr 1977, JN-6856. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Virgin Islands, St. 

Barthelemy, Guadeloupe, Isla las Aves, Martinique, Barbados, Grenada, Belize, Panama, Colombia, Netherlands Antilles, Venezuela, Trinidad (Taylor, 1960); Puerto Rico (Almodovar, 

1964); Grand Cayman, St. Kitts, Nevis, Antigua, 

Dominica, St. Lucia, St. Vincent (Taylor, 1969); 

Costa Rica (Dawson, 1962b); Isla San Andres 

(Kapraun, 1972). 

Stypopodium zonale (Lamouroux) Papenfuss 

Stypopodium zonale (Lamouroux) Papenfuss, 1940:205.—Taylor, 1960:232, pl. 28: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7007, and 21 Apr 1977, JN6931; sta 2, 4 Apr 1976, JN-6303; sta 8, 29 Mar 

1976, JN-6159; sta 9, 29 Apr 1977, JN-6821; sta 

10, 23 Apr 1977, JN-6897; sta 11, 25 Apr 1977, 

JN-6880; sta 15, 5 Apr 1976, JN-6328, and 27 

Apr 1977,JN-6682. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Dominica, Barbados, Panama, Colombia, Venezuela, Tobago (Taylor, 

1960); Nevis, St. Vincent (Taylor, 1969); Belize 

(Tsuda and Dawes, 1974); Curasao, Bonaire 

(Diaz-Piferrer, 1964b). 

REMARKS.—Stypopodium zonale was found growing in depths of 3 m to 28 m. Plants from greater 

depths (for example, JN-6328) were much thinner, about 165 /xm wide, whereas those from 

shallower water were thicker, to 480 /xm thick, 

and with more medullary cell layers (for example 

JN-6897). 

Order DICTYOSIPHONALES 

Family PUNCTARIACEAE 

Colpomenia sinuosa (Roth) Derbes and Solier 

Colpomenia sinuosa (Roth) Derbes and Solier, 1856:11.—Taylor, 1960:260, pl. 36: fig. 1.—Wynne and Norris, 1976:2, 

fig. la,b, 2a,b, 3, 11a. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976,JN-6284. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Puerto Rico, Virgin Islands, 

Guadeloupe, Martinique, Barbados, Panama, 

Netherlands Antilles, Venezuela (Taylor, 1960); 

Dominican Republic (Almodovar and Bonnelly, 

1977); St. Kitts, Nevis, Antigua, Dominica, St. 

Lucia (Taylor, 1969); Belize (Tsuda and Dawes, 

1974); Costa Rica (Dawson, 1962b); Trinidad 

(Richardson, 1975). 

REMARKS.—Collected only once, this species 

was uncommon, at least during our spring surveys. 

Order FUCALES 

Family SARGASSACEAE 

Sargassum filipendula C. Agardh 

Sargassum filipendula C. Agardh, 1824:300.—Taylor, 1960: 

270, pl. 37: fig. 3, pl. 40: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977, JN-6932, and 20 Apr 1977, JN7008. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Puerto Rico, Virgin 

Islands, St. Barthelemy, Nevis, Redonda Island, 

Guadeloupe, Dominica, Venezuela (Taylor, 
186 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

I960); St. Kitts (Taylor, 1969); Belize (Tsudaand 

Dawes, 1974); Costa Rica (Dawson, 1962b); Isla 

San Andres (Kapraun, 1972); Trinidad (Richardson, 1975). 

Sargassum fluitans B0rgesen 

Sargassum fluitans B0rgesen, 1914:66.—Taylor, 1960:281, pl. 

39: fig. 2, pl. 40: fig. 7. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, drift, 

31 Mar 1976, JN-6286. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Belize, Costa Rica, Panama (Taylor, 

1960); Grand Cayman, St. Kitts, Nevis, Antigua, 

St. Lucia, Barbados (Taylor, 1969). 

REMARKS.—Originally described from freefloating material (B0rgesen, 1914), this species is 

only known in the pelagic state (Taylor, 1960; 

Earle, 1969). 

CARIBBEAN DISTRIBUTION.—Bahamas, Virgin 

Islands (Taylor, 1928); St. Lucia, Barbados (Taylor, 1969). 

Sargassum polyceratium Montagne 

Sargassum polyceratium Montagne, 1837:356.—Taylor, 1960: 

276, pl. 40: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

29 Apr 1977, JN-6378; sta 9, 29 Apr 1977, JN6824; sta 15, 27 Apr 1977, JN-6683; sta 18, 25 

Apr 1977,JN-6736. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Puerto Rico, Virgin Islands, Guadeloupe, St. Lucia, Barbados, Old Providence Island, Panama, Colombia, Venezuela (Taylor, 

1960); Antigua (Taylor, 1969); Belize (Tsuda and 

Dawes, 1974); Isla San Andres (Kapraun, 1972); 

Curacao, Bonaire (Diaz-Piferrer, 1964b). 

* Sargassum hystrix J. Agardh 

Sargassum hystrix J. Agardh, 1847:7.—Taylor, 1960:279, pl. 

37: fig. 1, pl. 38: fig. 2, pl. 40: fig 6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

29 Mar 1976, JN-6169; sta 10, 23 Apr 1977, JN6898; sta 14, 27 Apr 1977, JN-6790; sta 15, 5 Apr 

1976,JN-6346. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Puerto Rico, Virgin Islands, 

Guadeloupe, Grenada, Costa Rica, Trinidad 

(Taylor, 1960); St. Kitts, Dominica, St. Lucia, 

Barbados (Taylor, 1969); Dominican Republic 

(Almodovar and Bonnelly, 1977); Curacao, Bonaire (Diaz-Piferrer, 1964b); Venezuela (Taylor, 

1976). 

* Sargassum hystrix var. buxifolium Chauvin 

Sargassum hystrix var. buxifolium Chauvin in J. Agardh, 1848: 

322.—Taylor, 1928:128, pl. 18: fig. 1, pl. 19: fig. 9; 1960: 

279, pl. 38: fig. 2, pl. 40: fig. 6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 10, 

23 Apr 1977, JN-6917; sta 11, 25 Apr 1977, JN6971. 

* Sargassum polyceratium var. ovatum 

(Collins) Taylor 

Sargassum polyceratium var. ovatum (Collins) Taylor, 1928:129, 

pl. 18: fig. 7, 10, pl. 19: fig. 16. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, 29 Apr 1977, JN-6379. Twin Cays: sta 24, 31 

Mar 1976,JN-6290. 

CARIBBEAN DISTRIBUTION.—Grand Cayman 

(Taylor, 1969); Jamaica (Collins, 1901); Hispaniola (Taylor and Arndt, 1929); Costa Rica (Dawson, 1962b); Colombia (Bula Meyer, 1977). 

Turbinaria tricostata Barton 

Turbinaria tricostata Barton, 1891:218, pl. 54: fig. 3. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7005 (fertile). 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Puerto 

Rico, Virgin Islands, Guadeloupe, Netherlands 

Antilles (Taylor, 1960); Belize (Tsuda and Dawes, 

1974); Isla San Andres (Kapraun, 1972); Venezuela (Diaz-Piferrer, 1970b). 
NUMBER 12 187 

Turbinaria turbinata (Linnaeus) Kuntze 

Turbinaria turbinata (Linnaeus) Kuntze, 1898:434.—Taylor, 

1960:285, pl. 39: figs. 3-5. 

LOCAL DISTRIBUTION.—South Wate r Cay: sta 

21, 28 Ap r 1977, JN-6718 (fertile); sta 23, 30 Apr 

1979, JN-7350. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Anguilla Islands, Cuba , Jamaica , Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Antigua, Guadeloupe , Belize, Panama , Colombia, Netherlands Antilles (Taylor, 1960); 

Grand Cayman , St. Kitts, Nevis, Dominic a (Taylor, 1969); Venezuela (Taylor, 1976). 

REMARKS.—This species was very common in 

the shallow waters behind the reef crest off the 

north end of South Wate r Cay. 

Division RHODOPHYT A 

Class BANGIOPHYCEA E 

Class FLORIDEOPHYCEA E 

Order NEMALIALE S 

Family ACROCHAETIACEA E 

* Acrochaetium corymbiferum (Thuret) Batters 

Acrochaetium corymbiferum (Thuret in Le Jolis) Batters, 1902: 

59.—Taylor, 1960:312 [as Acrochaetium bornetii].—Dixon 

and Irvine, 1977:88, fig. 21 [as Audoumella corymbifera]. 

LOCAL DISTRIBUTION.—Carri e Bow Cay: sta 17, 

epiphytic on Trichogloeopsis, 25 Apr 1977, JN -

7042b (<3 an d monospores on same thallus). 

CARIBBEAN DISTRIBUTION.—With this record 

from Belize, Acrochaetium corymbiferum is now added 

to the flora of the Caribbean Sea. 

REMARKS.—Thes e specimens are epiphytic on 

Trichogloeopsis, apparently a new host. Previously 

reported from Bermuda , North Carolina (Taylor, 

1960), the Mediterranean, and England (Dixon, 

1976; Dixon an d Irvine, 1977), it represents a new 

record of the genus in Belize. 

Order GONIOTRICHALES 

Family GONIOTRICHACEA E 

* Goniotrichum alsidii (Zanardini) Howe 

Goniotrichum alsidii (Zanardini) Howe, 1914:75.—B0rgesen, 

1915:4, fig. 2 [ as Goniotrichum elegans]. 

LOCAL DISTRIBUTION.—Carri e Bow Cay: sta 1, 

epiphytic on Polysiphonia, 30 Ma r 1976, JN-7265b; 

sta 9, epiphytic on Sargassum, 29 Apr 1977, JN -

6838b. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba , 

Jamaica , Virgin Islands, Guadeloupe , Barbados, 

Tobago (Taylor, 1960); Puerto Rico (Diaz-Piferrer, 1963); Curacao (Diaz-Piferrer, 1964b); Costa 

Rica (Dawson, 1962b); Venezuela (Diaz-Piferrer, 

1970b). 

REMARKS.—This is the first report of this genus 

in Belize. 

* Acrochaetium seriatum B0rgesen 

Acrochaetium seriatum B0rgesen, 1915:32, figs. 25-28. 

LOCAL DISTRIBUTION.—Carri e Bow Cay: sta 8, 

epiphytic on Halimeda, 27 Apr 1977, JN-6423b; 

sta 9, 29 Apr 1977, JN-6845a (monospores); sta 

14, epiphytic on Sargassum blade, 27 Apr 1977, 

JN-6809a (monospores), and epiphytic on Polysiphonia, 5 Apr 1976, JN-6356 (monospores). 

CARIBBEAN DISTRIBUTION.—Virgin Islands 

(Taylor, 1960); Costa Ric a (Dawson, 1962b). 

* Acrochaetium unipes B0rgesen 

Acrochaetium unipes B0rgesen, 1915:35, figs. 31-34. 

LOCAL DISTRIBUTION.—Carri e Bow Cay: sta 9, 

epiphytic on Stypopodium, 23 Apr 1977, JN-6923 

(monospores). 

CARIBBEAN DISTRIBUTION.—Hispaniola , Virgin 

Islands (Taylor, 1960). 
188 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Family HELMINTHOCLADIACEAE 

* Liagora farinosa Lamouroux 

FIGURES 96a, 97 

Liagora farinosa Lamouroux, 1816:240.—Taylor, 1960:326, 

pl. 43: fig. 3, pl. 45: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

29 Mar 1976, JN-6161a,b (°), JN-6165, JN-6166, 

JN-6214a-e (<3), and JN-6215a-c {6); sta 10, 25 

Apr 1977, JN-6967 {6); sta 19, 25 Apr 1977, JN7021a,b (?) and JN-7022 ($). 

CARIBBEAN DISTRIBUTION.—Bahamas, Turks Islands, Cuba, Cayman Islands, Jamaica, Hispani i 

2 cm 

'.r^y *•****. 

»•> ;^v,^* 4 5Tv''i^i'/<„V *•• 

1 cm 

2 cm 

FIGURE 96.—Habits of Carrie Bow Cay species of Liagora: a, L. farinosa (JN-6161); b, Liagora 

species (JN-6216); c, L. valida (JN-6945); d, L. pinnata (JN-6163). 
NUMBER 12 189 

FIGURE 97.—Liagora farinosa, details of reproductive structures: a, carpogonial branch (cb) with 

trichogyne (tr) borne laterally off an assimilatory filament (JN-6161); b, an assimilatory filament 

with capitate spermatangia QN-6215b6"); c, mature cystocarp, showing portion of dense 

involucre filaments and large carpospores (JN-6161). 
190 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

iola, Virgin Islands, Guadeloupe, Martinique, 

Barbados (Taylor, 1960); Puerto Rico (Diaz-Piferrer, 1963); St. Kitts, Nevis, St. Lucia (Taylor, 

1969); Venezuela (Taylor, 1976); Curacao (DiazPiferrer, 1964b; van den Hoek, 1969). 

REMARKS.—This species is widespread 

throughout tropical seas (Abbott, 1945). 

In describing a new genus, Ganonema, Fan and 

Wang (1974) segregated Liagora farinosa from 

other species of Liagora on the basis of differences 

in the female reproductive structure. The genus 

Liagora Lamouroux is described as having carpogonial branches borne laterally off ordinary assimilatory filaments (cf. Yamada, 1938; Abbott, 

1945). Ganonema farinosa (Lamouroux) Fan et 

Wang (1974:492) bears carpogonial branches 

that are restricted to short filaments initiated by 

the basal cells of the assimilatory filaments (Fan 

and Wang, 1974, figs. 1-3, 6-7). This feature has 

previously been illustrated and noted for L. farinosa by B0rgesen (1927:61, figs. 33b-d) from the 

Canary Islands, and Yamada (1938, fig. 15) for 

Japanese specimens. However, none of these earlier authors gave any taxonomic or evolutionary 

significance to the unusual position of the carpogonial branch in L. farinosa. In our Belize specimens (Figure 97) we have observed carpogonial 

branches borne on short filaments originating off 

the basal cells of assimilatory filaments. Most 

recently Abbott (1976:130) stated, "Liagora farinosa has been named at least eight times (Abbott, 

1945), a clear reflection of the variation shown by 

this species, and for lack of critical evaluation of 

this variation, further comparison and discussion 

of the generic attributes of Ganonema would be 

premature." 

* Liagora pinnata Harvey 

FIGURES 96d, 98 

Liagora pinnata Harvey, 1853:138.—B0rgesen, 1915:74, figs. 

76-81. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6941 (•?), JN-6942 (<?), and JN6943 {<?); sta 8, 29 Mar 1976, JN-6163 («?), JN6164 {<$), and JN-6213a,b (<?); sta 9, 29 Apr 1977, 

JN-6826 (<?). 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Virgin Islands, Guadeloupe, Venezuela (Taylor, 

1960); Cuba (Suarez, 1973); Grand Cayman 

(Taylor, 1969); Puerto Rico (Diaz-Piferrer, 1963), 

Curacao (Diaz-Piferrer, 1964b). 

REMARKS.—It is a widely distributed tropical 

species having been reported in the Pacific from 

China, Japan, and Hawaii (Abbott, 1945). 

Liagora valida Harvey 

FIGURE 96C 

Liagora valida Harvey, 1853:138.—Taylor, 1960:327, pl. 43: 

fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

28 Apr 1977, JN-6399 (6*); sta 7, 22 apr 1977, JN6939 ($), JN-6940 (<?), JN-6945 {6); sta 9, 29 Apr 

1977, JN-6828 (?); sta 10, 23 Apr 1977, JN-6904 

(?)• 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Puerto Rico, Virgin Islands, Guadeloupe, Barbados, Belize, Panama (Taylor, 1960); 

St. Kitts, Dominica, St. Vincent (Taylor, 1969); 

Curacao (Diaz-Piferrer, 1964b); Costa Rica 

(Dawson, 1962b); Isla San Andres (Kapraun, 

1972). 

REMARKS.—This is a wide ranging species, 

known from warmer waters of the Atlantic and 

Pacific (Abbott, 1945). Some of our Belize specimens, for instance, JN-6945 (<3), contained "monosporangial discs" among the assimilatory filaments as described by Howe (1920a) and B0rgesen (1920). These "discs," of unknown function, 

would make an interesting culture investigation. 

* Liagora species 

FIGURE 96b 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

29 Mar 1976, JN-6l76a ($), b (<$), JN-6216 (<3), 

JN-6217 ($), and 4 Apr 1976, JN-6300 ($), JN6301a ($), b (d); sta 9, 29 Apr 1977, JN-6827. 
NUMBER 12 191 

FIGURE 98.—Liagora pinnata (JN-6213^1), details of reproduction: a, carpogonial branch (cb) 

with trichogyne (tr; note spermatia near tip) borne laterally off an assimilatory filament, and 

showing developing involucral filaments (if); b, assimilatory filament bearing lateral spermatangia; c, mature cystocarp. 
192 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

REMARKS.—The Carrie Bow Cay specimens 

differ in habit from other species of Liagora reported from the Caribbean. The carpogonial 

branches are formed by the modification of one 

of the furcations of the assimilatory filaments. 

They seem to represent an undescribed species; 

however, further material is needed to evaluate 

their taxonomic status. 

* Trichogloeopsis species 

FIGURE 99 

Trichogloeopsis cf. pedicellata (Howe) Abbott and Doty, 1960: 

632-640, figs. 1-23. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6953 ($), JN-6954d (2); sta 8, 29 

Mar 1976, JN-6167 (9); sta 19, 25 Apr 1977, JN7023 ($), JN- 7024a (9), JN-7025 ($), and JN-7026 

REMARKS.—Collections from Carrie Bow Cay 

represent the first record of the genus Trichogloeopsis in Belize. Trichogloeopsis Abbott and Doty 

(1960) is distinguished from other genera of the 

Helminthocladiaceae by the production of sterile 

rhizoids, "gonimorhizoids," by the gonimoblast 

(Figure 99b). The Belizean material resembles T 

pedicellata (Howe) Abbott and Doty (1960) in 

internal measurements and in being monecious. 

10pm 

FIGURE 99.— Trichogloeopsis species (JN-6953), details of reproductive structures: a, carpogonial 

branch (cb) with trichogyne (tr) arising as a lateral (in the place of a vegetative branch) at the 

dichotomous branching; b, mature cystocarp with well-developed gonimoblastic rhizoids (gr). 
NUMBER 12 193 

The Belizean plants, however, differ in several 

respects: the assimilatory filaments are shorter in 

length; the cystocarps on the same thallus have 

both elongated (= "pedicellate" of Abbott and 

Doty, 1960) and non-elongated primary cells of 

the carpogonial branch; and the gonimoblastic 

rhizoids are well developed on mature cystocarps 

in contrast to the weakly developed ones of T. 

pedicellata (Abbott and Doty, 1960, fig. 20). A 

detailed study of variation in T. pedicellata and 

the Belize Trichogloeopsis species is needed before 

we can determine whether the Belize specimens 

are a new species or if they represent variation 

that could be included within T. pedicellata. The 

latter was originally described from the Bahamas 

as Liagora pedicellata Howe (1920b), and reported 

in Florida, Caicos Islands, Jamaica, Guadeloupe 

(Taylor, 1960), Barbados (Taylor, 1969), and 

Curasao (van den Hoek, 1969). 

* Galaxaura lapidescens (Ellis and Solander) 

Lamouroux 

Galaxaura lapidescens (Ellis and Solander) Lamouroux, 1816: 

264.—B0rgesen, 1916:95, figs. 102-104. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7013; sta 7, 22 Apr 1977, JN6946. South Water Cay: sta 21, 28 Apr 1977, JN6723; sta 23, 20 Apr 1979, JN-7323 and JN-7332. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Puerto Rico, Virgin Islands, 

Guadeloupe, Martinique, Barbados, Grenada, 

Old Providence Island, Panama, Colombia (Taylor, 1960); Dominican Republic (Almodovar and 

Bonnelly, 1977); St. Lucia (Taylor, 1969); Curacao (Diaz-Piferrer, 1964b); Costa Rica (Dawson, 

1962b); Venezuela (Diaz-Piferrer, 1970b). 

Family CHAETANGIACEAE 

* Galaxaura cylindrica (Solander) Kjellman 

Galaxaura cylindrica (Solander) Kjellman, 1900:64.—Taylor, 

1960:341, pl. 44: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

22 Apr 1979, JN-7404 and JN-7408. 

CARIBBEAN DISTRIBUTION.—Cuba, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Martinique, Barbados, 

Grenada, Costa Rica, Panama, Venezuela (Taylor, 1960); St. Kitts, Nevis, Dominica, St. Lucia 

(Taylor, 1969); Curacao (Diaz-Piferrer, 1964b). 

REMARKS.—The Carrie Bow Cay specimens are 

of larger diameter than is "typical" of the species, 

but they agree with this species in other respects. 

The branch diameters of Galaxaura cylindrica and 

G. oblongata (Ellis and Solander) Lamouroux overlap (Chou, 1947; Taylor, 1960). Chou (1947) 

suggested the two may prove to be one species, 

but Taylor (1960), while acknowledging that 

"branch diameters converge," stated there is little 

difficulty in distinguishing them. More specimens 

must be assembled and studied to evaluate their 

distinctness. 

* Galaxaura oblongata (Ellis and Solander) 

Lamouroux 

Galaxaura oblongata (Ellis and Solander) Lamouroux, 1816: 

262.—B0rgesen, 1927:71, figs. 39-41. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977, JN-7012, and 30 Apr 1977, JN7342, and 30 Mar 1976, JN-7255; sta 7, 22 Apr 

1977, JN-6948; sta 9, 29 Apr 1977, JN-6834; sta 

10, 25 Apr 1977, JN-6973, and 23 Apr 1977, JN6903, and 28 Apr 1979, JN-7565; sta 18, 25 Apr 

1977, JN-6744; sta 19, 28 Apr 1979, JN-7620. 

South Water Cay: sta 21, 29 Apr 1977, JN-6382; 

sta 23, 30 Apr 1979, JN-7330. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Puerto Rico, 

Virgin Islands, Guadeloupe, Martinique, Barbados, Costa Rica, Panama, Trinidad (Taylor, 

1960); St. Kitts, Antigua, Dominica, St. Lucia, 

Bequia (Taylor, 1969); Curasao (Diaz-Piferrer, 

1964a). 

REMARKS.—It is difficult to separate this species 

from Galaxaura cylindrica (see notes under that 

taxon), except that its branches are more than 1 

mm in diameter (Taylor, 1960). 
194 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

* Galaxaura obtusata 

(Ellis and Solander) Lamouroux 

Galaxaura obtusata (Ellis and Solander) Lamouroux, 1816: 

262.—Taylor, 1960: 342, pl. 44: figs. 4, 5, pl. 45: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

27 Apr 1977, JN-6678, and 27 Apr 1979, JN7600. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, St. Eustatius, 

Guadeloupe, Barbados, Tobago (Taylor, 1960); 

Nevis, St. Lucia (Taylor, 1969); Curacao (DiazPiferrer, 1964b); Panama (Earle, 1972); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—At Carrie Bow Cay, it has been 

found only in deep-water (18-37 m) habitats. 

Family BONNEMAISONIACEAE 

* Asparagopsis taxiformis (Delile) Trevisan 

"Falkenbergia hillebrandii" stage of Asparagopsis taxiformis (Delile) Trevisan, 1845:45.—B0rgesen, 1918:332, figs. 332, 

333 [as Falkenbergia hillebrandii].—Taylor, 1960:571, pl. 72: 

fig. 8 [as Falkenbergia hillebrandii]. 

LOCAL DISTRIBUTION.—Twin Cays: sta 26, 29 

Apr 1979, JN-7451. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cayman Islands, Virgin Islands, Guadeloupe, 

Martinique, Barbados (Taylor, 1960); Cuba 

(Suarez, 1973); Puerto Rico (Diaz-Piferrer, 1963); 

Jamaica (Chapman, 1963); Antigua, Dominica 

(Taylor, 1969); Curasao (Diaz-Piferrer, 1964b); 

Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—Only the sporophytic stage, "Falkenbergia hillebrandii,," was encountered at Twin 

Cays. This is a new Belizean record of the genus. 

Family GELIDIACEAE 

Gelidiella acerosa (Forsskal) Feldmann 

and Hamel 

Gelidiella acerosa (Forsskal) Feldmann and Hamel, 1934: 

533.—Taylor, 1960:351, pl. 46: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6947. South Water Cay: sta 21, 

entangled with Sargassum holdfasts, 28 Apr 1977, 

JN-6721, and 29 Apr 1977, JN-6380; sta 23, 30 

Apr 1979, JN-7322. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Salt Key Bank, Cuba, Cayman Islands, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

Guadeloupe, Martinique, Barbados, Grenadines, 

Grenada, Belize, Costa Rica, Panama, Netherlands Antilles, Venezuela, Trinidad, Tobago 

(Taylor, 1960); Nevis, Antigua, Dominica, St. 

Lucia, Bequia (Taylor, 1969). 

Order CRYPTONEMIALES 

Family DUMONTIACEAE 

* Dudresnaya crassa Howe 

Dudresnaya crassa Howe, 1905:572.—Taylor, 1960:364, pl. 43: 

fig. 6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

29 Mar 1976, JN-6168 {9,6); sta 10, 25 Apr 1977, 

JN-7054 {2,6); sta 19, 25 Apr 1977, JN-7124. 

CARIBBEAN DISTRIBUTION.—Puerto Rico (Almodovar, 1970). 

REMARKS.—Numerous specimens of this soft 

gelatinous species, representing a new Belizean 

record for the genus, were encountered subtidally 

at Carrie Bow Cay. Dudresnaya crassa, described 

from Bermuda by Howe (1905), has been reported to be dioecious (Taylor, 1950), but the 

Belizean plant is monoecious. The female and 

male reproductive structures and other morphological features of the Belizean specimens, however, agree with those described for D. crassa by 

Taylor (1950; 1960). One other species, D. verticillata, has also been reported to be both dioecious 

and monoecious (Taylor, 1950). 

Family RHIZOPHYLLIDACEAE 

Ochtodes secundiramea (Montagne) Howe 

Ochtodes secundiramea (Montagne) Howe 1920b: 583.—Joly 

and Ugadim, 1966:55, pl. 1: figs. 1-8, pl. 2: figs. 1-7, pl. 

3: figs. 1-4. 
NUMBER 12 195 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, 28 Apr 1977, coll. JN, KB, W. Fenical, and 

O. McConnell, JN-6714, and 29 Apr 1977, JN6895 ($). 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, St. Barthelemy, Guadeloupe, Martinique, Barbados, 

Grenada, Belize, Costa Rica, Panama, Trinidad, 

Tobago (Taylor, 1960); St. Kitts, Dominica, St. 

Lucia, Bequia (Taylor, 1969); Curacao (Diaz-Piferrer, 1964b); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—Many of the specimens were cystocarpic, and the cystocarps were clustered in 

pale wart-like nemathecia, reminiscent of parasitic algae. In fresh plants, numerous unicellular 

hairs issuing from the cortex of young portions 

were observed, as described by Joly and Ugadim 

(1966). Once preserved in 3% Formalin/seawater, 

these hairs become detached or disintegrate. The 

"gland cells" of fresh plants, viewed under a Wild 

M • 5 stereo microscope, were densely interspersed 

throughout the cortex, and appeared iridescent 

blue against the red pigmented cortical cells. 

After methanol extraction, the "gland cells" apparently lost their contents and no longer were 

iridescent blue. McConnell and Fenical (1978) 

discovered halogenated compounds in this plant, 

one of which, ochotodene, was up to 50% of the 

crude organic extract. These "gland cells" are the 

probable site of halogenation storage as they have 

been discovered to be in another red alga, Laurencia snyderae (Young et al., 1980). 

Family KALLYMENIACEAE 

* Kallymenia limminghii Montagne 

Kallymenia limminghii Montagne, 1860:173.—Taylor, 1960: 

432, pl. 80: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 12, 

29 Apr 1977, coll. M. Danty, JN-7105; sta 13, 23 

Apr 1979, JN-7701, JN-7702, JN-7703; sta 15, 27 

Apr 1979JN-7596; sta 18, 28 Apr 1979, JN-7622. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Guadeloupe (Taylor, 1969); Puerto Rico 

(Almodovar, 1970). 

REMARKS.—Our material was found only epizooic on dead gorgonians, and is the first record 

of the genus in Belize. 

Famil y CORALLINACEAE 

Articulate Corallinaceae 

* Amphiroa fragilissima (Linnaeus) Lamouroux 

Amphiroa fragilissima (Linnaeus) Lamouroux, 1816:298.— 

Taylor, 1960: 403, pl. 47: figs. 1, 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

23 Apr 1979, JN-767la; sta 19, 25 Apr 1977, JN7019; sta 20, 22 Apr 1979, JN-7745. Twin Cays: 

sta 25, epiphytic on Halimeda, 26 Apr 1977, JN6873; sta 26, 29 Apr 1979, JN-7311, JN-7534. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

St. Barthelemy, Guadeloupe, Dominica, Martinique, Barbados, Grenadines, Grenada, Belize, 

Costa Rica, Panama, Colombia, Netherlands Antilles, Venezuela, Tobago (Taylor, 1960); Grand 

Cayman, Nevis, Antigua, St. Lucia, Bequia (Taylor, 1969); Trinidad (Richardson, 1975). 

* Amphiroa rigida var. antillana B0rgesen 

Amphiroa rigida var. antillana B0rgesen, 1917:182.—Taylor, 

1960:404, pl. 47: fig. 3, pl. 48: fig. 1. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1979, JN-7752; sta 10, mixed with A. 

tribulus, 28 Apr 1979, JN-7567. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Hispaniola, Virgin Islands, Guadeloupe, Barbados, Belize, Panama (Taylor, 1960); Cuba (DiazPiferrer, 1964a); Puerto Rico (Almodovar, 1964); 

Antigua, Dominica (Taylor, 1969); Costa Rica 

(Dawson, 1962b); Curacao (van den Hoek, 1969); 

Venezuela (Taylor, 1976). 

* Amphiroa tribulus (Ellis and Solander) 

Lamouroux 

Amphiroa tribulus (Ellis and Solander) Lamouroux, 1816: 

302.—Taylor, 1960:406, pl. 47: figs. 4, 5. 
196 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1979, JN-7754; sta 6, 23 Apr 1979, JN7682; sta 9, 29 Apr 1977, JN-6835; sta 14, 5 Apr 

1976, coll. JN and W. Fenical, s.n.; sta 19, 28 Apr 

1979,JN-7625. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cuba, Jamaica, Hispaniola, Puerto Rico, Guadeloupe, Martinique, Belize, Panama (Taylor, 

1960); Nevis (Taylor, 1969); Costa Rica (Dawson, 

1962b); Trinidad (Richardson, 1975). 

* Jania capillacea Harvey 

Jania capillacea Harvey, 1853:84.—Taylor, 1960:412, pl. 49: 

fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

epiphytic on Turbinaria, 30 Mar 1976, JN-7268; 

sta 6, 23 Apr 1979, JN-7655b, JN-7672a; sta 7, 

21 Apr 1977, JN-7129a, and 23 Apr 1977, JN6993; sta 9, 23 Apr 1977, JN-6929; sta 10, 25 Apr 

1977,JN-7218b. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, Barbados, 

Belize, Costa Rica, Panama, Colombia (Taylor, 

1960); Puerto Rico (Diaz-Piferrer, 1963); Grand 

Cayman, St. Kitts, Antigua, Dominica, St. Lucia, 

Grenada (Taylor, 1969); Curagao (Diaz-Piferrer, 

1964b); Venezuela (Diaz-Piferrer, 1970b). 

Non-articulate Corallinaceaef 

* Fosliella farinosa (Lamouroux) Howe 

Fosliella farinosa (Lamouroux) Howe, 1920b:587.—Masaki 

and Tokida, 1960:39, pl. 1: figs. 4-5, pl. 2: figs. 8-12, pis. 

6-7.—Ugadim, 175:119, pl. 3: figs. 1-2 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, epiphytic on Laurencia, 29 Apr 1977, JN6381b. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

f Contributed by Roberta A. Townsend, School of Biological Sciences, University of Sydney, Sydney 2006, New 

South Wales, Australia (work completed while author was 

on a Smithsonian Institution Predoctoral Fellowship, National Museum of Natural History, Washington, 

D.C. 20560). 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

Guadeloupe, Barbados, Belize, Old Providence 

Island, Costa Rica, Panama, Colombia, Venezuela, Netherlands Antilles, Tobago (Taylor, 1960); 

St. Kitts, Antigua, St. Lucia, Dominica, Bequia 

(Taylor, 1969). 

* Fosliella species 1 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, epiphytic on Laurencia, 29 Apr 1977, JN-6381c. 

REMARKS.—This epiphyte has cells (7-15 /xm 

long) and tetrasporangia (30-42 /xm long) that 

are much smaller than those of Fosliella farinosa, 

which has larger cells (12-30 /xm long) and tetrasporangia (40-90 /xm long). Therefore it represents a different species. See generic comments 

below. 

* Fosliella species 2 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

epilithic on coral rubble, 23 Apr 1979, JN-7668 

(Z). 

REMARKS.—Before species names can be confidently placed on uniporate, thin crusted nonarticulate corallines, the problems with the circumscription of the genera Fosliella and Heteroderma must be resolved. At this time, it seems best 

to refer to the two unknown Belizean species of 

Fosliella as F. species 1 and F. species 2. 

* Lithoporella atlantica (Foslie) Foslie 

Lithoporella atlantica (Foslie) Foslie, 1909:59; 1929, pl. 73: fig. 

9.—Taylor, 1960: 387 [as Fosliella atlantica]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

epilithic on coral rubble, 28 Apr 1979, coll. J. 

Norris and I. Macintyre, JN-7512 (Z). 

CARIBBEAN DISTRIBUTION.—Virgin Islands, Hispaniola (Taylor, 1960). 

REMARKS.—From the Caribbean collections of 

W. H. Adey (Coralline Herbarium, US), this 

species appears in few localities and where found, 

is apparently not abundant. It is a new Belizean 

record for the genus. 
NUMBER 12 197 

* Hydrolithon boergesenii (Foslie) Foslie 

Hydrolithon boergesenii (Foslie) Foslie, 1909:56.—Taylor, 1960: 

397, pl. 76: fig. 2 [as Goniolithon boergesenii]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

epilithic on coral rubble, 23 Apr 1979, JN-7669 

and JN-7670a, and 10 May 1978, coll. K Riitzler, 

s.n. 

CARIBBEAN DISTRIBUTION.—Dominican Republic, Virgin Islands, Barbados, Costa Rica (Taylor, 

1969); Nicaragua (Dawson, 1962b). 

REMARKS.—In the Caribbean, this species is 

common throughout the intertidal zone and may 

be found to a depth of 67 m. This is the first 

report of the genus in Belize. 

* Mesophyllum species 1 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 14, 

on coral rubble, 27 Apr 1979, JN-7589. 

REMARKS.—Unfortunately this specimen is not 

reproductive; however, it has a leafy habit, thick 

coaxial hypothallium, "large-celled" meristem 

(cf. Adey and Macintyre, 1973), and rounded 

epithallial cells. It also lacks heterocysts; therefore 

I place it into the genus Mesophyllum. A new 

record of the genus in Belize, this specimen represents a new species that is under study by W. 

H. Adey (pers. comm.). 

seaward of the IMSWE laboratory. A new generic 

record for Belize, this is apparently the first record 

of this taxon from the Central America coast of 

the Caribbean. 

* Neogoniolithon species 1 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

23 Apr 1979,JN-7670b (Z). 

REMARKS.—This specimen is regularly 

branched, and grows sympatrically with Hydrolithon boergesenii on the same coral rubble. It possesses a coaxial hypothallium, single heterocysts, 

and conceptacles of 750 /xm outside diameter. 

Neogoniolithon species 1 and 2 are under study 

by W. H. Adey and represent undescribed species 

(pers. comm.) 

* Neogoniolithon species 2 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

on coral surface, 28 Apr 1979, coll. J. Norris and 

I. Macintyre, JN-7499. 

REMARKS.—A non-branched Neogoniolithon, 

with a parallel hypothallium that is coaxial in 

places. The conceptacles range from 250 to 500 

/xm outside diameter. See also comments on N. 

species 1 above. 

* Neogoniolithon strictum (Foslie) Setchell and 

Mason 

Neogoniolithon strictum (Foslie) Setchell and Mason, 1943:92.— 

Taylor, 1960:399, pl. 48: fig. 4, pl. 77: figs. 1-2 [as 

Goniolithon strictum]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

28 Apr 1979, coll. J. Norris and I. Macintyre, JN7501, and 23 Jun 1977, coll. K. Riitzler, CBC77.6.23. 

CARIBBEAN DISTRIBUTION.—Cuba, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands (Taylor, 

1960); Curacao (van den Hoek, 1969). 

REMARKS.—This material has the sloughing 

epithallium characteristic of Neogoniolithon strictum. A rather common species at Carrie Bow Cay, 

it is free living among Thalassia on the reef flat 

* Porolithon pachydermum 

(Weber-van Bosse and Foslie) Foslie 

Porolithon pachydermum (Weber-van Bosse and Foslie in Foslie) 

Foslie, 1909:57; 1929, pl. 67: figs. 9-10.—Taylor, 1960: 

401. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

on coral face, 28 Apr 1979, coll. J. Norris and I. 

Macintyre, JN-7498 (Z). 

CARIBBEAN DISTRIBUTION.—Jamaica, Puerto 

Rico, Virgin Islands, Barbados (Taylor, 1960); 

Curasao (van den Hoek, 1969). 

REMARKS.—This crust has the definite heterocyst fields found in the type of Porolithon pachydermum (isotype number FT-121, Coralline Herbarium, US). This species is common in shallow 

waters to depths of 20 m on Caribbean outer reefs 
198 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

(W. H. Adey, pers. comm.), and represents a 

genus new to Belize. 

Order GIGARTINALES 

up to 1.3 cm in thickness. It was growing on the 

reef crest in shallow water, attached to the underside of a large piece of dead Acropora palmata 

coral in an area of strong swells and surge. 

Family GRACILARIACEAE 

* Gracilaria crassissima P. and H. Crouan 

FIGURE 100 

Gracilaria crassissima P. and H. Crouan in Maze and 

Schramm, 1878:218.—Taylor, 1960: 443, pl. 55: fig. 4, pl. 

57: fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977,JN-6996. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Hispaniola, Puerto Rico, Guadeloupe (Taylor, 

1960); Cuba (Diaz-Piferrer, 1964a); St. Lucia, 

Grenada (Taylor, 1969); Barbados (Almodovar 

and Pagan, 1967); Curacao (Diaz-Piferrer, 

1964b); Costa Rica (Dawson, 1962b); Panama 

(Earle, 1972). 

REMARKS.—The fresh specimen measured 14 

cm in diameter, approximately 8 cm in height, 

and up to 5 cm in width, individual fronds were 

5 mm 

FIGURE 100.—Gracilaria crassissima, habit of a portion of a 

robust specimen (JN-6996). 

Family SOLIERIACEAE 

* Eucheuma? echinocarpum Areschoug 

Eucheuma echinocarpum Areschoug, 1854:349 [= reprint p. 

23].—Taylor, 1960:458. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977,JN-6960 (Z). 

CARIBBEAN DISTRIBUTION.—Cuba (Diaz-Piferrer, 1964a; Suarez, 1973); Jamaica (Collins, 1901; 

Chapman, 1963); Dominican Republic (Almodovar and Bonnelly, 1977); Puerto Rico (Almodovar, 1964); Guadeloupe (Taylor, 1960); Nevis, 

Dominica (Taylor, 1969); Barbados (Almodovar 

and Pagan, 1967); Curacao (van den Hoek, 

1969); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—The correct generic placement of 

this taxon is currently under study by D. Cheney 

(pers. comm.). 

* Eucheuma? cf. schrammii (P. and H. Crouan 

J. Agardh 

FIGURE 101 

Eucheuma schrammii (P. and H. Crouan in Schramm and 

Maze) J. Agardh, 1892:124.—Taylor, 1960:458.—DiazPiferrer, 1970a: 12, fig. 6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-7106 (9). 

CARIBBEAN DISTRIBUTION.—Virgin Islands, 

Guadeloupe (Taylor, 1960); Puerto Rico (Almodovar, 1970); Dominica (Taylor, 1969); Curasao 

(Diaz-Piferrer, 1964b); Costa Rica (Dawson, 

1962b); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—In habit, the Carrie Bow Cay plant 

resembles both Eucheuma schrammii and Meristotheca tobagensis Taylor (1962b: 55). Examination of 

mature cystocarps from the type of M. tobagensis 

(Taylor, 1962b, pl. 4: fig. 1; holotype number 

US-33935) revealed a structure like that described 

and illustrated for the type of the genus, M. 
NUMBER 12 199 

5 mm 1 mm 

FIGURE 101.—Eucheuma? cf. schrammii (JN-7106): a, habit of a portion of the thallus showing 

spiny nature of mature cystocarps; b, cross section through a mature cystocarp showing 

parenchymatous central tissue (pet), and the gonimoblast and carposporangia (c) developing 

outward towards the "Faserhiille'' (F). 

papulosa (Montagne) J. Agardh, by B0rgesen 

(1934b: 13-16, fig. 11). Mature cystocarps of both 

M. papulosa and M. tobagensis have gonimoblast 

tissue and carposporangia developing outward 

from the parenchymatous central tissue. Similarly, in cystocarps of the Carrie Bow Cay specimen, the gonimoblast and carposporangia appear 

to develop outwards towards the "Faserhiille" 

from the parenchymatous central tissue (Figure 

10lo). Therefore, we suggest that the taxon E. 

schrammii is probably a Meristotheca. However, until the type of Mychodea schrammii is re-examined 

to determine if its cystocarpic construction and 

carposporangial development is the same as our 

Belizean plant, we tentatively identify the latter 

as E.? cf. schrammii. 

Recently, Cheney and Dawes (in press) examined the "Originalexemplar" of Eucheuma schrammii selected by Kylin (1932, pl. 10: fig. 23) {Mychodea schrammii P. and H. Crouan in Schramm 

and Maze 1865:10; Herbarium Agardh number 

34680, LD) and concluded that this taxon is not 

a Eucheuma (a genus in which the mature cystocarps possess a single fusion cell). The generic 

placement of M. schrammii is under study by D. 

Cheney (pers. comm.). 

Taylor (1960:458) has described the cystocarps 

of Eucheuma schrammii as "hardly projecting from 

the surface, but the overlaying tissue often apiculate." Cystocarps of the Belizean plant differ in 

being confined to marginal portions of the blade; 

they also are very spiny (Figure 101a). More 

Caribbean material must be studied, particularly 

from the type locality in Guadeloupe, to assess 

the importance of these features for taxonomic 

purposes. 

Family CAULACANTHACEAE 

Guiry (1978:193) has recently pointed out that 

the family name Caulacanthaceae Kiitzing 

(1843) has nomenclatural priority over Rhabdoniaceae Kylin (1925). 

Catenella repens (Lightfoot) Batters 

Catenella repens (Lightfoot) Batters, 1902:69.—Taylor, 1960: 

462, pl. 66: fig. 13. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6293c and JN-6295, and 26 Apr 

1977,JN-6872. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 
200 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Jamaica, Virgin Islands, Tortola Island, Guadeloupe, Martinique, Belize, Panama (Taylor, 

1960); Puerto Rico (Almodovar, 1965); Curacao, 

Bonaire (Diaz-Piferrer, 1964b); Venezuela (DiazPiferrer, 1970b); Trinidad (Richardson, 1975). 

REMARKS.—This species, a new Belizean record 

for the genus, was found in association with Bostrychia, growing on mangrove roots; its common 

habitat in the Caribbean. 

Family HYPNEACEAE 

Hypnea cervicornis J. Agardh 

Hypnea cervicornis J. Agardh, 1852:451.—Taylor, 1960:466, 

pl. 73: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

entangled with Laurencia, 28 Apr 1977, JN-6395 

(9); sta 7, 21 Apr 1977, JN-6983. Twin Cays: sta 

24, 31 Mar 1976, JN-6299 ($); sta 25, 26 Apr 

1977,JN-6893. 

CARIBBEAN DISTRIBUTION.—Mexico, Cuba, 

Cayman Islands, Jamaica, Hispaniola, Virgin Islands, St. Barthelemy, Guadeloupe, Martinique, 

Belize, Panama, Colombia, Netherlands Antilles, 

Venezuela (Taylor, 1960); Puerto Rico (Diaz-Piferrer, 1963); Antigua, Barbados, Grenada (Taylor, 1969); Costa Rica (Dawson, 1962b); Trinidad 

(Richardson, 1975). 

REMARKS.—Bispores, 15-23 /xm X 45-52 /xm, 

borne in specialized branches were found on JN6299; the usual mode of sporophyte reproduction 

is by zonate tetraspores. 

Hypnea musciformis (Wulfen) Lamouroux 

Hypnea musciformis (Wulfen) Lamouroux, 1813:131.—Taylor, 

1960:467, pl. 73: fig. 1. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6271. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Martinique, Barbados, 

Grenada, Belize, Costa Rica, Panama, Colombia, 

Netherlands Antilles, Venezuela, Trinidad, Tobago (Taylor, 1960); St. Kitts, Nevis, Antigua, 

Dominica, St. Lucia, St. Vincent, Bequia (Taylor, 

1969); Isla San Andres (Kapraun, 1972). 

Order RHODYMENIALES 

Family RHODYMENIACEAE 

* Botryocladia shanksii Dawson 

Botryocladia shanksii Dawson, 1962b:385, pl. 2: figs, A, B, pl. 5: 

fig. B.—Schnetter, 1977:79, figs. 8-11. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

20 Apr 1977,JN-7107. 

CARIBBEAN DISTRIBUTION.—Barbados, Costa 

Rica (Dawson, 1962b); Colombia (Schnetter, 

1977). 

Family CHAMPIACEAE 

Champia parvula ( C Agardh) Harvey 

Champia parvula (C. Agardh) Harvey, 1853:76.—Taylor, 

1960:490, pl. 61: fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

epiphytic on Udotea, 28 Apr 1977, JN-6400; sta 6, 

epiphytic on Thalassia, 23 Apr 1979, JN-7651; sta 

8, epiphytic on Halimeda, 27 Apr 1977, JN-6420 

(0), JN-6435 (9); sta 9, 29 Apr 1977, JN-6846a; 

sta 10, 23 Apr 1977, JN-6922; sta 11, epiphytic 

on Sargassum, 29 Mar 1976, JN-6175; sta 14, 

epiphytic on Sargassum hystrix, 5 Apr 1976, JN6350 (0) and epiphytic on Dictyota dichotoma, JN6340 (0), and 27 Apr 1977, JN-6804; sta 15, 

epiphytic on Sargassum blade, 27 Apr 1977, JN6689 (9); sta 18, 25 Apr 1977, JN-6752 (9,0). 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Jamaica, Hispaniola, Puerto Rico, Virgin 

Islands, Guadeloupe, Dominica, Barbados, Venezuela (Taylor, 1960); Cuba (Suarez, 1973); 

Nevis, Antigua (Taylor, 1969); Belize (Tsuda and 

Dawes, 1974); Costa Rica (Wellington, 1973). 

REMARKS.—In 1976 and 1977 Champia parvula 

was found at several stations in the vicinity of 

Carrie Bow Cay, but it was always sparse in 

distribution and coverage. In the spring of 1979, 
NUMBER 12 201 

however, this species appeared in great abundance as an epiphyte on Thalassia at station 6, a 

dominant yellow coverage seen over the entire 

reef flat (compare Riitzler and Macintyre, herein: 

15, Figures 6b, lb). 

Coelothrix irregularis (Harvey) B0rgesen 

Coelothrix irregularis (Harvey) B0rgesen, 1920:389.—Taylor, 

1960:488, pl. 45: fig. 3, pl. 46: fig. 4. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 6, 

23 Apr 1979, JN-7658 and JN-7666. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Puerto Rico, Virgin Islands, 

Guadeloupe, Belize (Taylor, 1960); Grand Cayman, Antigua, Dominica, St. Lucia, Barbados 

(Taylor, 1969); Curacao (Diaz-Piferrer, 1964b); 

Trinidad (Richardson, 1975). 

* Chrysymenia cf. okamurai Yamada and 

Segawa 

Chrysymenia okamurai Yamada and Segawa, 1953:110, fig. 3. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

27 Apr 1977,JN-6428. 

REMARKS.—This species represents a genus not 

previously recorded in Belize, and is apparently 

not known elsewhere in the Caribbean. The plant 

resembles Chrysymenia okamurai Yamada and Segawa (1953:110, fig. 3; see also Abbott and Littler, 1969) from Japan and Hawaii in general 

morphology. More specimens, however, particularly cystocarpic ones, are needed before the identity of the Belize alga can be established. 

Order CERAMIALES 

Family CERAMIACEAE 

Antithamnion ogdeniae Abbott 

Antithamnion ogdeniae Abbott, 1979:218, figs. 11-14. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

epiphytic on Halimeda, 27 Apr 1977, JN-6429 and 

JN-6430; sta 14, epiphytic on Sargassum, 5 Apr 

1976,JN-6347. 

CARIBBEAN DISTRIBUTION.—This species was recently described from St. Croix, U.S. Virgin Islands, and Belize, with specimens (paratypes) 

from Carrie Bow Cay (Abbott, 1979). 

* Antithamnionella flagellata (B0rgesen) Abbott 

Antithamnionella flagellata (B0rgesen) Abbott, 

1979:222, figs. 18-20 [as

 lflagellatum,

].—B0rgesen, 

1945:5, figs. 1, 2 [as Antithamnion flagellatum]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

epiphytic, 27 Apr 1977, JN-6427 (0); sta 9, epiphytic, 29 Apr 1977, JN-6841d. 

CARIBBEAN DISTRIBUTION.—Virgin Islands (Abbott, 1979). 

REMARKS.—This species was originally described from Mauritius (B0rgesen, 1945) and only 

recently found near St. Croix, U.S. Virgin Islands 

(Abbott, 1979). Belizean specimens (the first record of the genus in Belize), while in general 

agreement with Antithamnionella flagellata, have 

branchlets reaching only 300 /xm in length, 

shorter than the 300-600 /xm length previously 

described. 

Centroceras clavulatum (C. Agardh) Montagne 

FIGURE 102 

Centroceras clavulatum (C. Agardh) Montagne, 1846:140.—Abbott and Hollenberg, 1976:604, fig. 547. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

epiphytic on Laurencia, 28 Apr 1977, JN-6394; sta 

7, entangled with turf, 23 Jun 1977, coll. K. 

Riitzler, CBC-77.6.23, and in algal turf, intermixed with Anotrichium tenue, Jania and Ceramium, 

22 Apr 1977, JN-6958b, and in algal turf, entangled with Taenioma, Herposiphonia, and Jania, JN6989a {6); sta 15, 27 Apr 1977, JN-6688a. South 

Water Cay: sta 21, on concrete block, 29 Apr 

1977, JN-6384. Twin Cays: sta 26, 22 Apr 1979, 

JN-7435. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 
202 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 102.—Centroceras clavulatum, upper portion of thallus, 

showing node with emergent tetrasporangia subtended by 

spines and involucres (JN-6384). 

Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Dominica, Martinique, 

Barbados, Grenada, Belize, Old Providence Island, Costa Rica, Panama, Colombia, Netherlands Antilles, Venezuela, Tobago (Taylor, 

1960); St. Kitts, Antigua, St. Lucia, St. Vincent, 

Bequia (Taylor, 1969); Isla San Andres (Kapraun, 1972); Trinidad (Richardson, 1975). 

REMARKS.—Centroceras clavulatum apparently 

has variable tetrasporangia as well as vegetative 

morphology. Tetrasporangia of the Belize specimens are emergent, subtended by 2-3 celled 

spines and occasionally by 4- to 5-celled involucres (Figure 102). Taylor (1960:537) describes the 

tetrasporangia as mostly exposed, generally 

formed in the whorl of spines, but does not mention involucres, while Joly (1965, pl. 39: fig. 502) 

illustrates the tetrasporangia of material from 

Brasil with involucres but without spines. 

* Ceramium brevizonatum var. caraibicum 

H. Petersen and B0rgesen 

Ceramium brevizonatum var. caraibicum H. Petersen and B0rgesen in B0rgesen, 1924:29 [as 'caraibica'].—Taylor, 1960: 

527, pl. 67: figs. 7-9. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6988 {6), and epiphytic on 

Liagora, JN-6955c (0,9). South Water Cay: sta 

21, 28 Apr 1977, JN-6732 (0). 

CARIBBEAN DISTRIBUTION.—Hispaniola (Taylor, 1960); Antigua, Barbados (Taylor, 1969). 

* Ceramium fastigiatum f. flaccidum 

H. Petersen 

Ceramium fastigiatum f. flaccidum H. Petersen in B0rgesen, 

1918:242 [as f. 'flaccida'].—Taylor, 1960:527, pl. 67: figs. 

4-6. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

5 Apr 1976, epiphytic on Dictyota dichotoma, JN6343 (9), and epiphytic on Dictyota cervicornis, JN6336, and 27 Apr 1977, JN-6688b {6) and JN6690 (0,6*). 

CARIBBEAN DISTRIBUTION.—Jamaica (Chapman, 1963); Hispaniola (Taylor and Arndt, 

1929); Virgin Islands (B0rgesen, 1918); Guadeloupe (Feldman and Lami, 1935). 

* Ceramium flaccidum (Kiitzing) Ardissone 

Ceramium flaccidum (Kiitzing) Ardissone, 1871:40.—Dawson, 

1962a:57, pl. 20: figs. 2-3, pl. 21: figs. 2-3 [as C. gracillimum 

var. byssoideum].—Womersley, 1978:234, figs. 4a-d. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

epiphytic on Udotea, 28 Apr 1977, JN-6402 (0); 

sta 7, 22 Apr 1977, epiphytic on Liagora, JN-6957 

(9); sta 8, epiphytic on Sargassum, 29 Mar 1976, 

JN-6173 (9), and epiphytic on Caulerpa, 27 Apr 

1977, JN-6415a (0) and JN-6431 (0); sta 9, 

epiphytic on Sargassum, 29 Apr 1977, JN-6843 

(0); sta 10, epiphytic on Dictyota, 23 Apr 1977, 

JN-6910 (0), JN-6912 (0), JN-7212 (0); sta 17, 

25 Apr 1977, JN-6754b (9), JN-6759a (0) and 

epiphytic on Dictyota, JN-6747 (0, 6). Twin Cays: 

sta 24, epiphytic on Stypopodium, 26 Apr 1977, JN6884 (0, 9). 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, Saba Bank, 

Old Providence Island, Colombia, Venezuela 

(Taylor, I960); Puerto Rico (Almodovar, 1964); 

Barbados (Taylor, 1969); Curacao (Diaz-Piferrer, 
NUMBER 12 203 

1964b); Costa Rica (Dawson, 1962b). 

REMARKS.—This minute species is commonly 

found on a variety of coarser algae. It is a wideranging species, found in the Pacific (Taylor, 

1945, as Ceramium byssoideum; Dawson, 1961, as C. 

gracillimum var. byssoideum), Gulf of California 

(Dawson, 1962a, as C. gracillimum var. byssoideum), 

Caribbean (Taylor, 1960, 1969, as C. byssoideum), 

Australia (Womersley, 1978), North Africa (Mazoyer, 1938, as C. gracillimum var. byssoideum), and 

Europe (Feldmann-Mazoyer, 1941, as C. gracillimum var. byssoideum). Womersley (1978) has found 

this to be a single, variable, and wide-ranging 

species reported under several names. He placed 

these taxa names in synonomy and noted that C 

flaccidum is apparently the earliest validly published name for the species. 

Ceramium nitens (C. Agardh) J. Agardh 

Ceramium nitens (C. Agardh) J. Agardh, 1851:130.—Taylor, 

1960:535, pl. 66, fig. 14. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

30 Mar 1976, JN-7262 and 25 Apr 1979, JN7728; sta 3, 28 Apr 1977, JN-6396; sta 8, growing 

on a gorgonian, 4 Apr 1976, JN-6302. South 

Water Cay: sta 21, 29 Apr 1977, JN-6383, coll. 

O. McConnell; sta 22, epiphytic on Sargassum 

stipe, 28 Apr 1977, JN-6716. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Puerto Rico, 

Virgin Islands, Guadeloupe, Martinique, Belize, 

Colombia (Taylor, 1960); Grand Cayman, St. 

Kitts, Dominica, Barbados (Taylor, 1969); Curacao (Diaz-Piferrer, 1964b); Venezuela (Taylor, 

1976). 

* Crouania attenuata (Bonnemaison) J. Agardh 

Crouania attenuata (Bonnemaison) J. Agardh, 1842:83.—B0rgesen, 1917:230, figs. 219-221. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

epiphytic on Stypopodium, 4 Apr 1976, JN-6306; 

sta 8, epiphytic on Liagora pinnata, 29 Mar 1976, 

JN-6212 (0), epiphytic on Valonia ventricosa, 27 

Apr 1977, JN-6407 {6), JN-6433 {6), and JN6436a, and epiphytic on Halimeda, JN-6421 a (9); 

sta 9, epiphytic on Sargassum blade, 29 Apr 1977, 

JN-6838a (0); sta 10, epiphytic on Galaxaura, 23 

Apr 1977, JN-6908 (0); sta 14, epiphytic on 

Halimeda, 27 Apr 1977, JN-6802 {6). 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Barbados (Taylor, 1960); Puerto Rico (Almodovar, 1964); Antigua (Taylor, 1969); Curacao (Diaz-Piferrer, 1964b); Venezuela (Diaz-Piferrer, 1970b). 

REMARKS.—This is the first report of this genus 

in Belize. 

* Griffithsia globulifera (Harvey) J. Agardh 

Gnffithsiaglobulifera (Harvey) J. Agardh, 1876:67.—B0rgesen, 

1917:202, figs. 191-193. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 2, 

on Strombus gigas (Queen conch) shell, 22 Apr 

1977, JN-7225 {6); sta 7, 22 Apr 1977, JN-6991a; 

sta 8, 27 Apr 1977, JN-6426 and JN-6432; sta 9, 

29 Apr 1977, JN-6836; sta 10, epiphytic on Galaxaura, 23 Apr 1977, JN-6909. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Virgin Islands, Guadeloupe, Barbados, Venezuela (Taylor, 1960); Jamaica 

(Chapman, 1963); Puerto Rico (Almodovar, 

1964); Curacao (van den Hoek, 1969). 

REMARKS.—The two morphologically similar 

species, Griffithsia globulifera and G. schousboei Montagne are most easily separated by their spermatangium. The Belizean plant is assigned to G. 

globulifera (JN-7225), because it lacks the involucre on the spermatangia, a feature ascribed to G. 

schousboei (Taylor, 1960). The spermatangia of the 

Carrie Bow Cay specimen are borne on terminal 

cells of the filaments as a cap, and in a ring about 

subterminal cells. Our specimens are the first 

record of the genus in Belize. 

* Anotrichium tenue (C. Agardh) Naegeli 

FIGURE 103 

Anotrichium tenue (C. Agardh) Naegeli, 1861:399.—B0rgesen, 
204 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 103.—Anotrichium tenue, upper portion of axes showing tetrasporangia borne terminally 

on pedicels (JN-6926). 

1920:462, fig. 423 [as Griffithsia tenuis].—Taylor, 1960:516 

[as Griffithsia tenuis].—Baldock, 1976:556, figs. 49-64. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

intermixed in an algal turf, with Jania and Ceramium, 22 Apr 1977, JN-6958 (0); sta 8, 27 Apr 

1977, JN-6434 (0); sta 10, epiphytic on Dictyota, 

23 Apr 1977,JN-6925 (0). 

CARIBBEAN DISTRIBUTION.—Hispaniola, Virgin 

Islands, Barbados (Taylor, 1960); Cuba (Suarez, 

1973); Jamaica (Chapman, 1963); Puerto Rico 

(Almodovar and Blomquist, 1965); St. Lucia 

(Taylor, 1969); Costa Rica (Dawson, 1962b); 

Curacao (van den Hoek, 1969). 

REMARKS.—Baldock (1976:550, figs. 60, 61) described and illustrated Australian specimens of 

Anotrichium tenue, in which the tetrasporangia were 

borne subterminally on pedicels; he suggested the 

position of the tetrasporangia be compared to 

material from other regions. Our specimens, a 

new record of the genus from Belize, bear mature 

and developing tetrasporangia terminally (Figure 

103). They also differ from Australian A. tenue by 

having fewer tetrasporangia per whorl. Belizean 
NUMBER 12 205 

plants have commonly 4-9, while specimens from 

Hong Kong (Tseng, 1942) and Australia (Baldock, 1976) have 8-10 (-12) tetrasporangia per 

whorl. Before placing taxonomic significance on 

these differences it will be necessary to study 

variation of tetrasporangial position within plant 

populations. It is also important to examine the 

tetrasporangia in the type of Griffithsia tenuis C. 

Agardh (1928) and in additional material from 

the type locality. 

* Seirospora occidentalis B0rgesen 

FIGURE 104 

Seirospora occidentals B0rgesen, 1909:14; 1917:222, figs. 209-

212. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6850a (9); sta 15, epiphytic on 

Dictyota, 5 Apr 1976, JN-6341 (9), and epiphytic 

on Sargassum, JN-6348 (seirospores), and 27 Apr 

1977, epiphytic on Sargassum blade, JN-6692 (9) 

andJN-6693b (<?). 

CARIBBEAN DISTRIBUTION.—Bahamas, Virgin 

Islands, Guadeloupe (Taylor, 1960); Puerto Rico 

(Almodovar and Blomquist, 1965). 

REMARKS.—Carposporangial, spermatangial, 

and thalli with the distinctive seirospores in 

chains (B0rgesen, 1909, 1917) were found in the 

Carrie Bow Cay specimens. They represent a new 

Belizean record for the genus. 

* Spermothamnion investiens 

(P. and H. Crouan) Vickers and Shaw 

Spermothamnion investiens (P. and H. Crouan in Maze and 

Schramm) Vickers and Shaw, 1905:64.—B0rgesen, 1917: 

200, figs. 189, 190. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6847; sta 18, epiphytic on Galaxaura, 25 Apr 1977, JN-6758a (0). 

CARIBBEAN DISTRIBUTION.—Bahamas, Virgin 

Islands, Guadeloupe, Barbados (Taylor, 1960); 

Puerto Rico (Diaz-Piferrer, 1963); Curacao 

(Diaz-Piferrer, 1964a). This is a new Belizean 

record for the genus. 

FIGURE 104.—Seirospora occidentalis, a small portion of the 

thallus showing the characteristic seirospores in chains (JN6348). 
206 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Spyridia filamentosa (Wulfen) Harvey ex 

Hooker 

Spyridia filamentosa (Wulfen) Harvey ex Hooker, 1833:337.— 

B0rgesen, 1917:233, fig. 222-226. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6274 (09), and 29 Apr 1979, JN7394 and JN-7397; sta 25, 26 Apr 1977, JN-6866 

(9); sta 26, 22 Apr 1979, JN-7439 {6). 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, St. Eustatius, Guadeloupe, Martinique, 

Barbados, Grenada, Belize, Costa Rica, Panama, 

Colombia, Netherlands Antilles, Isla las Aves 

(Taylor, 1960); Antigua, St. Lucia (Taylor, 1969); 

Venezuela (Diaz-Piferrer, 1970b); Trinidad 

(Richardson, 1975). 

Wrangelia argus Montagne 

Wrangelia argus Montagne, 1856:444.—Taylor, 1960:502, pl. 

66: figs. 7-8. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6986 (0); sta 8, 27 Apr 1977, 

JN-6419 (0); sta 9, epiphytic on Dictyota, 29 Apr 

1977, JN-6837; sta 18, 25 Apr 1977, JN-7226. 

South Water Cay: sta 21, 28 Apr 1977, JN-6731 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Jamaica, Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Barbados, Costa 

Rica, Panama, Colombia, Netherlands Antilles, 

Venezuela, Trinidad, Tobago (Taylor, 1960); 

Cuba (Suarez, 1973); Dominican Republic (Almodovar and Bonnelly, 1977); St. Kitts, Nevis, 

Dominica, St. Lucia (Taylor, 1969); Belize 

(Tsuda and Dawes, 1974). 

* Wrangelia penicillata C. Agardh 

Wrangelia penicillata C. Agardh, 1828:138.—Taylor, 1960: 

503, pl. 66: figs. 5-6, pl. 74: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

epiphytic on Liagora farinosa, 29 Mar 1976, JN6162 (0), and epiphytic on Caulerpa verticillata, 27 

Apr 1977, JN-6414 (0); sta 9, 29 Apr 1977, JN6841a {6). 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Jamaica, Virgin Islands, Guadeloupe 

(Taylor, 1960); Cuba (Diaz-Piferrer, 1964a); Dominican Republic (Almodovar and Bonnelly, 

1977); Puerto Rico (Almodovar, 1964); Grand 

Cayman, Nevis, Barbados (Taylor, 1969); Colombia (Schnetter and Bula Meyer, 1977); Curacao 

(van den Hoek, 1969); Venezuela (Diaz-Piferrer, 

1970b). 

Family DELESSERIACEAE 

Caloglossa leprieurii (Montagne) J. Agardh 

Caloglossa leprieurii (Montagne) J. Agardh, 1876:499.—Taylor, 1960: 544, pl. 68: fig. 1. 

LOCAL DISTRIBUTION.—Twin Cays: sta 26, 

growing on mangrove roots, 22 Apr 1979, JN7445. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, Guadeloupe, Martinique, Barbados, Belize, Panama, 

Colombia (Taylor, 1960); Cuba (Diaz-Piferrer, 

1964a); Grenada (Taylor, 1969); Curacao, Bonaire (Diaz-Piferrer, 1964b); Venezuela (Taylor, 

1976); Trinidad (Richardson, 1975). 

Martensia pavonia (J. Agardh) J. Agardh 

Martensia pavonia (J. Agardh) J. Agardh, 1863:831.—B0rgesen, 1919:348, figs. 344-346. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 9, 

29 Apr 1977, JN-6830; sta 11, 23 Apr 1979, JN7688; sta 14, epiphytic on Sargassum, 5 Apr 1976, 

JN-6352. 

CARIBBEAN DISTRIBUTION.—Mexico, Jamaica, 

Virgin Islands, Guadeloupe (Taylor, 1960); Cuba 

(Suarez, 1973); Puerto Rico (Almodovar, 1964); 

Antigua, Barbados (Taylor, 1969); Belize (Tsuda 

and Dawes, 1974); Venezuela (Diaz-Piferrer, 

1970b). 

* Taenioma nanum (Kiitzing) Papenfuss 

Taemoma nanum (Kiitzing) Papenfuss, 1952:179.—Joly, 1965: 

213, pl. 45: figs. 568, 569. 
NUMBER 12 207 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

in turf, 23 Jun 1977, coll. K. Riitzler, CBC77.6.23, and in turf with Centroceras and Herposiphonia, 22 Apr 1977, JN-6989b; sta 18, on Dictyota, 

25 Apr 1977,JN-6751b. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Virgin Islands (Taylor, 1960, as T macrourum); Cuba (Taylor, 1969); Curacao (van den 

Hoek, 1969); Venezuela (Taylor, 1976). 

REMARKS.—A small creeping species, it is found 

in the shallow-water turf community and represents a new generic record for Belize. Blades of 

specimen JN-6751b end mostly in two apical 

hairs, except for one that terminates in three 

hairs. All of its blades are 5 cells broad. Dawson 

(1962a:82-83), studying Taenioma perpusillum (J. 

Agardh) J. Agardh from Pacific Mexico, noted 

that their blades terminated in one, two or three 

hairs, "so that it is difficult to accept the number 

of terminal hairs as a basis for distinguishing 

these two species." More recently, Ganesan and 

Lemus (1969) observed mostly two or sometimes 

only one, but never three hairs in T. nanum from 

Venezuela. Clearly, critical studies are needed to 

evaluate the status of these two taxa. 

* Nitophyllum species 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 18, 

25 Apr 1977, JN-7170(0). 

REMARKS.—Small monostromatic blades, with 

entire margins, 1 cm tall, are tentatively referred 

to this genus. Only tetrasporangial thalli were 

found. Tetrasporangial sori are round to oblong, 

300 /xm to 1000 /xm in diameter, and located near 

the blade margins. This material may represent 

a new species but more specimens, particularly 

cystocarpic ones, are needed before its generic 

and specific placement can be determined. Similar specimens from St. Croix, U.S. Virgin Islands 

(J. L. Connor, pers. comm.) contain tetrasporic 

and cystocarpic thalli. The tetrasporangial thalli 

are undistinguishable from those from Belize. The 

cystocarps of the St. Croix specimen have carpospores borne terminally on gonimoblast filaments, 

a characteristic of the genus Nitophyllum, which 

separates it from the morphologically similar Mynogramme. 

* Hypoglossum tenuifolium var. carolinianum 

Williams 

Hypoglossum tenuifolium var. carolinianum Williams in Pearse 

and Williams, 1951:156 [as 'caroliniana'].—Williams 1948: 

693, figs. 12-17, [as//, tenuifolium].—Taylor, 1960:546. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

27 Apr 1977, JN-6418 (0), and on Valonia macrophysa, JN-6409; sta 10, 23 Apr 1977, JN-7213; sta 

14, 27 Apr 1977,JN-6801 (9). 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Jamaica, Virgin Islands, Guadeloupe, 

Netherlands Antilles, Venezuela (Taylor, 1960). 

REMARKS.—This is the first report of the genus 

in Belize. 

Family DASYACEAE 

* Dasya rigidula (Kiitzing) Ardissone 

Dasya ngidula (Kiitzing) Ardissone, 1878:140.—Taylor, 1960: 

558, pl. 72: fig. 4. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, entangled in Spyridia filamentosa, 31 Mar 1976, JN6275; sta 25, 26 Apr 1977, JN-6885 (0) and JN6874. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Cayman Islands, Jamaica, Venezuela (Taylor, 

1960); Cuba (Diaz-Piferrer, 1964a); Antigua, St. 

Lucia (Taylor, 1969); Curacao (van den Hoek, 

1969). 

REMARKS.—It represents a new Belizean record 

of the genus. 

* Dasyopsis antillarum Howe 

FIGURE 105 

Dasyopsis antillarum Howe, 1920b:577.—Taylor, 1960:564. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

27 Apr 1977, JN-6697 (0), and JN-7044 (0); sta 

14, epiphytic on Stypopodium zonale, 5 Apr 1976, 

JN-6337. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba (Taylor, 1960); Venezuela (DiazPiferrer, 1970b). 

REMARKS.—This is a new Belizean record for 

the genus. 
208 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 105.—Dasyopsis antillarum, portion of the thallus with tetrasporangial stichidia and 

some vegetative filaments (JN-7044). 

* Halodictyon mirabile Zanardini 

Halodictyon mirabile Zanardini, 1843:52.—Taylor, 1960:567, 

pl. 72: fig. 5. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

epiphytic on Stypopodium zonale, 4 Apr 1976, JN6304, and on Caulerpa verticillata, 27 Apr 1977, JN6416 and JN-6436b; sta 9, on Sargassum, 29 Mar 

1976, JN-6171, and on Dictyota, 23 Apr 1977, JN7211; sta 10, on Halimeda, 25 Apr 1977, JN-7219. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Barbados (Taylor, 1960); Curacao (van den 

Hoek, 1969). 

REMARKS.—Several nonreproductive specimens of this net-forming alga were discovered 

growing epiphytically on larger algae. These rep-
NUMBER 12 209 

resent a new Belizean record for the genus. 

* Heterosiphonia wurdemanni var. laxa 

B0rgesen 

Heterosiphonia wurdemanni var. laxa B0rgesen, 1919:326, figs. 

327, 328.—Taylor, 1960:565, pl. 72: fig. 9. 

LOCAL DISTRIBUTION.—Twin Cays: sta 26, 29 

Apr 1979, epiphytic on Caulerpa paspaloides, JN7286 (0), JN-7287 (0), JN-7288 (0), and JN7421 (0), and epiphytic on Acanthophora spicifera, 

JN-7425. 

CARIBBEAN DISTRIBUTION.—Hispaniola (Taylor 

and Arndt, 1929); Virgin Islands (B0rgesen, 

1919). 

REMARKS.—Morphologically this species is very 

similar to Dasya rigidula. However, the base of the 

monosiphonous laterals in Heterosiphonia is often 

polysiphonous, whereas the basal cell of the monosiphonous filaments in Dasya is always a single 

cell. 

Family RHODOMELACEAE 

Acanthophora spicifera (Vahl) B0rgesen 

Acanthophora spicifera (Vahl) B0rgesen, 1910:201.—Taylor, 

1960:620, pl. 71: fig. 3; pl. 72: figs. 1, 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 2, 

27 Apr 1979, JN-7636; on pebble on sandy beach, 

29 Jun 1972, coll. F.R. Fosberg-54442, and D. 

Spellman. South Water Cay: sta 23, 29 Apr 1979, 

JN-7542. Twin Cays: sta 24, 31 Mar 1976, JN6270; sta 25, 26 Apr 1977, JN-6878, and 29 Apr 

1979, JN-7583; sta 26, 29 Apr 1979, JN-7283. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, St. Barthelemy, 

Guadeloupe, Dominica, Martinique, Barbados, 

Grenada, Guatemala, Old Providence Island, 

Costa Rica, Panama, Colombia, Netherlands Antilles, Isla las Aves, Venezuela, Trinidad, Tobago 

(Taylor, 1960); Grand Cayman, Nevis, Antigua, 

St. Lucia, Bequia (Taylor, 1969); Belize (Tsuda 

and Dawes, 1974). 

REMARKS.—This plant is widely distributed 

throughout the Caribbean Sea, and the Atlantic, 

Pacific, and Indian oceans (Cordeiro-Marino, 

1978). In Belize it is common and forms dense 

expanding clumps. 

* Bostrychia binderi Harvey 

Bostrychia binderi Harvey, 1849:68.—Taylor, 1960:598.—Cordeiro-Marino, 1978:120, figs. 356-359. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 

growing on mangrove roots, 31 Mar 1976, JN6293b, and 26 Apr 1977, JN-6871 and JN-6888a. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Hispaniola, Virgin Islands, Tortola Island, Guadeloupe, Martinique, St. Vincent, Barbados, Panama, Venezuela, Trinidad (Taylor, 

1960); Puerto Rico (Diaz-Piferrer, 1963); St. 

Kitts, Antigua (Taylor, 1969); Curacao (DiazPiferrer, 1964b); Costa Rica (Dawson, 1962b). 

REMARKS.—This species is also known from 

other parts of the tropical western Atlantic, and 

from the tropical Pacific and Indian Oceans (Cordeiro-Marino, 1978). 

Bostrychia montagnei Harvey 

Bostrychia montagnei Harvey, 1853:55.—Taylor, 1960:598, pl. 

74: fig. 1. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 31 

Mar 1976, JN-6293a. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Belize (Taylor, 1960). 

Bostrychia tenella (Vahl) J. Agardh 

Bostrychia tenella (Vahl) J. Agardh, 1863:869.—Taylor, 1960: 

599.—Cordeiro-Marino, 1978:121, figs. 360-363. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, entangled with Catenella, 31 Mar 1976, JN-6279 0 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Martinique, 

Barbados, Grenada, Belize, Costa Rica, Venezuela (Taylor, 1960); Puerto Rico (Almodovar, 

1964); Antigua, Bequia (Taylor, 1969); Curasao, 

Bonaire (Diaz-Piferrer, 1964b); Trinidad (Richardson, 1975). 
210 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

* Chondria polyrhiza Collins and Hervey 

Chondria polyrhiza Collins and Hervey, 1917:121, pl. 2: fig. 

12.—Cordeiro-Marino, 1978:128, figs. 388-391. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 8, 

epiphytic on Halimeda, 29 Mar 1976, JN-6158, 

and epiphytic on Sargassum, JN-6174 (0); sta 10, 

epiphytic on Dictyota, 23 Apr 1977, JN-6027 (9); 

sta 14, epiphytic on Sargassum blade, 27 Apr 1977, 

JN-6800 (0); sta 15, epiphytic on Sargassum, 5 

Apr 1976, JN-6354. 

CARIBBEAN DISTRIBUTION.—Mexico (Humm 

and Hildebrand, 1962); Bahamas, Virgin Islands, 

(Taylor, 1960); Puerto Rico (Almodovar and 

Blomquist, 1965); Costa Rica (Dawson, 1962b); 

Curasao (van den Hoek, 1969); Venezuela (DiazPiferrer, 1970b). 

Digenia simplex (Wulfen) C. Agardh 

Digenm simplex (Wulfen) C. Agardh, 1822:389.—Taylor, 

1960:589.—Woelkerling, 1976:126, figs. 203-205. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 3, 

cast ashore, 22 Apr 1979, JN-7823. South Water 

Cay: sta 23, 30 Apr 1979, JN-7321. Twin Cays: 

sta 25, 26 April 1977, JN-6894b. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Turks Islands, Cuba, Jamaica, Hispaniola, 

Puerto Rico, Virgin Islands, St. Barthelemy, 

Guadeloupe, Martinique, Grenada, Belize, Panama (Taylor, 1960); Grand Cayman, St. Kitts, 

Nevis, Antigua, Dominica, St. Lucia, Bequia 

(Taylor, 1969); Barbados (Almodovar and Pagan, 1967); Curagao, Bonaire (Diaz-Piferrer, 

1964b); Costa Rica (Dawson, 1962b); Venezuela 

(Taylor, 1976). 

* Herposiphonia pecten-veneris (Harvey) 

Falkenberg 

Herposiphonia pecten-veneris (Harvey) Falkenberg, 1901:315.— 

Taylor, 1960:603. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 5, 

24 Apr 1977, JN-7220; sta 8, epiphytic on Udotea, 

29 Mar 1976, JN-6156 (0); sta 9, epiphytic on 

Sargassum, 29 Apr 1977, JN-6844. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Jamaica (Taylor, 1960); Cuba (Suarez, 

1973); Venezuela (Taylor, 1976). 

REMARKS.—B0rgesen (1920:470) thought this 

taxon could be a form of Herposiphonia tenella. 

Tseng (1944), while recognizing the species H. 

pecten-veneris, discussed its uncertain taxonomic 

relationship to H tenella and noted that examination of cystocarpic and spermatangial thalli 

would help to resolve the problem. Unfortunately, we have not encountered either in our 

collections. 

Herposiphonia secunda (C. Agardh) Ambronn 

Herposiphonia secunda (C. Agardh) Ambronn, 1880:197.— 

Taylor, 1960:604, pl. 72: figs. 10, 11.—Cordeiro-Marino, 

1978:125, figs. 378-381. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

21 Apr 1977,JN-6937a (0). 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Barbados, Grenada, Belize, 

Panama, Netherlands Antilles (Taylor, 1960); 

Puerto Rico (Almodovar, 1965); Nevis, Antigua, 

Dominica (Taylor, 1969). 

REMARKS.—B0rgesen (1918:290, fig. 289) 

found branching patterns attributed to Herposiphonia secunda on otherwise typical H tenella thalli 

and concluded that H. secunda was "most probably . . . nothing more than a reduced form or 

variety of H tenella:' Hollenberg (1968b: 556), 

after examining what was considered to be an 

isotype of H. secunda, shared the view that the 

branching pattern is variable and reduced the 

species to a form: H tenella f. secunda. B0rgesen 

(1920:469-474) later changed his earlier opinion 

and concluded the two taxa were separate species 

because they had "two very different types of 

antheridial plants." Most subsequent authors 

have followed this view and recognized not only 

differences in branching pattern, but also in dimensions, spermatangial features, and number of 

pericentral cells (for example, Taylor, 1960; Dawson, 1963; Oliveira Filho, 1969; Cordeiro-Marino, 

1978). We are adopting this approach until crit-
NUMBER 12 211 

ical studies have evaluated the taxonomic importance of these features. 

Our Belizean material has 7-8 pericentral cells 

and axes not exceeding 80 /xm in diameter. It is 

therefore closest in vegetative features to Herposiphonia secunda as described from elsewhere in the 

Caribbean (Taylor, 1960). 

* Lophosiphonia cristata Falkenberg 

Lophosiphonia cristata Falkenberg, 1901:499.—B0rgesen, 1918: 

297, figs. 295-298.—Taylor, 1960:606. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977,JN-7128a. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica, 

Virgin Islands (Taylor, 1960); Cuba (Suarez, 

1973); Puerto Rico (Diaz-Piferrer, 1963); Grand 

Cayman, Antigua (Taylor, 1969); Isla San Andres 

(Kapraun, 1972); Curacao (van den Hoek, 1969). 

REMARKS.—It is the first record of the genus in 

Belize. 

Polysiphonia atlantica Kapraun and Norris 

Polysiphonia atlantica Kapraun and Norris, herein: 226, Figure 

\Qla-c—B0rgesen, 1918:274, figs. 272-276 [as P. macrocarpa Harvey]. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, epiphytic on Ulva, 31 Mar 1976, JN-6292 (0). 

CARIBBEAN DISTRIBUTION.—Virgin Islands, 

Barbados (Taylor, 1960); Cuba (Diaz-Piferrer, 

1964a); Jamaica (Chapman, 1963); Costa Rica 

(Dawson, 1962b); Belize (Kapraun and Norris, 

herein); Curacao (Diaz-Piferrer, 1964b). 

Polysiphonia denudata (Dillwyn) 

Greville ex Harvey 

Polysiphonia denudata (Dillwyn) Greville ex Harvey in Hooker, 

1833:332.—Taylor, 1960:580.—Kapraun and Norris, 

herein: 228, Figure 107^-/. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 10, 

epiphytic on Dictyota, 29 Apr 1977, JN-6842a 

(0); sta 15, 27 Apr 1977, JN-6805 (9,0), and 

epiphytic on Stypopodium zonale, JN-6686 (0). 

CARIBBEAN DISTRIBUTION.—Virgin Islands, 

Guadeloupe, Barbados, Netherlands Antilles 

(Taylor, 1960); Cuba (Suarez, 1973); Belize (Kapraun and Norris, herein). 

Polysiphonia exilis Harvey 

Polysiphonia exilis Harvey, 1853:47.—Taylor, 1960:581.—Kapraun and Norris, herein: 228, Figure 108. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

epiphytic on Dictyota dichotoma, 5 Apr 1976, JN6342a-c {6). 

CARIBBEAN DISTRIBUTION.—Bahamas (Taylor, 

1960); Cuba (Diaz-Piferrer, 1964a); Belize (Kapraun and Norris, herein); Curacao (Diaz-Piferrer, 1964b). 

Polysiphonia ferulacea Suhr 

Polysiphonia ferulacea Suhr in J. Agardh, 1863:980.—Taylor, 

1960:578.—Kapraun and Norris, herein: 229, Figure 109. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, entangled with Jania, Centroceras, 

and Wrangelia turf, JN-6987; sta 8, 27 Apr 1977, 

epiphytic on Valonia ventricosa, JN-6410 (0), and 

epiphytic on Caulerpa verticillata, JN-6413; sta 10, 

23 Apr 1977, epiphytic on Udotea, JN-6938 {6); 

sta 15, 5 Apr 1976, epiphytic on Dictyota cervicornis, 

JN-6334 (9), JN-6335 (9), and epiphytic on Lobophora variegata, JN-6355 (0), and 27 Apr 1977, 

JN-6691b. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cayman Islands, Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Barbados, 

Costa Rica, Venezuela (Taylor, 1960); Cuba 

(Diaz-Piferrer, 1964a); Puerto Rico (Almodovar 

and Blomquist, 1965); St. Kitts, Antigua, Dominica (Taylor, 1969); Belize (Kapraun and Norris, 

herein); Curacao (Diaz-Piferrer, 1964b); Trinidad (Richardson, 1975). 

Polysiphonia flaccidissima Hollenberg 

Polysiphonia flaccidissima Hollenberg, 1942:783.—Hollenberg 

and Norris, 1977:4, fig. 2E.—Kapraun and Norris, herein: 

231, Figure 110. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 10, 
212 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

epiphytic on Dictyota, 29 Apr 1977, JN-6842b 

(0, 9, 6); sta 15, epiphytic on Sargassum hystrix, 5 

Apr 1976, JN-6351 {6,2), JN-6353 (0). Twin 

Cays: sta 24, epiphytic on Anadyomene stellata, 26 

Apr 1977,JN-6886 (9,0). 

CARIBBEAN DISTRIBUTION.—Belize (Kapraun 

and Norris, herein). 

Polysiphonia scopulorum var. villum 

(J. Agardh) Hollenberg 

Polysiphonia scopulorum var. villum (J. Agardh) Hollenberg, 

1968a:81.—Hollenberg and Norris, 1977:14, fig. 8B.—Kapraun and Norris, herein: 233, Figure 111. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 15, 

epiphytic on Lobophora variegata, 5 Apr 1976, JN6357 (9). 

CARIBBEAN DISTRIBUTION.—Belize (Kapraun 

and Norris, herein). 

Polysiphonia sphaerocarpa B0rgesen 

Polysiphonia sphaerocarpa B0rgesen, 1918:271.—Taylor, 1960: 

576.—Kapraun and Norris, herein: 233, Figure 112. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 10, 

29 Apr 1977, epiphytic on Sargassum, JN-6847a, 

and 23 Apr 1977, epiphytic on Dictyota, JN-6911 

(9); sta 18, epiphytic on Dictyota, JN-6746 (9), JN6747b {6), JN-6748 (0), and JN-6755 (9, 0). 

CARIBBEAN DISTRIBUTION.—Hispaniola, Virgin 

Islands (Taylor, 1960); Antigua, Dominica, Barbados (Taylor, 1969); Belize (Kapraun and Norris, herein). 

Murrayella periclados ( C Agardh) Schmitz 

Murrayella periclados (C. Agardh) Schmitz, 1893:227.—B0rgesen, 1919:314, figs. 318-320.—Taylor, 1960:593. 

LOCAL DISTRIBUTION.—Twin Cays: sta 24, 29 

Apr 1979, JN-7395 (0). 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Cayman Islands, Jamaica, Virgin Islands, Guadeloupe, Martinique, Barbados, Grenada, Belize, 

Guatemala, Panama, Venezuela (Taylor, 1960); 

Puerto Rico (Almodovar, 1964); Antigua (Taylor, 

1969); Curacao, Bonaire (Diaz-Piferrer, 1964b); 

Trinidad (Richardson, 1975). 

* Laurencia caraibica Silva 

Laurencia caraibica Silva, 1972:205.—Taylor, 1960:622 [as 

Laurencia nana]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

22 Apr 1977, JN-6997. South Water Cay: sta 21, 

29 Apr 1977, JN-6388. 

CARIBBEAN DISTRIBUTION.—Bahamas, Jamaica 

(Taylor, 1960); Antigua (Taylor, 1969). 

REMARKS.—The name Laurencia caraibica was 

introduced by Silva (1972) to replace L. nana 

Howe in Britton and Millspaugh (Howe, 1920b: 

566), a later homonym of L. nana (C. Agardh) 

Greville (1830: Iii). 

The plants from Belize are pink to light rose 

and form small decumbent clumps in shallow, 

wave-exposed portions of the reef crest. Lenticular 

thickenings occur in a few cell walls of the medulla and can be seen in both fresh and dried 

Carrie Bow Cay material. These were noted neither by Howe (1920b) nor by Yamada (1931) but 

re-examination of an isotype (US-68437, Howe5393) has revealed their presence. 

Laurencia intricata Lamouroux 

Laurencia intricata Lamouroux, 1813:43.—B0rgesen, 1918: 

251, figs. 241-242 [as L. implicata]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 4, 

25 Apr 1979, JN-7722; sta 19, 28 Apr 1979, JN7627. Twin Cays: sta 24, 29 Apr 1979, JN-7393. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Cayman Islands, Jamaica, Hispaniola, Virgin Islands, Guadeloupe, Grenada, Costa Rica, Panama (Taylor, 1960); St. Kitts, Dominica, Bequia 

(Taylor, 1979); Belize (Tsuda and Dawes, 1974); 

Curacao, Bonaire (Diaz-Piferrer, 1964b); Venezuela (Diaz-Piferrer, 1970b). 

* Laurencia microcladia Kiitzing 

Laurencia microcladia Kiitzing, 1865:22, pl. 60: figs, b, c.— 

Cordeiro-Marino, 1978:130, figs. 395-396. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 2, 

27 Apr 1979JN-7644. 
NUMBER 12 213 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos 

Islands, Cuba, Cayman Islands, Jamaica, Hispaniola, Virgin Islands, St. Barthelemy, Barbuda, 

Guadeloupe, Isla las Aves, Costa Rica, Panama, 

Netherlands Antilles, Venezuela (Taylor, 1960); 

Puerto Rico (Almodovar, 1964); Nevis, Antigua, 

St. Lucia (Taylor, 1969); Dominica (Taylor and 

Rhyne, 1970). 

Laurencia obtusa (Hudson) Lamouroux 

Laurencia obtusa (Hudson) Lamouroux, 1813:42.—B0rgesen, 

1918:247, figs. 237-240. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

21 Apr 1977, JN-6978, 22 Apr 1977, JN-6951 and 

JN-6961, 23 Apr 1976, JN-7667, and 28 Apr 1979, 

coll. J. Norris and I. Macintyre, JN-7487. South 

Water Cay: sta 21, 28 Apr 1977, JN-6728 and 

JN-6734. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Virgin Islands, St. Barthelemy, 

Guadeloupe, Barbados, Belize, Panama, Colombia, Netherlands Antilles, Isla las Aves, Tobago 

(Taylor, 1960); Puerto Rico (Almodovar, 1964); 

Antigua, Dominica, St. Lucia (Taylor, 1969); 

Costa Rica (Dawson, 1962b); Venezuela (Taylor, 

1976). 

Laurencia papillosa (Forsskal) Greville 

Laurencia papillosa (Forsskal) Greville, 1830: Iii.—Taylor, 

1960:623, pl. 74: fig. 2. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

28 Apr 1979, coll. J. Norris and I. Macintyre, JN7486. South Water Cay: sta 21, 28 Apr 1977, JN6720. Twin Cays: sta 24, 29 Apr 1979, JN-7392; 

sta 25, 26 Apr 1977, JN-6879. 

CARIBBEAN DISTRIBUTION.—Mexico, Bahamas, 

Caicos Islands, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Puerto Rico, Virgin Islands, St. Barthelemy, Guadeloupe, Barbados, Grenada, Belize, Costa Rica, Panama, Colombia, Netherlands 

Antilles, Venezuela, Trinidad (Taylor, 1960); St. 

Kitts, Nevis, Antigua, Dominica, St. Lucia, Bequia (Taylor, 1969); Isla San Andres (Kapraun, 

1972). 

* Wrightiella blodgettii (Harvey) Schmitz 

FIGURE 106 

Wrightiella blodgettii (Harvey) Schmitz, 1893:221.—Taylor, 

1960:591. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 1, 

on lower side of Diploria sp., 20 Apr 1977, JN7046 0 ; sta 11, 25 Apr 1977, JN-7055. 

CARIBBEAN DISTRIBUTION.—Bahamas, Guadeloupe (Taylor, 1960). 

REMARKS.—The genus has distinctive spinelike or "spur" branchlets, monosiphonous laterals, 

and spiral tetrasporangial stichidia. Taxonomists 

have questioned whether Wrightiella blodgettii is 

distinct from W. tumanowiczii (Gatty) Schmitz (for 

instance, B0rgesen, 1915; Schneider, 1975). The 

plants from Belize representing a new Belizean 

record for the genus, have numerous spur branchlets and are densely beset with monosiphonous 

branchlets; therefore, they agree with W. blodgettii 

as described by Taylor (1960). 

This species is rare in the Carrie Bow Cay area. 

We found it on a patch reef and noted that it 

superficially resembled a small gorgonian. 

Order (incertae sedis) 

Family WURDEMANNIACEAE 

Wurdemannia miniata (Lamouroux) Feldmann 

and Hamel 

Wurdemannia miniata (Lamouroux) Feldmann and Hamel, 

1934:544.—B0rgesen, 1919:368, figs. 360-361 [as Wurdemannia sectacea]. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

23 Apr 1979,JN-7678. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Jamaica, Virgin Islands, Guadeloupe, Isla las 

Aves (Taylor, 1960); Puerto Rico (Diaz-Piferrer, 

1963); Dominica, Barbados (Taylor, 1969); Belize 

(Tsuda and Dawes, 1974); Curacao, Bonaire 
214 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 106.— Wrightiella blodgettii (JN-7046): a, portion of the thallus showing branches with 

numerous spur branchlets; b, portion of main axis with a spiral tetrasporangial stichidia. 

(Diaz-Piferrer, 1964b); Panama (Earle, 1972); 

Venezuela (Taylor, 1976). 

Division MAGNOLIOPHYTA (= ANGIOSPERMAE) 

Class LlLIOPSIDA (= MONOCOTYLEDONEAE) 

Order NAJADALES 

Family CYMODOCEACEAE 

Halodule wrightii Ascherson 

Halodule wrightii Ascherson, 1868:19.—den Hartog, 1970: 

154.—Hanlon and Voss, 1975:18, fig. 4.—Howard, 1979: 

11, fig. 3b-c. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21, 16 Nov 1980, JN-9818. Twin Cays: SW of sta 

26, 25 May 1980, coll. K. Riitzler, JN-9815. 

CARIBBEAN DISTRIBUTION.—Cuba, Haiti, 

Puerto Rico, Jamaica, St. Martin, Guadeloupe 

(den Hartog, 1970); Belize (Tsuda and Dawes, 

1974). 

Syringodium filiforme Kiitzing 

Syringodium filiforme Kiitzing in Hohenacker, 1860: no. 426.— 

den Hartog, 1970:183.—Hanlon and Voss, 1975:20, figs. 

5, 9.—Howard, 1979:12, fig. 3a. 

LOCAL DISTRIBUTION.—South Water Cay: sta 

21,28 Apr 1977, JN-6733. 

CARIBBEAN DISTRIBUTION.—Bahamas, Caicos, 

Cuba, Hispaniola, Puerto Rico, Jamaica, Cay-
NUMBER 12 215 

man Islands, Virgin Islands, St. Martin, Antigua, 

Saba Island, Guadeloupe, Dominica, Martinique, 

Costa Rica, Panama, Isla de San Andres, Curacao, Tortuga Island (den Hartog, 1970); Belize 

(Tsuda and Dawes, 1974). 

REMARKS.—Rather common and usually intermixed with Thalassia in sandy areas. Flowers were 

observed in April 1979. 

Order HYDROCHARITALES 

Family HYDROCHARITACEAE 

Thalassia testudinum Banks ex Konig 

Thalassia testudinum Banks ex Konig, 1805:96.—den Hartog, 

1970:223.—Hanlon and Voss, 1975:12, figs. 1, 8.—Howard, 1979:24, Fig. 7a. 

LOCAL DISTRIBUTION.—Carrie Bow Cay: sta 7, 

23 Jun 1977, leg. K. Riitzler, CBC-77.6.23. Twin 

Cays: sta 25, 26 Apr 1977, JN-6869. 

CARIBBEAN DISTRIBUTION.—Bahamas, Cuba, 

Cayman Islands, Jamaica, Haiti, Puerto Rico, 

Virgin Islands, St. Barthelemy, Antigua, Guadeloupe, Martinique, Barbados, Grenada, Trinidad, Tortuga Island, Curasao, Belize, Costa Rica, 

Panama (den Hartog, 1970). 

REMARKS.—Widespread in the vicinity of Carrie Bow Cay, preferring sandy areas of the shallower depths. Flowers were observed in April 

1979. Blades were often heavily laden with epiphytes in spring. 

Summary and Conclusions 

Our collections of marine algae and seagrasses 

from the vicinity of Carrie Bow Cay yielded 34 

genera, 77 species, six varieties, and five forms 

that are new to Belize. Thus, the marine flora of 

Belize now totals 247 plant taxa, including 

nine Cyanophyta, 73 Chlorophyta, 32 Phaeophyta, 124 Rhodophyta, and four Magnoliophyta (= Angiospermae). 

Most major habitats of the lagoon, reef crest, 

and fore reef are characterized by certain common algae. A particularly interesting feature of 

the lagoon is the mangrove community of Twin 

Cays (stations 24-26). The intertidal stilt roots of 

the predominant red mangrove support three 

species of Bostrychia, along with Catenella repens 

and Caloglossa leprieurii, all known to be common 

associates of mangroves (see Taylor, 1960). Mudbanks of the subtidal zone are covered by extensive mats oi Caulerpa verticillata laden with detritus, 

and submerged roots provide substrate to two 

other Caulerpa species, C. paspaloides—an unusually large form of this species—and C. mexicana. 

Two other algae are conspicuous because they are 

commonly found on dead or even decaying mangrove roots, Acetabularia crenulata and Neomeris annulata. In the Thalassia bed adjacent to the mangroves (station 26), Penicillus dumentosus and Rhipocephalus phoenix are predominant, accompanied 

by less common Dictyosphaeria cavernosa. The seagrass beds closer to Carrie Bow Cay (station 4) 

are composed of Thalassia testudinum and Syringodium filiforme. They abound with Udotea flabeHum, 

Halimeda incrassata, and Avrainvillea longicaulis, 

along with some epiphytic species of Dasya. Both 

seagrasses were in flower during our surveys between March and May. 

The shallow reef flat (station 6) seaward of 

Carrie Bow Cay is also dominated by a Thalassia 

community. Two conspicuous components are 

the coralline algae Amphiroa fragillissima and Neogoniolithon strictum, the latter forming unattached 

nodules. Thalassia blades provide substrate for a 

number of small epiphytic algae. One of these, 

Champia parvula, was particularly abundant during spring 1979 when it appeared as a conspicuous yellow coverage over the greater part of the 

reef flat (see also Riitzler and Macintyre, herein: 

9, Figures 6b, lb). 

Shallow back-reef areas were also examined 

near South Water Cay, one kilometer to the north 

of Carrie Bow Cay. Their most distinctive features are widespread dense stands of Turbinaria 

turbinata and Sargassum polyceratium off the north 

tip of the cay (station 23). Rather patchy in 

distribution but unique to the area is Ochtodes 

secundiramea growing in the shallows of the reef 

crest east of the island (station 21). 

The wave-exposed intertidal reef crest off Car-
216 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

rie Bow Cay (station 7) has a rich and characteristic algal flora. The leeward coral rock is covered 

by extensive mats of Caulerpa racemosa and Halimeda opuntia. Laurencia obtusa and L. caraibica are 

most noticeable on the exposed parts of the crest, 

whereas numerous but small tufts of Wrangelia 

argus are found in surge areas. The abundant 

crustose corallines include Hydrolithon boergesenii 

and Porolithon pachydermum. Also common are turfs 

formed mainly by Acetabularia myriospora and Neomeris mucosa. Less abundant but characteristic for 

the reef crest and not found elsewhere are Liagora 

valida, Eucheuma echinocarpum, E. schrammii, and 

Gracilaria crassissima. 

On the fore reef, red algae of the genus Liagora 

characterize the spur and groove zone (stations 

8-10) in 4-12 m depth. Liagora farinosa and L. 

pinnata predominate, along with two possibly undescribed species, one belonging to Liagora, the 

other one to the related genus Trichogloeopsis. Dudresnaya crassa, generally considered rare in the 

Caribbean, is also common in this zone in the 

spring. Lobophora variegata, Stypopodium zonale, and 

Anadyomene stellata are abundant on the outer 

ridge and down the fore-reef slope to 40 m depth. 

All three species are also found in shallow water, 

although there they are morphologically different. Less common but restricted to this deepwater habitat are Galaxaura obtusa and Kallymenia 

limminghii. 

Collecting during different seasons and in the 

diverse marine environments throughout the entire barrier reef, numerous cays, and along the 

mainland coast would be expected to result in the 

discovery of many plants as yet unknown or 

unreported in Belize. 

Literature Cited 

Abbott, I. A. 

1945. The Genus Liagora (Rhodophyceae) in Hawaii. 

Occasional Papers of Bernice Bishop Museum, 17(10): 

145-169. 

1976. Dotyophycus pacificum gen. et sp. nov., with a Discussion of Some Families of Nemaliales (Rhodophyta). Phycologia, 15(2): 125-132. 

1979. Some Tropical Species Related to Antithamnion 

(Rhodophyta, Ceramiaceae). Phycologia, 18(3): 

213-227. 

Abbott, I. A., and M. S. Doty 

1960. Studies in the Helminthocladiaceae, II: Trichogloeopsis. American Journal of Botany, 47(8): 632-640. 

Abbott, I. A., and G. J. Hollenberg 

1976. Marine Algae of California, xii + [2] + 827 pages. 

Stanford, California: Stanford University Press. 

Abbott, I. A., and M. M. Littler 

1969. Some Rhodymeniales from Hawaii. Phycologia, 

8(3/4): 165-169. 

Adey, W. H., and I. G. Macintyre 

1973. Crustose Coralline Algae: A Re-evaluation in the 

Geological Sciences. Geological Society of America 

Bulletin, 84:883-994. 

Agardh, C. A. 

1822. Species algarum ... . Volume 1, part 2, pages [vvi] + 169-398. Lund: Berling. 

1823. Species algarum ... . Volume 1, part 2, pages [viiviii] + 399-531. Greifswald: Ernst Mauritius. 

1824. Systema algarum. xxxviii + 312 pages. Lund: Berling. 

1827. Aufzahlung einiger in den osterreichischen Landern gefundenen neuen Gattungen und Arten von 

Algen, nebst ihrer Diagnostik und beigefiigten 

Bemerkungen. Flora oder Botanische Zeitung, 10:625-

646. 

1828. Species algarum ... . Volume 2, part 1, lxxvi + 

189 pages. Greifswald: Ernst Mauritius. 

Agardh, J. G. 

1842. Algae Maris Mediterranei et Adriatici.... x + 164 

pages. Paris: Fortin, Masson. 

1847. Nya alger fran Mexico. Oversigh af Konglig 

Velenskaps-Akademiens Fbrhandlingar, 4(1):5— 17. 

1848. Species genera et ordines algarum ... . Volume 1, viii 

+ 363 pages. Lund: C.W.K. Gleerup. 

1851. Species genera et ordines algarum ... . Volume 2, 

part 1, xii + 336 -I- [14] pages. Lund: C.W.K. 

Gleerup. 

1852. Species genera et ordines algarum ... . Volume 2, 

part 2, pages [2] + 337-700 + [20] + 701-786. 

Lund: C.W.K. Gleerup. 

1863. Species genera et ordines algarum ... . Volume 2, 

part 3, pages [2]+ 787-1291. Lund: C.W.K. Gleerup. 
NUMBER 12 217 

1873. Till algernes systematik . . . [Part 1, Numbers iiii]. Lunds Universitels Arsskrift, ny foeljd, 9:1-71. 

1876. Species genera et ordines algarum ... . Volume 3, 

part 1, viii -I- 724 pages. Leipzig: T. O. Weigel. 

1882. Till algernes systematik . . . [Part 2, Numbers ivv]. Lunds Universitets Arsskrift, ny foeljd, 17: 1-134 

+ [2], plates 1-3. 

1883. Till algernes systematik . . . [Part 3, Number vi]. 

Lunds Universitets Arsskrift, ny foeljd, 19:1-177 + 

[4], plates 1-4. 

1885. Till algernes systematik . . . [Part 4, Number vii]. 

Lunds Universitets Arsskrift, ny foeljd, 21:1-177 + 

[3], plate 1. 

1892. Analecta algologica ... . Lunds Universitets Arsskrift, ny foeljd, 28(6): i -I- 182, plates 1-3. 

Almodovar, L. R. 

1964. The Marine Algae of Guanica, Puerto Rico. Revue 

Algologiue, 7(2): 129-150. 

1965. The Unnamed Rhodophyta of the Marshall A. 

Howe Collection of Marine Algae from Puerto 

Rico. Nova Hedwigia, 9(1-4): 1-19. 

1970. Deep-Water Algae New to Puerto Rico. Quarterly 

Journal of the Florida Academy of Sciences, 33(1): 23-

28. 

Almodovar, L. R., and H. L. Blomquist 

1965. Some Marine Algae New to Puerto Rico. Nova 

Hedwigia, 9:63-71. 

Almodovar, L. R., and I. Bonnelly de Calventi 

1977. Notas sobre las Algas Marinas Bentonicas Macroscopicas 

de la Republica Dominicana. 22 pages. Santo Domingo, Dominican Republic: Editora Taller. 

[Universidad Autonoma de Santo Domingo, Investigaciones Marinas.] 

Almodovar, L. R., and F. A. Pagan 

1967. Notes on the Algae of Barbados. Nova Hedwigia, 

13:111-115. 

Ambronn, H. 

1880. Uber einige Falle von Bilateralitat bei den Florideen. Botanische Zeitung, 38(12): 193-200. 

Ardissone, F. 

1871. Rivista dei Ceramii della flora Italiana. Nuovo 

Giornale Botanico Italiano, 3:32-50. 

1878. Studi sulle alghe italiche della famiglia della Rhodomelacee. Atti della Societd Crittogamologica Italiana, 

1:41-159. 

Areschoug, J. E. 

1854. Phyceae novae et minus cognitae ... . Nova Acta 

Regiae Societatis Scientiarum Upsaliensis, series iii, 1: 

328-372 [reprint pages 1-46]. 

Ascherson, P.F.A. 

1868. [Untitled]. In Anonymous, [Report of a meeting 

on 16 June 1868]. Sitzungs-Bericht der Gesellschaft 

Naturforschender Freunde zu Berlin, 1868:18-19. 

Baldock, R. N. 

1976. The Griffithsieae Group of the Ceramiaceae (Rhodophyta) and Its Southern Australian Representatives. Australian Journal of Botany, 24(4):509-593. 

Barton, E. S. 

1891. A Systematic and Structural Account of the Genus 

Turbinaria Lamouroux. Transactions of the Linnean 

Society of London, 2nd series, 3:215-226. 

Batters, E.A.L. 

1902. A Catalogue of the British Marine Algae. Journal 

of Botany (London), 40(supplement): 1-107. 

Blainville, H.M.D. de 

1834. Manuel d'Actinologie ... . viii -I- 694 pages. Paris: 

F. G. Levrault. 

Bliding, C. 

1963. A Critical Survey of European Taxa in Ulvales, I: 

Capsosiphon, Percursaria, Blidingia, Enteromorpha. Opera Botanica, 8(3): 1-160. 

1968. A Critical Survey of European Taxa in Ulvales, 

II: Ulva, Ulvaria, Monostroma, Kornmannia. Botaniska 

Notiser, 121:535-629. 

B0rgesen, F. 

1905. Contributions a la connaissance du genre Siphonocladus Schmitz. Oversigl over del Kongelige Danske 

Videnskabernes Selskabs Forhandlinger og dels Medlemmers Arbeider, 1905(3):259-291. 

1907. An Ecological and Systematic Account of the 

Caulerpas of the Danish West Indies. Del Kongelige 

Danske Videnskabernes Selskabs Naturvidenskabelige og 

Mathematiske Afhandlinger, series iv, 4(5):337-392. 

1908. The Species oi Avrainvillea Hitherto Found on the 

Shores of the Danish West Indies. Videnskabelige 

Meddelelser fra dansk Naturhistorisk Forening i Kj0benhavn, 1908:27-44, plate 3. 

1909. Some New or Little Known West Indian Florideae, Botanisk Tidsskrift, 30:1-19. 

1910. Some New or Little Known West Indian Florideae, II. Botanisk Tidsskrift, 30:117-207. 

1911. Some Chlorophyceae from the Danish West Indies, I. Botanisk Tidsskrift, 31:127-152. 

1913. The Marine Algae of the Danish West Indies, Part 

1: Chlorophyceae. Dansk Botanisk Arkiv, 1(4): 1-158 

+ [2] + map. 

1914. The Marine Algae of the Danish West Indies, Part 

2: Phaeophyceae. Dansk Botanisk Arkiv, 2(2): 1-66 

+ [2]. 

1915. The Marine Algae of the Danish West Indies, Part 

III: Rhodophyceae (1). Dansk Botanisk Arkiv, 3(1 a): 

1-80. 

1916. The Marine Algae of the Danish West Indies, Part 

III: Rhodophyceae (2). Dansk Botanisk Arkiv, 3(lb): 

81-144. 

1917. The Marine Algae of the Danish West Indies, Part 

III: Rhodophyceae (3). Dansk Botanisk Arkiv, 3(lc): 

145-240. 

1918. The Marine Algae of the Danish West Indies, Part 
218 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

IV [sic]: Rhodophyceae (4). Dansk Botanisk Arkiv, 

3(ld):241-304. 

1919. The Marine Algae of the Danish West Indies, Part 

IV [sic]: Rhodophyceae (5). Dansk Botanisk Arkiv, 

3(le):305-368. 

1920. The Marine Algae of the Danish West Indies, Part 

III: Rhodophyceae (6), with Addenda to the Chlorophyceae, Phaeophyceae and Rhodophyceae. 

Dansk Botanisk Arkiv, 3(If):369-498 + [6]. 

1924. Marine Algae. In C. H. Ostenfeld, Plants from 

Beata Island, St. Domingo (Botanical Results of 

the Dana-Expedition 1921-22, No. 1). Dansk Botanisk Arkiv, 4(7): 14-35. 

1927. Marine Algae from the Canary Islands, Especially 

from Teneriffe and Gran Canaria, III: Rhodophyceae, Part I: Bangiales and Nemalionales. Biologiske Meddelelser, Det Kongelige Danske Videnskabernes 

Selskab, 6(6): 1-97. 

1932. A Revision of Forsskal's Algae Mentioned in Flora 

Aegyptico Arabica and Found in His Herbarium 

in the Botanical Museum of the University of 

Copenhagen. Dansk Botanisk Arkiv, 8(2): 1-15. 

1934a. Some Marine Algae from the Northern Part of the 

Arabian Sea with Remarks on their Geographical 

Distribution. Biologiske Meddelelser, Det Kongelige 

Danske Videnskabernes Selskab, 11:1-72. 

1934b. Some Indian Rhodophyceae Especially from the 

Shores of the Presidency of Bombay, IV. Bulletin 

of Miscellaneous Information (Royal Botanic Gardens, 

Kew), 1(1934): 1-30, figures, 4 plates. 

1945. Some Marine Algae from Mauritius, III: Rhodophyceae, Part 4: Ceramiales. Biologiske Meddelelser, 

Det Kongelige Danske Videnskabernes Selskab, 19(10): 

1-68. 

Bula Meyer, G. 

1977. Algas marinas benticas indicadoras de un area 

afectada por aguas de surgencia frente a la costa 

Caribe de Colombia. Anales del Instituto Investigaciones Marinas, Punta Betin (Santa Marta, Colombia), 9:45-71. 

Chapman, V. J. 

1961. The Marine Algae of Jamaica, Part I: Myxophyceae and Chlorophyceae. Bulletin of the Institute of 

Jamaica, Science Series, 12(1): 159 pages. 

1963. The Marine Algae of Jamaica, Part II: Phaeophyceae and Rhodophyceae. Bulletin of the Institute of 

Jamaica, Science Series, 12(2): 201 pages. 

Cheney, D. P., and C. J. Dawes 

In press. The Genus Eucheuma (Rhodophyta; Gigartinales) 

in Florida and the Caribbean: Morphological 

Variation, Ecotypes, and Taxonomic Treatment. 

Phycologia. 

Chou, R.C.-Y. 

1947. Pacific Species of Galaxaura, II: Sexual Types. Papers of the Michigan Academy of Science, Arts, and 

Letters, 31(1945):3-24, plates 1-13. 

Colinvaux, L. H. 

1968. New Species of Halimeda: A Taxonomic Reappraisal. Journal of Phycology, 4(l):30-35. 

Collins, F. S. 

1901. The Algae of Jamaica. Proceedings of the American 

Academy of Arts and Sciences, 39(9) :231-270. 

1909. The Green Algae of North America. Tufts College 

Studies, 2(3): 79-480, plates 1-18. 

Collins, F. S., and A. B. Hervey 

1917. The Algae of Bermuda. Proceedings of the American 

Academy of Arts and Sciences, 53:1 -195. 

Cordeiro-Marino, M. 

1978. Rhodoficeas bentonicas marinhas do Estado de 

Santa Catarina. Rickia, 7:1-243. 

Crouan, P. L., and H. M. Crouan 

1859. Notes sur quelques especes et genres nouveaux 

d'algues marines de la rade de Brest. Annates des 

Sciences Naturelles (Botanique), series 4, 12 (5): 288-

292. 

Dawson, E. Yale 

1961. A Guide to the Literature and Distributions of 

Pacific Benthic Algae from Alaska to the 

Galapagos Islands. Pacific Science, 15(3):370-461. 

1962a. Marine Red Algae of Pacific Mexico, Part 7: 

Ceramiales; Ceramiaceae, Delesseriaceae. Allan 

Hancock Pacific Expeditions, 26(1): 1-207. 

1962b. Additions to the Marine Flora of Costa Rica and 

Nicaragua. Pacific Naturalist, 3(13):375-395. 

1963. Marine Red Algae of Pacific Mexico, Part 8: 

Ceramiales; Dasyaceae, Rhodomelaceae. Nova 

Hedwigia, 6:401-481. 

Decaisne, J. 

1842. Memoire sur les corallines ou polypiers calciferes. 

Annates des Sciences Naturelles (Botanique), series 2, 

18:96-128. 

den Hartog. See Hartog, C. den 

Derbes, A. A., and A.J.J. Solier 

1856. Memoire sur quelques points de la Physiologie des 

Algues. Supplement aux Comptes Rendus Seances de 

l'Academie des Sciences, Paris, 1:1-120, plates 1-23. 

Diaz-Piferrer, M. 

1963. Adiciones a la flora marina de Puerto Rico. Caribbean Journal of Sciences, 3(4) :215-235. 

1964a. Adiciones a la flora marina de Cuba. Caribbean 

Journal of Science, 4(2/3): 353-371. 

1964b. Adiciones a la flora marina de las Antillas Holandesas, Curazao y Bonaire. Caribbean Journal of Sciences, 4(4) :413-543. 

1970a. Nuevas adiciones a la flora marina de Puerto Rico. 

Caribbean Journal of Science, 10(1/2) :9-15. 

1970b. Adiciones a la flora marina de Venezuela. Caribbean Journal of Sciences, 10 (3/4): 159-198. 

Dickie, G. 

1874. On the Algae of Mauritius. Journal of the Linnean 
NUMBER 12 219 

Society [of London] (Botany), 14:190-202. 

Dixon, P. S. 

1976. Appendix I. In M. Parke and P. S. Dixon, Checklist of British Marine Algae—Third Re vision. Journal of the Marine Biology Association, U.K., 56:590. 

Dixon, P. S., and L. S. Irvine 

1977. Introduction, Nemaliales, Gigartinales. In Seaweeds 

of the British Isles, 1(1): xi -I- 252 pages. London: 

British Museum (Natural History). 

Earle, S. A. 

1969. Phaeophyta of the Eastern Gulf of Mexico. Phycologia, 7(2): 71-254. 

1972. A Review of the Marine Plants of Panama. Bulletin 

of the Biological Society of Washington, 2:69-87. 

Egerod, L. E. 

1952. An Analysis of the Siphonous Chlorophycophyta 

with Special Reference to the Siphonocladales, 

Siphonales, and Dasycladales of Hawaii. University 

of California Publications in Botany, 25 (5): 325-454. 

Eubank, L. 

1946. Hawaiian Representatives of the Genus Caulerpa. 

University of California Publications in Botany, 18(18): 

409-432. 

Falkenberg, P. 

1901. Die Rhodomelaceen des Golfes von Neapel und 

der angrenzenden Meeres-Abschnitte. Fauna und 

Flora des Golfes von Neapel, Monographic, 26: xvi + 

1-754 pages, 24 plates. Berlin: R. Friedlander & 

Sohn. 

Fan, K., and Y. Wang 

1974. Studies on the Marine Algae of Hsisha Islands, 

China, I: Ganonema gen. nov. Acta Phytotaxonomica 

Simca, 12(4): 489-495. 

Feldmann, J., and G. Hamel 

1934. Observations sur quelques Gelidiacees. Revue Generate de Botanique, 46(1934):528-550. 

Feldmann, J., and R. Lami 

1935. La Vegetation marine. In Stehle, H., Flore de la 

Guadeloupe et de'pendances: Essai d'e'cologie et geographic 

botanique, pages 251-260. Basse-Terre. 

Feldmann-Mazoyer, G. 

1941. Recherches sur les Ceramiacees de la Mediterranee occidentale. [ii] + 1-510 pages, 4 plates. Alger: Imprimerie Minerva. 

Foslie, M. H. 

1909. Algologiske Notiser, VI. Det Kongelige Norske Videnskabers Selskab Skrifter, 1909(2): 1-63. 

Foslie, M. H. [H. Printz, editor] 

1929. Contributions to a Monograph of the Lithothamnia. 60 

+ [2] pages, 75 plates. Trondheim: Det Kongelige 

Norske Videnskabers Selskab Museet. 

Ganesan, E. K., and A. J. Lemus 

1969. Notes on Venezuelan Marine Algae, 1: On the 

Occurence of the Red Algal Genus Taenioma J. 

Ag. Nova Hedwigia, 18:765-767. 

Gepp, A., and E. S. Gepp 

1905. Notes on Penicillus and Rhipocephalus. The Journal of 

Botany, British and Foreign, 43:1-5, plate 468. 

1911. The Codiaceae of the Siboga Expedition, Including a Monograph of the Flabellarieae and the 

Udoteae. Siboga-Expeditie, Uitkomsten op Zoologisch, 

Botanisch, Oceanographisch en Geologisch Gebied uitgegeven door. . . M. Weber, Monographic, 62:1-150, 22 

plates. Leiden: E. J. Brill. 

Goreau, T. F., and E. A. Graham 

1967. A New Species oi Halimeda from Jamaica. Bulletin 

of Marine Science of the Gulf and Caribbean, 17(2) :432-

441. 

Greville, R. K. 

1830. Algae Britanmcae . . . . lxxxviii + 218 pages, 19 

plates. Edinburgh: MacLachlan and Stewart. 

Guiry, M.D. 

1978. Notes on Some Family Names of Florideophyceae 

(Rhodophyta). Taxon, 27(2/3): 191-195. 

Hanlon, R., and G. Voss 

1975. A Guide to the Sea Grasses of Florida, the Gulf of 

Mexico, and the Caribbean Region. Sea Grant Field 

Guide Series, 4: 30 pages. Miami, Florida: University of Miami Sea Grant Program. 

Hartog, C. den 

1970. Sea-Grasses of the World. Verhandelingen der Koninklijke Nederlandse Akademie van Wetenschappen, Afdeeling Natuurkunde, tweede reeks, deel 159(1): 275 

pages + 20 pages of photographs. Amsterdam: 

North-Holland Publishing Company. 

Harvey, W. H. 

1849. Nereis Australis [Part 2], pages 65-124, plates 

26-50. London: Reeve Brothers. 

1853. Nereis Boreali-Americana .. . Part II: Rhodospermeae. 

[First issue] ii + 258 pages, plates 13-36. Washington, D.C: Smithsonian Institution [London: 

John Van Voorst]. [Third issue, 1853 [1858], 

Smithsonian Contributions to Knowledge, 5(5): ii + 258 

pages, plates 13-36.] 

Hauck, F. 

1884. Die Meeresalgen Deutschlands und Osterreichs 

[Part 8]. In L. Rabenhort, Kryptogamen-Flora von 

Deutschland, Osterreich und der Schweiz, edition 2, 2: 

385-448. Leipzig: Edward Kummer. 

Heydrich, F. 

1892. Beitrage zur Kenntniss der Algenflora von KaiserWilhelmsland (Deutsch-New-Guinea). Berichte der 

Deutschen Botanischen Gesellschaft, 10:458-485. 

Hoek, C. van den 

1963. Revision of the European Species of Cladophora. vii + 

248 pages, 55 plates. Leiden: E. J. Brill. 

1969. Algal Vegetation—Types Along the Open Coasts 

of Curacao, Netherlands Antilles, II. Proceedings of 

the Kuninklijke Nederlandse Akademie Van Wetenschappen, series C (Biological and Medical Sciences), 

72:559-577. 
220 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Hohenacker, R. F. 

1860. Algae marinae exsiccatae. Fascicle 9, number 426. 

[Distributed by author]. 

Hollenberg, G. J. 

1942. An Account of the Species of Polysiphonia on the 

Pacific Coast of North America, I: Oligosiphonia. 

American Journal of Botany, 29:772-785. 

1968a. An Account of the Species of Polysiphonia of the 

Central and Western Tropical Pacific Ocean, I: 

Oligosiphonia. Pacific Science, 22(1):56-98. 

1968b. An Account of the Species of the Red Algae Herposiphonia Occurring in the Central and Western 

Tropical Pacific Ocean. Pacific Science, 22(4) :536-

559. 

Hollenberg, G. J., and J. N. Norris 

1977. The Red Alga Polysiphonia (Rhodomelaceae) in the 

Northern Gulf of California. Smithsonian Contributions to the Marine Sciences, 1: iii + 21 pages. 

Hooker, W. J. 

1833. Mosses, Hepatiae, Lichens, Characeae and Algae. 

In The English Flora of Sir James Edward Smith, 

edition 2, 5(1): x + 4 + 432 pages. London: Rees, 

Orme, Brown, Green, Longman. [Also issued in 

volume 2 of W. J. Hooker, The British Flora, 

1838]. 

Howard, R. A. 

1979. Monocotyledoneae. In Flora of the Lesser Antilles, 3: 

xi + 586 pages. Jamaica Plain, Massachusetts: 

Arnold Arboretum, Harvard University. 

Howe, M. A. 

1904. Notes on Bahaman Algae. Bulletin of the Torrey 

Botanical Club, 31(2):93-100, plate 6. 

1905. Phycological Studies, II: New Chlorophyceae, 

New Rhodophyceae, and Miscellaneous Notes. 

Bulletin of the Torrey Botanical Club, 32:563-586, 

plates 23-29. 

1907. Phycological Studies, III: Further Notes on Halimeda and Avrainvillea. Bulletin of the Torrey Botanical 

Club, 34:491-516, plates 25-30. 

1909. Phycological Studies, IV, The Genus Neomeris and 

Notes on Other Siphonales. Bulletin of the Torrey 

Botanical Club, 36:75-104, plates 1-8. 

1914. The Marine Algae of Peru. Memoirs of the Torrey 

Botanical Club, 15: [i] + 185 pages, plates 1-66. 

1920a. Observations on Monosporangial Discs in the Genus Liagora. Bulletin of the Torrey Botanical Club, 47: 

1-8. 

1920b. Class 2, Algae. In N. L. Britton and C. F. Millspaugh, The Bahama Flora, pages 553-618. New 

York: Published by the authors. 

Humm, H. J., and H. H. Hildebrand 

1962. Marine Algae from the Gulf Coast of Texas and 

Mexico. Publications of the Institute of Marine Science, 

University of Texas, 8:227-268. 

Joly, A. B. 

1965. Flora marinha do litoral norte do Estado de Sao 

Paulo e regioes circunvizinhas. Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Boletim 

294, Serie Botanica, 21:1-393, plates 1-59. 

Joly, A. B., M. Cordeiro-Marino, Y. Ugadim, N. Yamaguishi-Tomita, and F. C. Pinheiro 

1965. New Marine Algae from Brazil. Arquivos da Estacao 

de Biologia Marinha da Universidade do Ceard, 5(2): 

79-92. 

Joly, A. B., and Y. Ugadim 

1966. The Reproduction of Ochtodes secundiramea (Montagne) Howe (Gigartinales, Rhizophyllidaceae). 

Boletim do Instituto Oceanogrdfico, Universidad de Sao 

Paulo, 15(l):55-64. 

Kapraun, D. F. 

1972. Notes on the Benthic Marine Algae of San Andres, 

Colombia. Caribbean Journal of Science, 12(3/4): 199-

203. 

Kjellman, F. R. 

1900. Om Floride-slagtet Galaxaura, dess organografi och 

systematik. Kongelige Svenska Vetenskapsakademiens 

Handlingar, 33(1): 1-110, plates 1-20. 

K6nig, C. [K.D.E.] 

1805. Additions to M. Cavolini's Treatise on Zostera 

oceanica. In C.D.E. Konig and J. Sims, editors, 

Annals of Botany, 2 (1): 91 -98. 

Kuntze, C.E.O. 

1898. Revisio generum planlarum ... . Part 3(3), 576 

pages. Leipzig: Arthur Felix. 

Kiitzing, F. T. 

1843. Phycologia generalis ... . xxxii + 458 pages. Leipzig: F. A. Brockhaus. 

1849. Species algarum. vi + 922 pages. Leipzig. F. A. 

Brockhaus. 

1858. Tabulae phycologicae ... . Volume 8, parts 1-2, ii 

+ 48 pages, 100 plates. Nordhausen: Kosten des 

Verfassers. 

1859. Tabulae phycologicae ... . Volume 9, parts 1-2, viii 

+ 42 pages, 100 plates. Nordhausen: Kosten des 

Verfassers. 

1865. Tabulae phycologicae ... . Volume 15, parts 1-2, ii 

+ 36 pages, 100 plates. Nordhausen: Kosten des 

Verfassers. 

Kylin, H. 

1925. The Marine Red Algae in the Vicinity of the 

Biological Station at Friday Harbor, Washington. 

Lunds Universitets Arsskrift, ny foeljd, avdeling 2, 

21 (9): 1-87. [Also issued as Kunglig Fysiografiska 

Sdllskapets Handlingar, ny foeljd, 36(9): 1-87.] 

1932. Die Florideenordnung Gigartinales. Lunds Universitels Arsskrift, ny foeljd, avdeling 2, 28(8): 1-88, 

plates 1-28. [Also issued as Kunglig Fysiografiska 

Sdllskapets Handlingar, ny foeljd, 43(8): 1-88, plates 

1-28.1 
NUMBER 12 221 

1940. Die Phaeophyceenordnung Chordariales. Lunds 

Universitets Arsskrift, ny foeljd, avdeling 2, 36(9): 1-

67, plates 1-8. [Also issued as Kunglig Fysiografiska 

Sdllskapets Handlinger, ny foeljd, 51(9): 1-67, plates 

1-8.] 

Lamarck, J.B.A.P.M. de 

1813. Sur les Polypiers empates. Annates du Museum 

d'Histoire Naturelle, Paris, 20(7):294-312. 

Lamouroux, J.V.F. 

1809a. Exposition des caracteres du genre Dictyota (1) et 

tableau desespeces qu'il renferme. Journal de Botanique, Paris, 2:38-44. 

1809b. Observations sur la physiologie des algues marines, 

et description de cinq nouveaux genres de cette 

famille. Nouveau Bulletin des Sciences, Public' par la 

Societe Philomatique de Paris, 1:330-333. 

1809c. Memoire sur trois nouveaux genres de la famille 

des algues marines. Journal de Botanique (Desvaux), 

2:129-135, plate 1. 

1809d. Memoire sur les Caulerpes. Journal de Botanique 

(Desvaux), 2:136-146. 

1812. Extrait d'un memoire sur la classification des polypiers coralligenes non entierement pierreux. Nouveau Bulletin des Sciences, Pub lie par la Societe Philomatique de Paris, 3(63): 181-188. 

1813. Essai sur les genres de la famille des thalassiophytes 

non articulees. Annates du Museum d'Histoire Naturelle, Paris, 20:21-47, 115-139, 267-293, plates 7-

13. 

1816. Histoire des polypiers coralligenes flexibles, vulgairement 

nommes zoophytes, lxxxiv + 560 pages, 19 plates. 

Caen: F. Poisson. 

Masaki, T., and J. Tokida 

1960. Studies on the Melobesioideae of Japan, III. Bulletin of the Faculty of Fisheries, Hokkaido University, 

11 (2):37-42, plates 1-7. 

Maze, H., and H. Schramm 

1878. Essai de classification des algues de la Guadeloupe, xix 

+ 283 + [iii] pages. Basse-Terre, Guadeloupe: 

Imprimerie du Gouvernement. 

Mazoyer, G. 

1938. Les Ceramiaees de l'Afrique du Nord. Bulletin de la 

Societe d'Histoire Naturelle de l'Afrique du Nord, 29: 

317-331. 

McConnell, O. J., and W. Fenical 

1978. Ochtodene and Ochtodiol: Novel Polyhalogenated Cyclic Monoterpenes from the Red Seaweed 

Ochtodes secundiramea. Journal of Organic Chemistry, 

43:4238-4241. 

Meneghini, G. 

1840. Lettera del Prof. Giuseppe Meneghini al Doltora Iacob 

Corinaldi a Pisa. [4]pages. Padova: Tipografia Prosperi. 

Montagne, [J.P.F.] C. 

1837. Centurie de plantes cellulaires exotiques nouvelles. 

Annates des Sciences Naturelles (Botanique), series 2, 

8:345-370. 

1842. Algae. In Ramon de la Sagra, Histoire physique, 

politique et naturelle de file de Cuba: Botanique—Plantes 

cellulaires, pages 1-104, plates 1-5. Paris. 

1846. Phyceae. In M. C. Durieu de Maisonneuve, Exploration scientifique de I'Algerie . . . , Botanique, 1 (Cryptogamie), pages 1-197, plates 1-16. Paris: Imprimerie Royale. 

1856. Sylloge generum specierumque Cryplogamarum quas in 

variis operibus descriptas iconibusque illustratas, nunc ad 

diagnosim reductas, nonnullasque novas inferjectas, ordine 

systematico disposuit. xxiv + 497 pages. Paris: J.-B. 

Bailliere. 

1860. Neuvieme centurie de plantes cellulaires nouvelles, 

tant indigenes qu'exotiques: Decades I, II. Annates 

des Sciences Naturelles (Botanique), series 4, 14:167-

185, plates 10-11. 

Murray, G , and L. A. Boodle 

1889. A Systematic and Structural Account of the Genus 

Avrainvillea. Journal of Botany, British and Foreign, 27: 

67-72. 

Naegeli, C. 

1861. Bietrage zur Morphologie und Systematik der Ceramiaceae. Sitzungsberichte der Bayerischen Akademie 

der Wissenschaften in Miinchen, 2:297-415, plate 1. 

Norris, J. N. 

1976. Enteromorpha. In I. A. Abbott and G J. Hollenberg, 

Marine Algae of California, pages 73-77. Stanford, 

California: Stanford University Press. 

Okamura, K. 

1897. On the Algae from Ogasawarajima (Bonin Islands). Botanical Magazine (Tokyo), 11:1-16. 

Oliveira Filho, E. C. de 

1969. Algas marinhas do sul do Estado do Espirito Santo 

(Brasil), 1: Ceramiales. Faculdade de Filosofia, Ciencias e Letras, Universidade de Sao Paulo, Boletim 343, 

Serie Botanica, 26:1-277, plates A-D and 1-30, 1 

map. 

Papenfuss, G. F. 

1940. Notes on South African Marine Algae, I. Botaniska 

Notiser, Lund. 1940:200-226. 

1952. Notes on South African Marine Algae, III. The 

Journal of South African Botany, 17:167-188. 

1977. Review of the Genera of Dictyotales (Phaeophycophyta). The Bulletin of Japanese Society of Phycology, 

25(Supplement) :271-287. 

Pearse, A. S., and L. G. Williams 

1951. The Biota of the Reefs off the Carolinas. Journal of 

the Elisha Mitchell Scientific Society, 67(1): 133-161. 

Piccone, A. 

1884. Crociera del Corsaro alle hole Madera e Canarie del 

Capitano Enrico d'Albertis: Alghe. 60 pages, 1 plate. 

Genova: R. Instituto Sordo-Muti. 
222 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Richardson, W. D. 

1975. The Marine Algae of Trinidad, West Indies. Bulletin of the British Museum (NaturalHistory), (Botany), 

5(3):71-143, 27 plates. 

Scagel, R. F. 

1966. Marine Algae of British Columbia and Northern 

Washington, Part I: Chlorophyceae (Green Algae). National Museum of Canada, Bulletin 207, Biological Series, 74: viii + 257 pages. 

Schmitz, F. 

1893. Die Gattung Lophothalia J. Agardh. Berichte der 

Deutschen Botanischen Gesellschaft, 11(3):212-232. 

Schneider, C. W. 

1975. North Carolina Marine Algae, V: Additions to the 

Flora of Onslow Bay, Including the Reassignment 

of Fauchea peltata Taylor to Weberella Schmitz. 

British Phycological Journal, 10:129-138. 

Schnetter, R. 

1977. Notas sobre las especies Caribes del genero Botryocladia (Rhodymeniales, Rhodophyceae) con referenda especial a los taxa de al Costa Atlantica de 

Colombia, Anales del Instituto Investigaciones Marinas, 

Punta Betin, Santa Maria, Colombia, 9:73-80. 

Schnetter, R., and G. Bula Meyer 

1977. Rodoficeas nuevas para la costa atlantica de Colombia. Anales del Instituto Investigaciones Marinas, 

Punta Betin, Santa Marta, Colombia, 9:81-90. 

Schramm, A., and H. Maze 

1865. Essai de classification des algues de la Guadeloupe. [First 

edition], ii + 52 pages. Basse-Terre, Guadeloupe: 

Imprimerie du Gouvernement. 

1866. Essai de classification des algues de la Guadeloupe. [Second edition], v + 144 pages. Cayenne, French 

Guyana: Imprimerie du Gouvernement. 

Setchell, W. A. 

1925. Notes on Microdictyon. University of California Publications in Botany, 13:101-107. 

Setchell, W. A., and L. R. Mason 

1943. Goniolithon and Neogoniolithon. Proceedings of the National Academy of Sciences, 29(3/4) :87-92. 

Silva, P. C. 

1960. Codium (Chlorophyta) in the Tropical Western 

Atlantic. Nova Hedwigia, l(3/4):497-536, plates 

107-123. 

1972. Remarks on Algal Nomenclature, V. Taxon 21(1): 

199-212. 

Sosa, E. H. 

1977. Adiciones a la flora marina de Cuba. Ciencias 

Biologicas (Cuba), 1:158-160. 

Stafleu, F. A., V. Demoulin, W. Greuter, P. Hiepko, I. A. 

Linczevski, R. McVaugh, R. D. Meikle, R. C. Rollins, 

R. Ross, J. M. Schopf, and E. G. Voss 

1978. International Code of Botanical Nomenclature ... . xiv + 457 pages. Utrecht: Bohn, Scheltema & Holkema. 

Suarez, A. M. 

1973. Catalogo de algas Cubanas. Ciencias, Universidad de 

la Habana, series 8 (Investigaciones Marinas), 2: 

107 + 10 pages (Suplemento). 

Taylor, W. R. 

1928. The Marine Algae of Florida, with Special Reference to the Dry Tortugas. In Papers from the 

Tortugas Laboratory, 25. Carnegie Institution of 

Washington Publication, 379: v + 219, plates 1-37. 

1935. Botany of the Maya Area: Miscellaneous Papers, 

VII: Marine Algae from the Yucatan Peninsula. 

Carnegie Institution of Washington Publication, 461: 

115-124. 

1942. Caribbean Marine Algae of the Allan Hancock 

Expedition, 1939. Allan Hancock Atlantic Expedition, 

Report, 2: 193 pages. 

1945. Pacific Marine Algae of the Allan Hancock Expeditions to the Galapogos Islands. Allan Hancock 

Pacific Expeditions, 12: [vi] + iv + 528. 

1950. Reproduction oi Dudresnaya crassa Howe. Biological 

Bulletin, 99:272-284. 

1960. Marine Algae of the Eastern Tropical and Subtropical 

Coasts of the Americas, ix + [iii] + 870 pages. Ann 

Arbor: University of Michigan Press. 

1962a. Two Undescribed Species of Halimeda. Bulletin of 

Torrey Botanical Club, 89(3): 172-177. 

1962b. Marine Algae from the Tropical Atlantic Ocean, 

V: Algae from the Lesser Antilles. Contributions from 

the United States National Herbarium, 36(2):43-62, 

plates 1-4. 

1969. Notes on the Distribution of West Indian Marine 

Algae Particularly in the Lesser Antilles. Contributions from the University of Michigan Herbarium, 9(2): 

125-203. 

1976. A Check-List of Venezuelan Marine Algae. Boletin 

de la Sociedad Venezolana de Ciencias Naturales, 

22(132/133):71-101. 

Taylor, W. R., and C. H. Arndt 

1929. The Marine Algae of the Southeastern Peninsula 

of Hispaniola. American Journal of Botany, 16:651-

662. 

Taylor, W. R., and C. F. Rhyne 

1970. Marine Algae of Dominica. Smithsonian Contributions 

to Botany, 3: i + 16 pages. 

Trevisan, V.B.A. 

1845. Nomenclator algarum, ou collection des noms imposes aux 

plantes de la famille des algues. Volume 1, 80 pages. 

Padua. 

Tseng, C. K. 

1942. Studies on Chinese Species of Griffithsia. Papers oj 

the Michigan Academy of Science, Arts, and Letters, 

27(1941):105-116. 

1944. Marine Algae of Hong Kong, V: The Genus Herposiphonia. Papers of the Michigan Academy of Sciences, 

Arts, and Letters, 29(1943):55-65, plates 1-2. 
NUMBER 12 223 

Tsuda, R. T., and C. J. Dawes 

1974. Preliminary Checklist of the Marine Benthic 

Plants from Glover's Reef, British Honduras. Atoll 

Research Bulletin, 173:1-13. 

Ugadim, Y. 

1975. Algas marinhas bentonicas do litoral sul do Estado 

de Sao Paulo e do litoral do Estado do Parana, 

III: Divisao Rhodophyta(2): Cryptonemiales, Gigartinales e Rhodymeniales. Boletim da Botanica, 

Universidade de Sao Paulo, 3:115-163. 

Valet, G. 

1969. Contributions a l'etude des Dasycladales: Revision 

systematique. Nova Hedwigia, 17:575-637. 

van den Hoek. See Hoek, C. van den 

Vickers, A., and M.H. Shaw 

1905. Liste des algues marines de la Barbade. Annates des 

Sciences Naturelles, Paris, (Botanique), series 9, 1:45-

66. 

Wellington, G. M. 

1973. Additions to the Atlantic Benthic Flora of Costa 

Rica. Brenesia, 2:17-20. 

Williams, L. G. 

1948. Seasonal Alternation of the Marine Floras at Cape 

Lookout, North Carolina. American Journal of Botany, 135(10):682-695. 

Woelkerling, W. J. 

1976. South Florida Benthic Marine Algae: Keys and 

Comments. Sedimenta, 5: i + 148 pages. [Publication of the Comparative Sedimentology Laboratory, University of Miami.] 

Womersley, H.B.S. 

1967. A Critical Survey of the Marine Algae of Southern 

Australia, II: Phaeophyta. Australian Journal of Botany, 15:189-270. 

1978. Southern Australian Species of Ceramium Roth 

(Rhodophyta). Australian Journal of Marine and 

Freshwater Resources, 29:205-257. 

Wynne, M. J., and J. N. Norris 

1976. The Genus Colpomenia Derbes et Solier (Phaeophyta) in the Gulf of California. Smithsonian Contributions to Botany, 35: iii + 18. 

Yamada, Y. 

1931. Notes on Laurencia, with Special Reference to the 

Japanese Species. University of California Publications 

in Botany, 16(7): 185-310. 

1938. The Species of Liagora from Japan. Scientific Papers 

of the Institute of Algological Research, Faculty of Science, 

Hokkaido Imperial University, 2(1): 1-34. 

Yamada, Y., and S. Segawa 

1953. On Some New or Noteworthy Algae from Hachijo 

Island. Records of Oceanographic Works in Japan, new 

series, 1(1): 109-114. 

Young, D. N , B. M. Howard, and W. Fenical 

1980. Subcellular Localization of Brominated Secondary Metabolites in the Red Alga, Laurencia snyderae. 

Journal of Phycology, 16:182-185. 

Zanardini, G.A.M. 

1843. Saggio di classificazione naturalle delle Ficee. 64 + [i] 

pages, 1 plate. Venezia: Girolamo Tasso. 
The Red Alga Polysiphonia Greville 

(Rhodomelaceae) from Carrie Bow Cay 

and Vicinity, Belize 

Donald F. Kapraun 

and James N. Norris 

ABSTRACT 

Seven taxa of the red alga Polysiphonia Greville 

(Rhodomelaceae; Ceramiales) were found near 

Carrie Bow Cay and Twin Cays in the central 

region of the Belizean barrier reef. Polysiphonia 

exilis and P flaccidissima are new records for the 

Caribbean marine flora, P. atlantica, P. denudata, 

P. ferulacea, P. scopulorum var. villum, and P. sphaerocarpa are new to Belize. Polysiphonia atlantica is 

given as a new name for the preoccupied name 

P. macrocarpa Harvey. Most species occur as epiphytes on other algae, commonly on Sargassum 

and Dictyota. Polysiphonia ferulacea and P. sphaerocarpa are also found entangled in algal turfs, and 

P. scopulorum var. villum also occurs on mangrove 

roots. 

Introduction 

Only two species oi Polysiphonia Greville (1824) 

have been recorded previously from Belize (British Honduras): P. havanensis Montagne (Taylor, 

1935; 1960) and P. scopulorum Harvey (Tsuda and 

Dawes, 1974). In our present study oi Polysiphonia 

from Carrie Bow Cay and vicinity, we report six 

additional species and one variety new to the 

marine flora of Belize. 

Donald F. Kapraun, Department of Biology, University of North 

Carolina, Wilmington, N. C. 2840J. James N Norris, Department 

of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 

Several detailed studies of the taxonomy, distribution, and geographic variation oi Polysiphonia 

have increased our knowledge of its members. In 

the Pacific Ocean, studies have covered the coasts 

of North America (Hollenberg, 1942, 1944), including Mexico (Hollenberg, 1961), and the 

northern Gulf of California (Hollenberg and Norris, 1977); Hawaii (Menez, 1964); the central and 

western tropical Pacific (Hollenberg, 1968a, 

1968b); southern Australia (Womersley, 1979); 

and Japan (Segi, 1951). In the Atlantic Ocean, 

work has focused on the coasts of North Carolina 

and Texas (Kapraun, 1977, 1979); Brazil (Oliveira Filho, 1969); Portugal (Ardre, 1970); and the 

western Mediterranean (Lauret, 1967, 1970). 

We relied mainly on the following morphological features in distinguishing the taxa of Polysiphonia: number of pericentral cells; presence or 

absence of cortication; kind and position of origin 

of the attachment rhizoids; nature and position 

of trichoblasts; and nature of the spermatangial 

branchlets. Material for this study was collected 

from intertidal and subtidal habitats near Carrie 

Bow Cay and Twin Cays. These habitats are 

described and illustrated by Riitzler and Macintyre (herein: 9), Riitzler and Ferraris (herein: 

77) and Norris and Bucher (herein: 167). Specimens, which were obtained by J. Norris (MarchApril, 1976), J. Norris and K. Bucher (April 

1977), and D. Kapraun (March 1978), were 

mainly epiphytic on other algae. Collections are 

designated as follows: DK for D. F. Kapraun, JN 

225 
226 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

for J. N. Norris, and KB for K. E. Bucher. The 

prefix initials of the numbers cited correspond to 

the collector's field notebooks. Reproductive condition is designated by 0 for tetrasporangia, 9 for 

cystocarps, and 6 for spermatangia. Specimens 

studied are deposited in the Algal Collection, 

United States National Herbarium, Smithsonian 

Institution (US). 

ACKNOWLEDGMENTS.—We wish to thank K. E. 

Bucher for assistance in collecting and processing 

Belizean epiphytes, including Polysiphonia. We are 

grateful to G. J. Hollenberg, C. Schneider, and 

E. Menez for their reviews and comments on the 

manuscript. Finally, we thank D. Nicholson for 

discussion and review of the nomenclatural remarks, particularly on "P. macrocarpa." 

Key to the Species of Polysiphonia from Carrie Bow Cay and Vicinity 

1. Pericentral cells 4 (subgenus Oligosiphonia) 2 

Pericentral cells 5 or more (subgenus Polysiphonia) 6 

2. Branches arising in the axils of trichoblasts P. flaccidissima 

Branches developmentally replacing trichoblasts 3 

3. Rhizoids in open connection with pericentral cells 4 

Rhizoids cut off from pericentral cells 5 

4. Scar cells common in decumbent axes; trichoblasts in erect filaments 

branched 1-2 times P. atlantica 

Scar cells absent in decumbent axes; trichoblasts in erect filaments highly 

branched to several orders P. scopulorum var. villum 

5. Plants small, to 1.5 cm tall, and delicate, main filaments less than 100 jum 

in diameter; often epiphytic, erect filaments arising from a rhizoidal 

disc P. sphaerocarpa 

Plants larger, to 4 cm tall, and coarse, decumbent axes to 250 jum in 

diameter; erect filaments from entangled decumbent branches 

P. ferulacea 

6. Pericentral cells 5 (-6) P. denudata 

Pericentral cells 8 P. exilis 

Polysiphonia atlantica, new name 

FIGURES 107a-c 

Polysiphonia macrocarpa Harvey in Mackay, 1836:206 [non 

Polysiphonia macrocarpa (C. Agardh) Sprengel 1827:350].— 

Collins and Hervey, 1917:123.—B0rgesen, 1918:274, figs. 

272-276.—Taylor, 1928:184; 1960:578.—Oliveira Filho, 

1977:153.—Kapraun, 1977:317, figs. 14-21,57; 1979:107, 

figs. 10-12. 

DESCRIPTION.—Thalli minute, forming entangled mats to 1 cm tall; erect filaments sparsely 

subdichotomously branched; branches replacing 

trichoblasts in development; pericentral cells 4; 

erect filaments arising in unilateral fashion from 

prostrate axes; rhizoids remaining in open connection with pericentral cells. Tetrasporangia in 

long straight series in the branch tips. 

TYPE-LOCALITY.—Portstewart; Miltown Malbay, Ireland. 

DISTRIBUTION.—Tropical western Atlantic 

(Virgin Islands, Barbados, Curacao, Costa Rica, 

Cuba, Jamaica); temperate eastern Atlantic; Indian Ocean. 

SPECIMENS STUDIED.—Twin Cays: JN-6292 

(©), epiphytic on Ulva, 1 m depth, among mangroves on E side of eastern half of Twin Cays, 31 

Mar 1976. 

REMARKS.—Reluctantly we abandon the name 

Polysiphonia macrocarpa Harvey, because it has been 

widely applied to this neotropical species. Recently, Womersley (1979:471) noted that P. macrocarpa Harvey in Mackay (1836) is a later hom-
NUMBER 12 227 

50yum 

50/um 

FIGURE 107.—Polysiphonia atlantica: a, rhizoids in open connection with pericentral cells (JN6292); b, prostrate axis with unilateral development of erect filaments (JN-6292); c, tetrasporangia in long straight series (JN-6292). Polysiphonia denudata: d, rhizoids cut off at or near the 

end of pericentral cells (JN-6805); e, branch arising in axil of trichoblast (JN-6805);/, mature 

pericarp (JN-6805). 
228 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

onym of P. macrocarpa (C. Agardh) Sprengel 

(1827:350) [basionym: Hutchinsia macrocarpa C. 

Agardh, 1824:157]. Pending further studies, we 

propose a new name, P. atlantica Kapraun and J. 

Norris, for Harvey's taxon. Parke and Dixon 

(1976:537) pointed out that this taxon is in need 

of taxonomic and nomenclatural re-investigation. 

In this treatment we follow B0rgesen (1918), 

Ardre (1970), and Kapraun (1977, 1979) in their 

interpretation of P. macrocarpa Harvey. Reported 

elsewhere in the Caribbean as P. macrocarpa (for 

instance, Taylor, 1960), the occurrence of this 

taxon in Belize is not surprising. 

Polysiphonia macrocarpa (C. Agardh) Sprengel 

(1827) may well be an overlooked name for one 

of the several taxa described or reported in the 

Caribbean. Hutchinsia macrocarpa C. Agardh 

(1824) was originally described "In mari Antillarum, 'Port au Pray'." It is interesting that for 

other taxa in his Systema algarum, C. Agardh latinized localities, yet for H. macrocarpa he placed 

Port au Pray in quotation marks, leaving it in 

French form. By not using Latin for its locality, 

he may have wished to suggest something was 

wrong with the locality name. Could it be Port 

de Paix or Port au Prince (Haiti)? The type 

specimen (probably located at the Botanical Museum Lund) should be re-examined to determine 

the identity of this taxon, and compared to those 

taxa recorded from the Caribbean. 

Polysiphonia denudata (Dillwyn) 

Greville ex Harvey 

FIGURE 107 d-f 

Conferva denudata Dillwyn 1809:85. 

Polysiphonia denudata (Dillwyn) Greville ex Harvey in W. J. 

Hooker, 1833:332. 

Polysiphonia denudata (Dillwyn) Kiitzing, 1849:824; 1863:28, 

pl. 90.—Taylor, 1960:580.—Joly, 1965:221, pl. 47: figs. 

586-590.—Taylor and Bernatowicz, 1969:37.—Taylor, 

1969:183.—Lauret, 1970:123, pl. 1: figs. 1-20, pl. 2: figs. 

20-22.—Kapraun, 1977:321, figs. 38-42, 58; 1979:109, 

figs. 24-27.—Oliveira Filho, 1977:151.—Schnetter and 

Bula Meyer, 1977:89, fig. 17. 

DESCRIPTION.—Thalli dark red, to 3 cm tall 

from a discoidal base; erect filaments dichotomously branched; branches widely divergent, becoming decumbent in older parts and attached 

to the substratum; rhizoids cut off at or near the 

end of pericentral cells; pericentral cells 5 (-6); 

branches arising in axils of trichoblasts. Pericarps 

spherical, 200-250 jum in diameter. 

TYPE-LOCALITY.—Southhampton, England. 

DISTRIBUTION.—Atlantic: North America, Caribbean (Virgin Islands, Guadeloupe, Barbados, 

Netherland Antilles, Colombia), England, Portugal. Mediterranean. 

SPECIMENS STUDIED.—Carrie Bow Cay: JN6686 (©), epiphytic on Stypopodium zonale, 20-23 

m depth, fore reef slope, 27 Apr 1977; JN-6805 

(9, 0), 18 m depth, outer ridge of fore reef, legit 

JN and KB, 27 Apr 1977; JN-6842a (©), epiphytic on Dictyota, 10 m depth, spur and groove zone 

of inner fore reef, legit JN and KB, 29 Apr 1977. 

REMARKS.—The publication date of Dillwyn's 

British Confervae, in which Conferva denudata was 

first validly published, has been the subject of 

debate. We follow Dixon's (1960:309, 317) interpretation that publication date of the work is 

probably the late autumn (?) of 1809. 

Apparently a physiologically tolerant species, 

Polysiphonia denuda occurs in North American and 

European temperate waters (Taylor, 1962; Parke 

and Dixon, 1976), as well as in the American 

tropics. 

Polysiphonia exilis Harvey 

FIGURE 108 

Polysiphonia exilis Harvey, 1853:47.—Howe, 1920:570.—Taylor, 1928:183; 1960:581—Hollenberg, 1968b:200, figs. Ic, 

3c—Oliveira Filho, 1977:151. 

DESCRIPTION.—Thalli dark brown, with an extensive creeping system; trichoblasts formed in 

radial sequence in erect and prostrate axes, but 

erect filaments arising cicatrigenously and more 

or less unilaterally from adaxial scar cells; 

branches replacing trichoblasts developmentally; 

rhizoids cut off from pericentral cells; pericentral 

cells 8 (-10). Spermatangial branches oblong, 50 

X 150 jum, and sometimes subtended by trichoblasts, which arise one per segment. 

TYPE-LOCALITY.—Key West, Florida. 

DISTRIBUTION.—Tropical Atlantic (except Car-
NUMBER 12 229 

50,um 

FIGURE 108.—Polysiphonia exilis: a, prostrate axis with adventitious branch development, rhizoids 

cut off from pericentral cells (JN-6342); b, spermatangial branches, note trichoblasts arising 

one per segment (JN-6342). 

ibbean, up to now) and Pacific oceans. 

SPECIMENS STUDIED.—Carrie Bow Cay: JN6342a-c ((?), epiphytic on Dictyota dichotoma, ~20 

m depth, outer ridge of fore reef, 5 Apr 1976. 

REMARKS.—This is the first record of this species in the Caribbean Sea. The Carrie Bow Cay 

specimens apparently also represent the first spermatangial plants observed in this species (Taylor, 

1960; Hollenberg, 1968b). 

Polysiphonia ferulacea Suhr 

FIGURE 109 

Polysiphonia ferulacea Suhr in J. Agardh, 1863:980.—Collins 

and Hervey, 1917:124.—B0rgesen, 1918:277, figs. 277-

280.—Howe, 1920:570.—Taylor, 1928:183, pl. 24: figs. 

16-18, pl. 25: fig. 15, pl. 26: figs. 11, 15; 1960:578.— 

Oliveira Filho, 1969:128, pl. 24: figs. 141-144; 1977:151.— 

Taylor and Bernatowicz, 1969:36.—Taylor, 1969:183.— 

Taylor and Rhyne, 1970:15.—Yoneshigue-Beraga, 1972: 

27, pl. 6: figs. 30-31.—Richardson, 1975:130, pl. 23: fig. 

7.—Kapraun, 1977:320, figs. 32-37. 

DESCRIPTION.—Thalli erect, to 4 cm tall, from 

a discoidal base; alternately branched from a 

distinct main axis; older plants with extensive 

decumbent branches secondarily attached by rhizoids cut off from pericentral cells; branches replacing trichoblasts in development; pericentral 

cells 4; branches characteristically basally constricted; spermatangial branches cylindrical, 50 
230 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

50yum 

FIGURE 109—Polysiphonia ferulacea: a, b, variation in erect filaments on the same plant in respect 

to cell height-width ratios (DK-s.n., 21 Mar 1978); c, rhizoids cut off from pericentral cells 

(DK-s.n., 21 Mar 1978); d, tetrasporangia in spiral series (DK-s.n., 21 Mar 1978); e, spermatangial branches with characteristic enlarged sterile tip cells (JN-6938). 
NUMBER 12 231 

X 150 jum, with 1-2 conspicuous, thick-walled 

sterile tip cells. Tetrasporangi a in spiral series in 

branch tips. 

TYPE-LOCALITY.—Atlanti c Coast of Mexico. 

DISTRIBUTION.—Widespread in subtropical and 

tropical seas; Caribbean: Puerto Rico, Virgin 

Islands, St. Kitts, Antigua , Guadeloupe , Dominica, Barbados, Trinidad, Venezuela, Curacao, 

Costa Rica, Cayma n Islands, Jamaica , and Hispaniola. 

SPECIMENS STUDIED.—Carri e Bow Cay: JN -

6334 (9) and JN-6335 (2), epiphytic on Dictyota 

cervicornis, 20 m depth, outer ridge of fore reef, 5 

Apr 1976; JN-6335 (©), epiphytic on Lobophora 

variegata, 25 m depth, fore reef slope, 5 Apr 1976; 

JN-6410 (©), epiphytic on Valonia ventricosa, and 

JN-6413, epiphytic on Caulpera verticillata, both 

from high-relief spur and groove zone (vicinity of 

IMSWE transect, between 300-400 m markers), 

5-6 m depth, legit J N and KB, 27 Apr 1977; JN -

6691b, 20 m depth, fore reef slope, 27 Apr 1977; 

JN-6938 (6), epiphytic on Udotea, 10 m depth, 

low-relief spur and groove zone of inner fore reef 

("10 m S of IMSW E transect line between the 

400-500 m markers), legit J N and KB, 23 Apr 

1977; JN-6987, 0.5 m depth, entangled with turf 

oi Jania, Centroceras, an d Wrangelia, reef crest in 

front of Carrie Bow Cay Laboratory, legit J N and 

KB, 22 Apr 1977; DK-s.n. (6, ©), eulittoral on 

cinderblock wall, 21 Ma r 1978. 

REMARKS.—Th e close resemblance an d hence 

possible taxonomic confusion between Polysiphonia 

ferulacea Suhr in J. Agardh and P. sparsa (Setchell) 

Hollenberg has been previously noted (Hollenberg, 1968a; Kapraun , 1977). In the present 

study, specimens of P. ferulacea varied greatly in 

trichoblast development and cell length to width 

ratios of erect filaments. Although some individuals had short segments as in P. sparsa, the presence of characteristic spermatangial branches 

with sterile tip cells seems to confirm our identification as P. ferulacea. 

Polysiphonia flaccidissima Hollenberg 

FIGURE 110 

Polysiphonia flaccidissima Hollenberg, 1942:783, figs. 8, 19; 

1961:351, pl. 2: fig. 2; 1968a:63, figs. 2a, 11.—Brauner, 

1975:128, figs. 2-4.—Abbott and Hollenberg, 1976:688, 

fig. 634.—Hollenberg and Norris, 1977:4, fig. 2E.—Kapraun, 1979:107, figs. 8-9. 

DESCRIPTION.—Thalli minute , extensive creeping system giving rise to erect filaments up to 1.5 

cm tall; pericentral cells 4; erect and prostrate 

axes with radial development of branch primordia; branches arising in axils of trichoblasts; rhizoids cut off from the proximal end of pericentral 

cells. Tetrasporangi a to 75 jum in diameter, in 

spiral series in branch tips; spermatangial 

branches oblong, 40 X 170 jum, lacking sterile tip 

cells and subtended by a trichoblast; matur e 

pericarps oval, 150 X 175 jum. 

TYPE-LOCALITY.—Lagun a Beach, Orang e 

County, California. 

DISTRIBUTION.—Atlantic : North Carolina, Gulf 

of Mexico; Pacific: Southern California, Hawaii, 

Phoenix and Marshall Islands. 

SPECIMENS STUDIED.—Carri e Bow Cay: JN -

6351 (6\ 2) and JN-6353 (©), both epiphytic on 

Sargassum hystrix, 16-20 m depth, outer ridge and 

fore reef slope, 5 Apr 1976; JN-6842b (©, ?, 6), 

epiphytic on Dictyota, 10 m depth, spur and 

groove zone of inner fore reef, legit J N and KB, 

29 Apr 1977. Twin Cays: JN-6886 (?, ©), epiphytic on Anadyomene stellata, 1 m depth, amon g mangroves on E side of eastern half of Twin Cays, 26 

Apr 1977. 

REMARKS.—Polysiphonia flaccidissima is recorded 

for the first time in the Caribbean Sea. Superficially, it resembles P. havanensis sensu B0rgesen 

(1918:266) (see Kapraun , 1977:316), which is 

widely distributed in the Caribbean; however, P. 

flaccidissima is clearly distinguished from this 

taxon by its rhizoids, which are cut off from the 

proximal end of the pericentral cells rather than 

occurring in open connection with pericentral 

cells as in P. havanensis. Taylor's (1935) specimens 

identified as P. havanensis could possibly belong 

here; they need to be re-examined to verify its 

presence in Belize. 

Womersley (1979:479) compared descriptions 

of Polysiphonia flaccidissima (Hollenberg, 1942, 

1968a; Abbott and Hollenberg, 1976) with southern Australian and Mediterranean specimens 

identified as P. sertularioides (Grateloup) J. Agardh 
232 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 110.—Polysiphonia flaccidissima: a, tetrasporangia in spiral series (JN-6351); b, rhizoids 

cut off from pericentral cells (JN-6351); c, spermatangial branches (JN-6351); d, developing 

pericarps, note developing vegetative branch in axil of trichoblast (JN-
NUMBER 12 233 

and concluded that the two species may be synnonymous. Until the exact relationship of P. flaccidissima to P. sertularioides has been determined, 

we refer material from our present study to P. 

flaccidissima, the name well established in the New 

World literature (see for instance, Hollenberg, 

1942, 1961, 1968a; Hollenberg and Norris, 1977; 

Brauner, 1975; Kapraun, 1979). 

Polysiphonia scopulorum var. villum 

(J. Agardh) Hollenberg 

FIGURE 111 

Polysiphonia villum]. Agardh, 1863:941. 

Lophosiphonia villum (J. Agardh) Setchell and Gardner 1903: 

329. 

Polysiphonia scopulorum var. villum (J. Agardh) Hollenberg, 

1968a:81, fig. 7A.—Hollenberg and Norris, 1977:14, fig. 

8B., figs. 640-641. 

DESCRIPTION.—Thalli minute, to 0.5 cm tall, 

extensive creeping system giving rise to erect filaments; pericentral cells 4; erect filaments arising 

endogenously in unilateral fashion, at regular 

intervals of 2-4 segments, from prostrate axes; 

erect filaments with conspicuous trichoblasts dichotomously branching up to the fourth order; 

erect branches replacing trichoblasts in development; rhizoids in open connection with pericentral cells. Tetrasporangia in long straight series. 

TYPE LOCALITY.—" .. . ad littus [sic] americae 

tropicae" (J. Agardh, 1863:941); probably the 

Pacific Coast of Mexico (Hollenberg 1968a:81). 

DISTRIBUTION.—Wildely distributed in northern Pacific and western Atlantic oceans. 

SPECIMENS STUDIED.—Carrie Bow Cay: JN6357 (2), epiphytic on Lobophora variegata, 20-25 

m depth, fore reef slope, 5 Apr 1976. Twin Cays: 

DK-s.n. (©), eulittoral, epiphytic on mangroves, 

24 Mar 1978. 

REMARKS.—Although this taxon is usually described as having few, slightly branched trichoblasts, our material closely resembles the illustrations in Hollenberg (1968a) showing highly 

branched, conspicuous trichoblasts in erect filaments. Tsuda and Dawes (1974) list Polysiphonia 

scopulorum in their report on Glover's Reef, Belize, 

but make no comments on infraspecific relationships. The presence of var. villum in the Caribbean 

is now confirmed. 

Polysiphonia sphaerocarpa B0rgesen 

FIGURE 112 

Polysiphonia sphaerocarpa B0rgesen, 1918:271, figs. 267-271.— 

Taylor, 1960:576.—Hollenberg, 1968a:87, figs. 21, 26.— 

Taylor, 1969:183.—Taylor and Rhyne, 1970:15.—Kapraun, 1977:318, figs. 26-31, 56. 

DESCRIPTION.—Thalli small epiphytes, to 1.5 

cm tall from a discoidal base; branches dichotomous, becoming decumbent and attached to the 

substratum by rhizoids cut off from pericentral 

cells; branches replacing trichoblasts in development; pericentral cells 4. Spermatangial branches 

cylindrical, 50 X 250 jum, lacking sterile tip cells, 

with subtending trichoblasts; pericarps spherical, 

250 jum diameter, with ostiole of large cells. Tetrasporangia in spiral series. 

TYPE-LOCALITY.—Stor e Nordsidebugt, St. 

Thomas, U.S. Virgin Islands. 

DISTRIBUTION.—Widespread in tropical Atlantic, including the Caribbean (Virgin Islands, Antigua, Dominica, Barbados, Hispaniola), and 

tropical Pacific oceans. 

SPECIMENS STUDIED.—Carrie Bow Cay: JN6746 (?), JN-6747 (6), JN-6748 (©), and JN-6755 

($, ©), all epiphytic on Dictyota, 8 m depth, patch 

reef, SW of Carrie Bow Cay, legit JN and KB, 25 

Apr 1977; JN-6847a (5), epiphytic on Sargassum, 

10 m depth, spur and groove zone of inner fore 

reef, legit JN and KB, 29 Apr 1977; JN-6911 

($), epiphytic on Dictyota, 9-10 m depth, lowrelief spur and groove of inner fore reef vicinity 

of IMSWE transect (Riitzler and Macintyre, 

herein), legit JN and KB, 23 Apr 1977; DK-s.n. 

(6, ©), eulittoral on reef, 21 Mar 1978. 

REMARKS.—This species is apparently a member of the "pan-tropical flora" (Kapraun, 1977). 

Several Carrie Bow Cay specimens have bifurcate 

spermatangial branches (Figure 1126), which are 

atypical for this species. Hollenberg (1968a:80, 

fig. 6F) reported similar variation in male gametophytes of another species of Polysiphonia, P. scopulorum Harvey from Hawaii. 
234 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 111.—Polysiphonia scopulorum var. villum: a, prostrate axis with rhizoids in open connection 

with pericentral cells (DK-s.n., 24 Mar 1978); b, erect filament with conspicuous trichoblasts 

and tetrasporangia in straight series (DK-s.n., 24 Mar 1978). 
NUMBER 12 235 

-.1 a o J o,'< 

o?°°a "S A ^ r

 Jo0 

50/jm 

0°Q, 

,£p o ° OD D O OO ° O O e V 

r> J SotsO Do"-'-oo° 0 » & 

Oj.O O <>Q - Q _" • "/JO 

50jum 

FIGURE 112.—Polysiphonia sphaerocarpa: a, tetrasporangia in spiral series (DK-s.n., 21 Mar 1978); 

b, spermatangial branches (DK-s.n., 21 Mar 1978); c, mature pericarp (JN-6911); d, rhizoids 

cut off from pericentral cells (DK-s.n., 21 Mar 1978). 
236 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Conclusions 

Our collections from Carrie Bow Cay add seven 

taxa of Polysiphonia to the previously reported 

marine flora of Belize. Five species belong to the 

subgenus Oligosiphonia—P. atlantica, P. ferulacea, P. 

flaccidissima, P. scopulorum var. villum, and P. sphaerocarpa—and two to the subgenus Polysiphonia—P. 

denudata and P. exilis. Polysiphonia exillis and P. 

flaccidissima are found in the Caribbean Sea for 

the first time. Prior to this study only P. havanensis 

and P. scopulorum were known from Belize, neither 

of which appeared in our collections. Although 

we cannot exclude the possibility that P. scopulorum reported by Tsuda and Dawes (1974) from 

Glover's reef, only 25 km east of Carrie Bow Cay, 

may belong to var. villum. 

Polysiphonia atlantica is a new name proposed for 

P. macrocarpa Harvey, a later homonym for P. 

macrocarpa (C. Agardh) Sprengel. The identity of 

C. Agardh's Hutchinsia macrocarpa, described from 

the Antilles but never reported since, remains 

unknown. 

Although members of the genus Polysiphonia 

occur predominantly in tropical waters, P. ferulacea ranges into the subtropical Atlantic and P. 

atlantica and P. denudata extend even into the 

temperate Atlantic. The latter two species, at 

least, are remarkable for their physiological tolerance of such a large range of environmental 

conditions. 

All our collections were made close to Carrie 

Bow Cay, during March and April. We expect 

that material sampled during other seasons and 

at different localities along the barrier reef, on the 

atolls, and along the mainland coast will contain 

interesting new finds and will help solve taxonomic problems. 

Literature Cited 

Abbott, I. A., and G. J. Hollenberg 

1976. Marine Algae of California, xii + (2) + 827 pages. 

Stanford, California: Stanford University Press. 

Agardh, C. A. 

1824. Systema algarum. xxxvii + 312 pages. Lund: Berling. 

Agardh, J. G. 

1863. Species, genera el ordines algarum, seu descriptiones succinctae specierum generum et ordinum, quibus algarum 

regnum constituitur. Volume 2, part 3, pages [2] + 

787-1291. Lund G.W.K. Gleerup. 

Ardre, F. 

1970. Contribution a l'etude des algues marines du Portugal, I: La Flore. Portugaliae Acta Biologica (B), 

10(1/4):5-423, 56 plates. 

B0rgesen, F. 

1918. The Marine Algae of the Danish West Indies, Part 

IV [sic]: Rhodophyceae (4). Danske Botanisk Arkiv, 

3(ld):241-304. 

Brauner, J. 

1975. Seasonality of Epiphytic Algae on Zostera marina at 

Beaufort, North Carolina. Nova Hedwigia, 26:125-

133. 

Collins, F. S., and A. B. Hervey 

1917. The Algae of Bermuda. Proceedings of the American 

Academy of Arts and Sciences, 53(1): 1-195. 

Dillwyn, L. W. 

[1809.] British Confervae 87 pages. London: W. Phillips. 

Dixon, P. S. 

1960. Taxonomic and Nomenclatural Notes on the Florideae, II. Botamska Notiser, 113(3):295-319. 

Greville, R. K. 

1824. Flora Edinensis. lxxxi -I- 478 pages. Edinburgh: W. 

Blackwood. 

Harvey, W. H. 

1853. Nereis Boreali-Americana . . . Part II: Rhodospermeae. 

[First issue, ii +] 258 pages, plates 13-36. Washington, D.C: Smithsonian Institution [London: 

John Van Voorst]. [Third issue, 1853 [1858], 

Smithsonian Contributions to Knowledge, 5(5): ii + 258 

pages, plates 13-36.] 

Hollenberg, G. J. 

1942. An Account of the Species oi Polysiphonia on the 

Pacific Coast of North America, I: Oligosiphonia. 

American Journal of Botany, 29:772-785. 

1944. An Account of the Species of Polysiphonia on the 

Pacific Coast of North America, II: Polysiphonia. 

American Journal of Botany, 31:474-483. 

1961. Marine Red Algae of Pacific Mexico, Part 5: The 
NUMBER 12 237 

Genus Polysiphonia. Pacific Naturalist, 2(6):345-375. 

1968a. An Account of the Species of Polysiphonia of the 

Central and Western Tropical Pacific Ocean, I: 

Oligosiphonia. Pacific Science, 22(l):56-98. 

1968b. An Account of the Species of the Red Alga Polysiphonia of the Central and Western Tropical Pacific 

Ocean, II: Polysiphonia. Pacific Science, 22(2): 198-

207. 

Hollenberg, G. J., and J. N. Norris 

1977. The Red Alga Polysiphonia (Rhodomelaceae) in the 

Northern Gulf of California. Smithsonian Contributions to the Marine Sciences, 1: iii + 21 pages. 

Hooker, W.J. 

1833. Mosses, Heptiae, Lichens, Characeae and Algae. 

In J. E. Smith, The English Flora, edition 2, 5(1): x 

+ 4-1- 432 pages. London: Longman, Rees, Orme, 

Brown, Green & Longman. [Also issued in volume 

2 of W. J. Hooker, The British Flora.] 

Howe, M. A. 

1920. Class 2, Algae In N. L. Britton and C. F. Millspaugh, The Bahama Flora, pages 553-618. New 

York: Published by the authors. 

Joly, A. B. 

1965. Flora marinha do litoral norte do Estado de Sao 

Paulo e regioes circunvizinhas. Boletim Faculdade de 

Filosofia, Ciencias e Letras da Universidade de Sao Paulo, 

Boletim (Botanica), 21:1-393, plates 1-59. 

Kapraun, D. 

1977. The Genus Polysiphonia in North Carolina, U.S.A. 

Botanica Marina, 20:313-331. 

1979. The Genus Polysiphonia (Rhodophyta, Ceramiales) 

in the Vicinity of Port Aransas, Texas. Contributions 

in Marine Science, 22:105-120. 

Kiitzing, F. T. 

1849. Species algarum. vi + 922 pages. Lipsiae: F. A. 

Brockhaus. 

1863. Tabulae phycologicae. Volume 13, [i] + 31 pages. 

Nordhausen. 

Lauret, M. 

1967. Morphologie, phenologie, repartition des Polysiphonia marins du littoral languedocien, I: Section 

Oligosiphonia. Naturalia Monspeliensia (Botany), 18: 

347-373. 

1970. Morphologie, phenologie, repartition des Polysiphonia marins du littoral languedocien. Naturalia 

Monospeliensia (Botany), 21:121-163. 

Mackay, J. T. 

1836. Flora Hibernica. Volume 2, xxxiv + [3] + 354 + 279 

pages. Dublin: Wm. Curry Jun. & Co. 

Meiiez, E. G. 

1964. The Taxonomy of Polysiphonia in Hawaii. Pacific 

Science, 18(2):207-222. 

Oliveira Filho, E. C. de 

1969. Algas marinhas do sul do Estado do Espirito Santo 

(Brasil), I: Ceramiales. Universidade de Sao Paulo, 

Faculdade de Filosofia, Ciencias e Letras, Boletim, 343 

(Serie Botanica, 26): 277 pages, plates A-D + 1-

30, 1 map. 

1977. Algas marinhas bentonicas do Brasil. [iii] + 407 pages. 

Sao Paulo, Brasil: Universidade de Sao Paulo 

(Instituto de Biociencias). 

Parke, M., and P. S. Dixon 

1976. Check-List of British Marine Algae—Third Revision. Journal of the Marine Biological Association, U.K., 

56:527-594. 

Richardson, W. D. 

1975. The Marine Algae of Trinidad, West Indies. Bulletin of the British Museum (Natural History), Botany, 

5(3):73-143, plates 16-27. 

Schnetter, R., and G. Bula Meyer 

1977. Rodoficeas nuevas para la Costa Atlantica de 

Colombia. Anales del Instituto de Investigaciones Marinas Punta Betin, Santa Maria, Colombia, 9:81-90. 

Segi, T. 

1951. Systematic Study of the Genus Polysiphonia from 

Japan and its Vicinity. Journal of the Faculty of 

Fisheries, Prefectural University of Mie, 1(2): 169-272. 

Setchell, W. A., and N. L. Gardner 

1903. Algae of Northwestern America. University of California Publications, Botany, 1:165-418. 

Sprengel, K.PJ. 

1827. Classis 24: Cryptogamia. In Caroli Linnaei . . . Systema vegetabihum, editio decima sexta, 4:592 pages. 

Gottingen: Sumtibus Librariae Dieterichianae. 

Taylor, W. R. 

1928. The Marine Algae of Florida, with Special Reference to the Dry Tortugas. Papers from the Tortugas Laboratory, 25. Carnegie Institution of Washington Publication, 379: v + 219, plates 1-37. 

1935. Botany of the Maya Area, Miscellaneous Papers, 

VII: Marine Algae from the Yucatan Peninsula. 

Carnegie Institution of Washington Publication, 461: 

115-124. 

1960. Marine Algae of the Eastern Tropical and Subtropical 

Coasts of the Americas, ix + [iii] + 870 pages. Ann 

Arbor: University of Michigan Press. 

1962. Marine Algae of the Northeastern Coast of North America. 

Second edition with corrections, viii + 509 pages. 

Ann Arbor: University of Michigan Press. 

1969. Notes on the Distribution of West Indian Marine 

Algae Particularly in the Lesser Antilles. Contributions from the University of Michigan Herbarium, 9(2): 

125-203. 

Taylor, W. R., and A. J. Bernatowicz 

1969. Distribution of Marine Algae about Bermuda. 

Bermuda Biological Station for Research, Special Publication, 1: 42 pages. Bermuda: St. George's West. 

Taylor, W. R., and C. F. Rhyne 

1970. Marine Algae of Dominica. Smithsonian Contributions 

to Botany, 3: i + 16. 
238 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Tsuda, R. T., and C. J. Dawes 

1974. Preliminary Checklist of the Marine Benthic 

Plants from Glover's Reef, British Honduras. Atoll 

Research Bulletin, 173:1-13. 

Womersley, H.B.S. 

1979. Southern Australian Species of Polysiphonia Greville (Rhodophyta). Australian Journal of Botany, 

27(4):459-528. 

Yoneshigue-Braga, Y. 

1972. Flora marinha bentonica da Baia de Guanabara 

e Cercanias, III: Rhodophyta; 3 Ceramiales. Publicacao do Instituto de Pesquisas da Marinha (Rw de 

Janeiro), 065:1-49, plates 1-9, map. 
Hydroidea (Cnidaria: Hydrozoa) 

from Carrie Bow Cay, Belize 

Barry W. Spracklin 

ABSTRACT 

Forty-five species of hydroids were found living 

on the reef and in the lagoon at Carrie Bow Cay. 

Collections were made each spring (March-June) 

from 1974 to 1978. Information was gathered on 

distribution, abundance, substrates, depth ranges, 

and reproductive maturity. Halcordyle disticha 

(Goldfuss), Halecium bermudense Congdon, and Haloptens carinata Allman are the dominant hydroids 

in the outer reef areas and on patch reefs. Dynamena cornicina McCrady was common in all areas 

of the Thalassia (turtle grass) beds. Eleutheria dichotoma Quatrefages, Halecium speciosum Nutting, 

and Egmundella grandis Fraser are recorded from 

the Caribbean Sea for the first time. Plumularia 

species is probably a growth form of P. floridana 

Nutting, the gonosome of which is undescribed. 

Halecium species is undescribed; corynid species, 

corymorphid species, and sertularid species are 

undescribed and appear to belong to new genera. 

Introduction 

This report on the hyrdroids of the Belizean 

barrier reef is based on collections made near 

Carrie Bow Cay during the spring months of each 

year from 1974 to 1978. Collections and observations were made at nearly all areas during each 

of the five years. Many of the hydroids have been 

photographed in situ and almost all have been 

observed alive, prior to fixation. 

No major collection of hydroids has previously 

been made from the coast of Belize and little 

Barry W. Spracklin, Zoology Department, University of New Hampshire, Durham, NH 03824. 

material from this area of the Caribbean Sea was 

included by Nutting (1900, 1904, 1915), Fraser 

(1944, 1947), Gemerden-Hoogeveen (1965), or 

Vervoort (1968). In addition, most of the previous 

collections were made over a short period of time 

during a single year and were examined only 

after preservation. The major exception was Wedler's (1975) ecological study of the hydroids of 

Santa Marta (Colombia), which covered a 16-

month period. 

ACKNOWLEDGMENTS.—I thank all of my numerous diving partners. S. Earle, P. M. Kier, M. E. 

Rice, and K. Riitzler collected some of the hydroids; sponges were identified by K. Riitzler. C. 

W. Walker read the manuscript and made numerous useful suggestions. 

Methods 

During the spring months (March-June) of a 

5-year period (1974-1978) 103 collections of hydroids were made by skin diving and SCUBA 

diving. Conventional transecting techniques 

could not be applied readily because of the patchy 

distribution of hydroids in many areas, the cryptic 

occurrence of many hydroids, and the problem of 

identifying species with certainty underwater. 

Since, in most cases, a portion of each colony was 

collected for identification, the number of observations of each species has been used for abundance estimates. The large number of hydroids 

(33 species) discovered on dead gorgonians and 

scleractinean corals does not necessarily indicate 

a preference for these substrates. Many small 

hydroids are commonly found on algae, but these 

are readily overlooked. 

239 
240 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Man y of the hydroids were photographed underwater or on the island before fixation, an d 

most of the athecate hydroids were drawn from 

photographs. Th e thecate hydroids were drawn 

from photographs and by camer a lucida from 

fixed specimens. A reference collection is being 

prepared. 

Annotated Species List 

Suborder ATHECATA 

Family CORYMORPHIDAE 

Corymorphid species. Figure 113a. 

This hydroid is very similar to Corymorpha symmetrica Hargitt from the Phillipines; however, 

solitary polyps grow out of the ends of branches 

of the gorgonians Muriceopsis flavida (Lamarck), 

Pseudopterogorgia acerosa (Pallas), Gorgonia flabellum Linnaeus, and G. ventalina Linnaeus rather 

tha n on coralline algae. A matur e hydranth has 

16 to 22 proximal tentacles (usually coiled at 

the tip), 25 to 30 oral tentacles and seven to 

nine nematocyst clusters, each bearing one to 

three lateral gonophores with no evidence of 

tentacles. Although corymorphid sp. and C. 

symmetrica clearly belong in the family Corymorphidae , these two species could easily be 

placed in a separate genus since they both have 

nematocyst clusters, attach to solid substrates, 

and lack tentacles on the gonophores or medus a 

buds. 

Family TUBULARIIDAE 

Ectopleura grandis Fraser. Figure 113c. 

T h e identification was mad e from a colony 

that had only immatur e medusa buds. 

Family HALOCORDYLIDAE 

Halocordyle disticha (Goldfuss). Figure 113c. 

Family CORYNIDAE 

Corynid species. Figure H3d,e. 

This hydroid is common in the spur and groove 

zones on the encrusting sponge Monanchora barbadensis Hechtel and in the reef crest it was 

found on the boring sponge Cliona caribbea 

Carter. In the outer reef areas it was found on 

an unidentified sponge, and once on an unidentified alga growing on a dead gorgonian. It 

has an oral whorl of four tentacles and a second 

whorl of eight to ten clusters of three tentacles 

each. Th e gonophores, which develop between 

the two whorls, were released in finger bowls 

immediately after collection with four apical 

nematocyst clusters, but no tentacles. 

Family ELEUTHERIIDAE 

Eleutheria dichotoma Quatrefages. Figure 113/. 

First record from the Caribbean Sea. Th e medusa has been recorded from the Mediterranean Sea, British Isles, Sweden, and France 

(Brinkmann-Voss, 1970). Only a single polyp 

with seven capitate tentacles was found on a 

blade of turtle grass. 

Family ZANCLEIDAE 

Zanclea costata Gegenbaur. Figure 113g. 

T h e perisarc at the base of the hydrant h was 

annulated in the material on Sargassum and 

smooth in the material from the reef crest. 

Family CLAVIDAE 

Corydendrium parasiticum (Linnaeus). Figure 113/?. 

Not listed by Vervoort (1968) from the Caribbean, this hydroid was recorded by Wedler 

(1975) from Colombia. 

Turritopsis nutricula (McCrady). Figure 113?. 

Family BOUGAINVILLIIDAE 

Garveia humilis (Allman). Figure 113;'. 

Family EUDENDRIIDAE 

Eudendrium attenuatum Allman. Figure \l3k,J. 

Eudendrium eximium Allman. Figure 114a. 

Myrionema hargitti (Congdon). Figure 1146. 

Suborder THECATA 

Family CALICELLIDAE 

Egmundella grandis Fraser. Figure 114c. 

First Caribbean Sea record. 

Family HALECIIDAE 

Halecium bermudense Congdon. Figure \\4d. 

Halecium namum Alder. Figure 114c. 
NUMBER 12 241 

FIGURE 113.—Athecate hydroids, Corymorphidae-Eudendriidae: a, corymorphid species, some 

proximal tentacles removed to show the nematocyst clusters, X 10; b, Eclopleura grandis, some 

proximal tentacles removed to show the medusa buds, X 40; c, Halocordyle disticha, X \2; d, 

corynid species, X 45; e, corynid species, oral view, X 60;/, Eleuthena dichotoma, contracted after 

fixation, X 60; g, Zanclea costata, X 30; h, Corydendnum parasiticum, X 10; i, Turntopsis nutncula, X 

10; j , Garveia humilis, X 50: k, Eudendrium attenuatum, main stem and side branch, X 8; /, Eudendrium 

attenuatum, male, X 32. 
242 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 114.—Athecate hydroids, Eudendriidae (a,b), and thecate hydroids, Calicellidae-Haleciidae: (c-j) a, Eudendrium eximium, X 10; b, Myrionema hargitti, main stem and branch of female 

colony, X 13; c, Egmundella grandis, X 130; d, Halecium bermudense, X 30; e, Halecium namum, X 40; 

/, Halecium speciosum, X 50; g, Halecium speciosum, female gonophore, X 50; h, Halecium species, 

X 30; i, Halecium species, female gonophore, X 100; 7, Ophiodissa mirabilis, X 60. 
NUMBER 12 243 

FIGURE 115.—Thecate hydroids, Campanulariidae-Sertulariidae: a, Clytia hemisphaerica, X 30; 

b, Clytia laxa, X 25; c, Clytia noliformis, X 25; d, Obelia dichotoma, X 40; e, Hebella calcarala, X 80;/, 

Hebella venusta, X 50; g, Scandia mutabilis, X 80; h, Cmdoscyphus marginalus, X 15; i, Diphasia tropica, 

X 30; / Dynamena cornicina, X 40. 
244 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 116.—Thecate hydroids, Sertulariidae-Plumulariidae: a, Dynamena crisiodes, X 20; b, 

Sertularella parvula, X 50; c, Sertularella speciosa, X 15; d, Serlulana stookeyi, X 25; e, Sertularia 

turbinata, X 25; / sertularid species, X 40; g, sertularid species, showing tentacle-like nematophore, X 130; h, Antenella gracilis, X 70; i, Antenella quadriaurita, X 70; / Halopteris carinata, X 36; 

k, Halopteris cannata, male gonophore, X 36. 
NUMBER 12 245 

FIGURE 117.—Thecate hydroids, Plumulariidae: a, Halopteris diaphana, X 35; b, Monostaechas 

quadridens, X 15; c, Plumularia halecioides, X 35; d, Plumulana halecioides, gonophore from base of 

stem or stolon, X 35; e, Plumularia margaretta, X 70; / Plumularia setacea, X 60; g, Plumularia species, 

X 50; h, Aglaophenia latecarinata, X 130; i, Aglaophenia pluma pluma, X 130. 
246 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Halecium speciosum Nutting. Figure 114/g. 

Described from the western coast of North 

America, this is the first record from the Atlantic Ocean. 

Halecium species. Figure 1 \4h,i. 

The colonial growth form and strongly flared 

hydrophoral margin match the material that 

Vervoort (1968) referred to Halecium reflexum 

Stechow. Cornelius (1975a) reduced//, reflexum 

to a junior synonym of H. labrosum Alder. Vervoort's material was not reproductive and the 

single gonophore present on my material is 

similar to H. nanum (Figure 114c) and not at all 

like the gonophore of H. labrosum. My material 

of H. nanum from the Carrie Bow Cay area has 

the normal growth form and hydrophoral margin, and thus H. species probably represents an 

undescribed species. 

Ophwdissa mirabilis (Hincks). Figure 114;'. 

Cornelius (1975a) reduced Ophiodissa caciniformis (Ritchie) to a junior synonym of 0. mirabilis. 

Family CAMPANULARIIDAE 

Clytia hemisphaerica (Linnaeus). Figure 115a. 

Listed as Campanularia {Clytia) johnstoni Alder 

by Vervoort (1968); see Millard (1966) for 

synonymy. 

Clytia laxa Fraser. Figure 1156. 

Listed as Laomedea (Phialidium) laxa by Vervoort (1968). The genus Laomedea was reduced 

by Cornelius (1975b) to a junior synonym of 

Obelia. 

Clytia noliformis (McCrady). Figure 115c. 

Obelia dichotoma (Linnaeus). Figure ll5d. 

Listed by Vervoort (1968) as Laomedea (Obelia) 

congdoniHarght. Cornelius (1975b) referred numerous described species in the genus Obelia to 

three valid species. 

Family LAFOEIDAE 

Hebella calcarata (A. Agassiz). Figure 115c. 

Hebella venusta (Allman). Figure 115/. 

Scandia mutabilis (Ritchie). Figure 115g. 

Family SERTURLARIIDAE 

Cnidoscyphus marginatus (Allman). Figure 115a. 

Diphasia tropica Nutting. Figure 115z. 

Dynamena cornicina McCrady. Figure 115/ 

Dynamena crisioides (Lamouroux). Figure 116a. 

Sertularella parvula (Allman). Figure 1166. 

Sertularella speciosa Congdon. Figure 116c. 

The distinct hydrothecal base present in all of 

the shallow water specimens (31 m or less) was 

totally absent in the colony from 67 m. 

Sertularia stookeyi Nutting. Figure 116a". 

Sertularia turbinata Lamouroux. Figure 116c. 

Sertularid species. Figure 116/g. 

Superficially, this hydroid appears to be a normal sertularid except that, in addition to the 

hydranth, each hydrotheca contains a large 

tentacle-like nematophore. In living material, 

either the nematophore or the hydranth, or 

both at once, may extend out of the hydrotheca. The operculum is composed of a single 

plate. This hydroid was collected in 1974 and 

not seen again until 1978 when it was found to 

be abundant on the underside of coral rubble 

in a different area of the reef crest. No reproductive material was evident. 

Family PLUMULARIIDAE 

Antenella gracilis Allman. Figure 116a. 

Antenella quadriaurita Ritchie. Figure 116z. 

Halopteris carinata Allman. Figure 116//:. 

Halopteris diaphana (Heller). Figure 117a. 

Monostaechas quadridens (McCrady). Figure 1176. 

Plumularia halecwdes (Alder). Figure 117c/ 

Plumularia margaretta (Nutting). Figure 117c. 

Plumularia setacea (Linnaeus). Figure 117/ 

Plumularia species. Figure 11Ig. 

This hydroid is probably conspecific with P. 

floridana Nutting; however, the gonophores 

were not described and the annulations were 

somewhat different than in my material. A 

more reliable identification will have to await 

an examination of the type material. 

Aglaophenia latecarinata Allman. Figure 117a. 

Aglaophenia pluma pluma (Linnaeus). Figure 117z. 

Distribution 

In order to facilitate collection and recording 

of data, the lagoon and reef around Carrie Bow 

Cay were subdivided into 10 areas. The lagoon 

includes patch reefs, mangrove islands, and the 
NUMBER 12 247 

turtle grass (Thalassia testudinium Banks ex Konig) 

beds. Th e reef was divided into the back reef, reef 

crest, high- an d low-relief spur and groove, sand 

trough, outer ridge, and fore reef slope zones 

(compare Riitzler and Macintyre , herein: 9). Th e 

dat a in Tabl e 18 are the combined results of 103 

collections mad e over a five-year period (1974-

1978) and show the distribution, abundance , substrates, reproductive maturity, an d depth ranges 

of the hydroids found in these 10 areas. 

PATCH REEFS.—Th e patch reefs have the most 

diverse hydroid fauna (22 species) in the Carrie 

Bow Cay area an d even adjacent reefs vary considerably in composition and abundanc e of species. Density of hydroid colonies on the patch 

reefs is generally greater than that in comparabl e 

habitats in the spur and groove, sand trough, and 

outer ridge zones of the barrier reef. Halocordyle 

disticha, Halecium bermudense, Halopteris carinata, Antenella gracilis, and Plumularia setacea are found 

commonly on dead gorgonians and corals on 

patch reefs. Cnidoscyphus marginatus is unusually 

abundan t at one of the sites, overgrowing large 

areas of coral rock. A single hydroclad oi Antenella 

quadriaurita, the only record of this species in the 

present collection, occurred on a piece of coral 

rubble covered with numerous hydrocladia of A. 

gracilis. 

MANGROVE ISLANDS.—Dynamena cnsioides is 

found abundantl y and exclusively on the mangrove roots at Twi n Cays and We e We e Cay. 

Myrionema hargitti and Halecium bermudense are concentrated on the mangrove roots and banks at 

the north end of the channel dividing Twin Cays. 

Halecium bermudense, Clytia hemisphaerica, Obelia dichotoma, Dynamena cornicina, and Plumularia halecioides are common on mangrove roots and algae 

at other locations around the mangrove islands. 

TURTLE GRASS BEDS.—I n view of the large 

numbe r of turtle grass blades examined and the 

enormous surface area available for settlement it 

is surprising that only 12 species of hydroids are 

found here. Halecium bermudense and Dynamena cornicina are the only species commo n throughout 

and both appea r to be most abundan t at the 

eastern sides of Twi n Cays an d the patch reefs. 

Plumularia haleciodes, and to a lesser extent P. 

setacea, are also commo n on the east side of Twin 

Cays. Th e only specimens of Eleutheria dichotoma 

(a single polyp) and Sertularia stookeyi (one colony) 

were collected in this habitat. In man y areas of 

the lagoon, algae replace hydroids as the major 

epihytic growth on turtle grass. 

BACK REEF.—Dynamena cornicina and Plumularia 

halecioides are common on the turtle grass in the 

back reef area between Carrie Bow Cay and the 

reef crest (0-0.5 m). In 1974 Myrionema hargitti was 

found in a single patch of sand and rubble near 

the reef crest. Since then, this hydroid has steadily 

increased in abundanc e and in 1978 occurred in 

almost all the sand patches in this area, growing 

attached to small pieces of rubble in the sand. At 

the north end of the island, Halecium speciosum 

Nuttin g was collected on the calcareous green 

alga Halimeda species at a depth of approximately 

0.5 m. Halecium species and H. nanum are also 

present here on Halimeda species; Halecium bermudense was collected from the brown alga Turbinaria 

species. 

REE F CREST.—Most of the nine hydroids from 

the reef crest were discovered on the underside of 

coral rubble slightly deeper than the normal lowtide level and were represented, for the most part, 

by small stolonal colonies. Th e only common 

hydroid in this area is "sertularid species." A few 

hydranths of "corynid species" were collected 

from one of the reef-crest channels on the boring 

sponge, Cliona caribbaea. Halocordyle disticha, Plumularia inermis, and P. setacea were collected in 2 

m of water on dead gorgonians off the end of the 

reef crest south of Carrie Bow Cay. 

HIGH-RELIE F SPUR AND GROOVE ZONE.—Onl y 

four hydroids were found amon g the high-relief 

coral spurs (2-6 m). Th e most common one in 

this zone is "corynid species," which occurs unde r 

overhanging coral on the sponge Monanchora barbadensis. Halocordyle disticha and "corymorphid species" were collected on gorgonians at the seaward 

edge of the high spurs, whereas Halecium bermudense was found in crevices amon g the corals. 

LOW-RELIE F SPUR AND GROOVE ZONE.—Non e 

of the 14 species collected here between depths of 

7 an d 12 m is abundant, although Halocordyle 

disticha, Halecium bermudense, Halopteris carinata, and 
248 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

V 

IS > 

B 

o 

cj5 

5 

T> 

TJ 

u bin 

S 

o 

G 

O 

T J ase 

-O 

w 

TJ 

'3 

y hydr 

U « Bow 

.ii Carr 

o 

0 

3 

; f l 

'C -Dis 1. 

CO 

— rABLE 

OJ rare 

II sent 

-O 

rt 

II 

1 ates 

B 

'£ 

V 

u 

c 

C3 

"0 

c 3 

of ab 

c 

o entati 

-r. 

1-(rep 1978 

r-. 

O) 

CO ason 

-

be 

c ;ollecti 

o 

c 

II 

# 

<u" 

<« bstr 

3 1/5 lar icu part 

RI 

c 

o 

c 

OJ r 

u C CL 

II 

.. undant 

_Q 

rt 

II on, 

B 

£ 

o 

u 

II 

co 

"5 

C 

0 

'33 

= occa 

c 

o vati ser 

J2 

O ring 

3 

-a tes 

T I test 

JS and 

U 

nj U gives 

GO 

TJ 3 

03 

3 < 

TJ OJ 

s !_ ores o 

^ - £ matur uctive prod 

OJ 

O 

-a 

_o per 

c o nformat 

Q. gono 

-C 

TJ 

OJ llect 

o 

3 

£ 

-a droi 

>-

J G 

tfl :ime th 

J= u 

O Z £• 

"5 fl'S 

°TJ 3 

h P « 

CJ U |H 

OH CL E 

spiojpAj-i 

SUBIUOSJOS 

pUB SJBJOD pB3Q 

utnssvBivs 

§UI}BO|j[ 

SJOOJ 3AOJ§UBJAJ 

s3pB|q mssvjvqj^ 

3BSJ Y 

sdojs JSSJ-SJO^ 

sSp u J3jn o 

qSnojj puBg 

3UOZ 3AOOJ§ 

puB jnd s Moq 

3UOZ 3AOOJ§ 

puB jnd s qS| H 

JS3J0 J33"£J 

J33J >|3Bg 

spsq vissvjm/j^ 

SpUB[Sl 3AOJ§UBJ/\J 

SJSSJ q^JBj 

CL 

t/3 

r^ r-^ co — r-~ ~ co —• i— - H ^-i 

-CM-HM i r !

0

i-.co-'iocn(Ncocnco l f !a)S ni:r)-

H 

IIIIOOIIIIIIIIIOI^II I * CM CN io lOCMcn-^r^cMCMCMc o uo in co * 

> - > s 

& 2 

>C3 S , i 

Jun 

CM 

1 

ri * 

G 

3 

•* 

a>! * -

G C 

3 3 

O £• — O >^ >^ 

CM TM CM « « w 

l I l l l l I l l I l 

^K^ky * I S> K> S> S> S/1 S^ ^ 

A n n I rS /S rS rS rS rS i*S 

2 b 

C . pi eg 

rt 

a 

< 

CM CM 

I I I I X I I I I I I 

I I I I I I X I X I X 

I I I X I I l l l I I 

I I I I I X I X 

X I I I I X l I 

X X I I I X I l 

I IXIXIX I I I IXXXXXXXI X 

XXI I 

XXI X 

X X I 

X I X 

I I I CM I I —i CO CM I 

CO I CO N I I |CMCM | j I 

CO CM CO CM I |CMCM | I I I 

CO I CO CO I I I I CM I —' | 

CM I CM * I I I I I I I I 

I I I * l I CM i i i i i i 

I I I I I I I I I I I -

I I I I - I I I I I I I 

llllll|CM|CM|C O 

CO | CO | I I OI N I CM I | 

CO CM | | 

« I I I 

co | | -

N

 I I I 

I I I I 

CM CM — CM 

CO CM | | 

M

 I I I 

CO CM | | 

CM | CM 

CM | CM 

I I I 

I I I 

I I I 

I I I 

CM | | 

CO | | 

CM | | 

t/3 Cl 

.ii "S 

y -e 

CL "5 

• 2 K 

^3 J" 

•f" !S -E 

o ii ^ 

O fe3 tt! 

s J 1'E s 

•c -2 .» S 

.2 "fe £ R 

•J3 S 

s E .a .a § £ i 1 ^ =1 •« s .uis-s s § 
NUMBER 12 249 rate 

t o 

XI 

3 

C/3 

a 

< 

_C 4J 

•Si bo--—s 

CL c G 

Q >-

0 *3 C* 

TJ a' C 

2 T J 5 

•fc 2 2 

a, a E 

u 

spiojpAj-j 

SUBIUOSJOS 

p UB SJBJOD pB3 Q 

utnssvSxvg 

§UUBO[j 

SJOOJ SAOJSUBJAJ 

sspBjq VISSV]VI{JL 

SBSJ V 

3dojS J33J-3JO j 

3§p u jsm o 

qSnoj j puBg 

3UOZ 3AOOJ§ 

puB jnd s Moq 

3UOZ 3AOOJ§ 

puB jnd s qSiH 

JS3JD J33>J 

J33 J T\OVQ 

spaq mssvjvqj^ 

SpUBJSI 3AOJ§UBJAJ 

SJ33J qojB j 

s 

CL 

C/2 

CO 

1 

CM 

O 

l_ 

a 

< 

CM 

1 

S « CM S CM m d 

1 " O 1 O 1 1 

o mo o 

rt G 

o **< 

—< CM 

1 1 

* * * * * Q . CL 

< < •n cn 

1 X 1 1 1 1 1 

X X 1 X X X X 1 

X 

X 

X 

1 

CM 

CM 

-

1 

1 

1 

1 

O J 

CO 

CO 

s 

5 

"5 

'-5 

.« 

o 

1 1 1 1 1 1 1 

I 1 1 1 1 X X 

1 1 1 1 1 X 1 

1 1 1 1 1 1 1 

CM | | CO | CM | 

CM | | CM | | | 

CM - H | CM | CM | 

| | | CM | CM j 

1 1 1 1 1 1 1 

CM | - | - | | 

1 1 1 1 1 M 1 

1 1 1 1 1 * 1 

| | | | | CO • * 

1 1 1 •* 1 ^ 1 

3 

a 

S r, 2 

Hebella calcarata H. venusta Scandia mutabilis 

Cnidoscyphus mar 

Diphasia tropica Dynamena cornici, D. crisioides 

CO 

1 

CM 

d 

# 

1 

1 

1 

1 

X 

X 

CM 

1 

1 

| 

1 

C J 

1 

O J 

1 

CM 

J3 

3 5 

5! 

a 

"£: 

t>3 

<X 

1 

UD 

# 

1 

X 

1 

1 

1 

1 

O J 

O J 

O l 

O J 

•3 

" 1 

. o 

So 

05 

•n 2 

© 1 

O 

>-

t3 

o 

1 1 

i n ^ 

d co 

i i i- » 

O CO 

> N 

cd 

* * 

cn 

1 1 1 

« CO C N 

CO — CM 

1 1 1 

i n - ^ C M 

CM 2 

O J 1 

Q .

 rt * 

CM C M 

1 1 1 

<JD 

1 

O 

c 

3 

O J 

1 

U 

CL 

< O l 

1 

CO —• 

— CO 

1 1 

•n o 

* * 

1 1 

CO 

1 

*£ 

G 

3 

!—S 

O l 

2 

O J 

1 

1xxxxxxxxxx x 

1 X 

1 1 

X 1 

1 1 1 

1 1 1 

1 X 1 

1 X X X 1 

1 1 

1 1 

1 1 

1 c* 

1 1 

1 1 

1 CM 

- 1 

1 1 

1 CM Sertularia slookeyi S. turbinata 

1 CM | 

1 1 1 

1 CM | 

1 1 1 

1 1 1 

CO | | 

1 1 1 

1 CM | 

i 1 1 

| CO -

[fl 

cj Sertularid speci 

Antenella gracilis A. quadriaunta 

1 1 1 

J 1 1 

I X I 

1 X 1 

CO CM | 

CO CM | 

CO CM CM 

CO CM | 

1 1 1 

1 1 1 

1 1 1 

1 CM | 

1 1 1 

CO CM | 

s 

•"§ 

« "« Haloptens cannat H. diaphana Monostaechas que 

1 

X 

X 

X 

O J 

co 

CO 

co 

O l 

J V 

. 5 

a 

.a 

IK 

5 

a 

1 X 

1 X 

I X 

1 X 

1 CM 

1 CM 

— O J 

1 CM 

1 1 

1 CM 

1 1 

1 CM 

1 CM 

— CO P. margaretta P. setacea 

1 

1 

1 

1 

O J 

O J 

1 

1 

1 

1 

1 

1 

1 

O J 

V 

'u 

V 

a 

O J 

d 

i 

o 

>. 

Bl 

1 

1 

X 

1 

1 

1 

1 

1 

1 

1 

1 

1 

1 

1 

1 

1 

<3 

a 

,g 

S 

-s: 

co 

i 

<x> 

* 

1 

1 

1 

1 

1 

X 

1 

1 

O J 

O J 

a 

5 

s 

"5, 

« 

s 

<L 

i n * 

co * 

CO 

r~- <£> 

o * 

O J * 

Cn * 

•—i r -

O l 

r-» CO 

OJ co 

O J 

• * <y> 

• * • * 

a> o 

CO o 

CM — 

o cn 

CM — 

CM CM 

« 

c 

o tfl ' £ 

CJ y 

CJ 3 2 

CL O 

5i u 

1 M <— 

O O 

u u 

Xi X Total num Total num 
250 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

"corymorphid species" were conspicuous during 

most dives. The small number of species present 

in this large area is surprising in view of the 

greater diversity in deeper water at the adjacent 

sand trough and in shallower water at the lagoon 

patch reefs. 

SAND TROUGH.—The eight collections made 

here range from a depth of approximately 13 m 

at the upper edge of the inner reef slope to about 

28 m at the bottom of the trough. This area is 

called the sand trough, but the 22 species collected here are all from the flanks of the trough, 

not from the sand fields at the bottom. Although 

the hydroid fauna is more diverse here than in 

the spur and groove zones, none of the species is 

particularly abundant. Halocordyle disticha, Halecium bermudense, Halopteris carinata, and "corymorphid species" are again the most frequently observed hydroids. 

OUTER RIDGE.—Seventeen hydroids were collected mostly on dead gorgonians and corals on 

the outer ridge at 13-15 m, a depth equivalent to 

that of the upper level of the sand-trough zone. 

Halocordyle disticha, Halecium bermudense, Halopteris 

carinata, and "corymorphid species" are the most 

common hydroids. 

FORE-REEF SLOPE.—Seven collections were 

made on the fore-reef slope between 15 and 46 m 

in depth. Halecium bermudense, Cnidoscyphus marginatus, and Halopteris carinata are common down to 

31 m, whereas Halocordyle disticha was collected 

only occasionally in the upper part of this zone, 

to a depth of 18 m, and "corymorphid species" 

was not seen at all. The only hydroid found below 

31 m was Sertularella speciosa, collected during a 

bounce dive at 67 m. 

SARGASSUM FLOATS.—Hydroids from this substrate, collected whenever the currents carried 

large amounts of free-floating brown algae of the 

genus Sargassum past the island, are not local in 

origin. Although present on almost every piece of 

alga, only seven pieces were recorded from six 

collections. Clytia hemisphaerica, Obelia dichotoma, 

and Aglaophenia latecannata abound in all of the 

collections. Aglaophenia latecarinata is not found on 

any other substrate. 

Discussion and Conclusions 

Forty-five species of hydroids representing 15 

families and 29 genera are identified and figured 

(Figure 113-117). Three species, Eleutheria dichotoma Quatrefages, Halecium speciosum Nutting, and 

Egmundella grandis Fraser, have not previously 

been reported from the Caribbean. A number of 

unidentified and apparently undescribed species 

have been collected. Halecium species appears to 

be undescribed. Three other hydroids, a corymorphid, a corynid, and sertularid, cannot easily 

be placed in any known genera and await description (Spracklin, in prep.); in the body of this 

paper they are referred to as "corymorphid species," "corynid species," and "sertularid species." 

At Carrie Bow Cay, a number of hydroids are 

sharply restricted in depth of occurrence or substrate preference. For example, Halocordyle disticha 

and "corymorphid species" do not occur below 

18 m. Zanclea costata is found only on the reef crest 

and on Sargassum at a maximum depth of approximately 0.2 m, Aglaophenia latecarinata utilizes only 

Sargassum, and Dynamena crisioides grows abundantly and exclusively on mangrove roots. In 

contrast, Clytia hemisphaerica was collected from 

all six of the substrates examined and Halecium 

bermudense, the most widely distributed hydroid, 

was collected from every substrate examined except Sargassum and from every area except the 

reef crest. Only Halecium bermudense commonly 

utilizes sponges as a substrate. 

The depth limitation of Halocordyle disticha, to 

18 m or less, is particularly interesting since it is 

reported to 29 m along the eastern United States 

(Gosner, 1971), but only to approximately 8 m in 

Colombia (Wedler, 1975). This species is abundant in shallow water and is one of the few 

hydroids easily recognizable underwater, so that 

it could not have been overlooked at greater 

depths. Because this is also the first hydroid to 

settle on the mooring lines each spring, with 

colonies appearing usually in three to four weeks, 

it is very suitable for field studies and manipulation. 

The three lagoon areas near Carrie Bow Cay— 
NUMBER 12 251 

the turtle grass beds, mangrove islands, and patch 

reefs—contain hydroid populations that overlap 

only slightly. Of the 26 species collected in the 

lagoon only six {Halecium bermudense, Clytia hemisphaerica, Obelia dichotoma, Dynamena cornicina, Plumularia haleciodes, and P. setacea) are found in all 

three areas. Only Halecium bermudense, and Dynamena cornicina are common throughout the lagoon. 

The back reef, reef crest, and high-relief spur 

and groove zones have relatively distinct hydroid 

populations with a different species dominating 

each zone. In the back reef, Myrionema hargitti is 

the dominant hydroid, whereas "sertularid species" is most abundant in the reef crest, and 

"corynid species" in the high-relief spur and 

groove zone. In contrast, the outer reef areas 

appear much more uniform. The same group of 

four species {Halocordyle disticha, Halecium bermudense, Halopteris carinata, and "corymorphid species") dominates the low-relief spur and groove, 

sand trough, and outer ridge zones, with Halecium 

bermudense, Cnidoscyphus marginatum, and Haloptens 

carinata also the most abundant hydroids on the 

fore-reef slope. With the exception of the fore-reef 

slope, the outer reef zones can be considered as a 

single area for future hydroid studies. 

Literature Cited 

Brinkman-Voss, A. 

1970. Anthomedusae/Athecata (Hydrozoa, Cnidaria) of 

the Mediterranean, Part 1: Capitata. Edizione della 

Stazione Zoologica di Napoli, 39:1-96, figures 1-106, 

plates 1-11. 

Cornelius, P.F.S. 

1975a. A Revision of the Species of Lafoeidae and Haleciidae (Coelenterata: Hydroida) Recorded from 

Britain and Nearby Seas. Bulletin of the British 

Museum (Natural History), Zoology, 28:373-426, figures 1-14. 

1975b. The Hydroid Species of Obelia (Coelenterata, Hydrozoa: Campanulariidae), with Notes on the Medusa Stage. Bulletin of the British Museum (Natural 

History), Zoology, 28(6):249-293, figures 1-5. 

Fraser, CM . 

1944. Hydroids of the Atlantic Coast of North America. 640 

pages, 94 plates. Toronto: University of Toronto 

Press. 

1947. Hydroids of the 1939 Allan Hancock Caribbean 

Sea Expedition. Allan Hancock Atlantic Expedition, 

4:1-24, plates 1-3. 

Gemerden-Hoogeveen, G.C.H. van 

1965. Hydroids of the Caribbean: Sertulariidae, Plumulariidae and Aglaopheniidae. Studies on the 

Fauna of Curacao and Other Caribbean Islands, 22(84): 

1-87, figures 1-45. 

Gosner, K. L. 

1971. Guide to Identification of Marine and Estuarine Invertebrates. 693 pages. New York: Wiley-Interscience. 

Hargitt, C. W. 

1924. Hydroids of the Philippine Islands. The Philippine 

Journal of Science, 24(4):467-507, plates 1-6. 

Millard, N.A.H. 

1966. The Hydrozoa of the South and West Coasts of 

South Africa, Part III: The Gymnoblastea and 

Small Families of Calyptoblastea. Annals of the 

South African Museum, 48(18):427-487, figures 1-

15, plate 1. 

Nutting, C. 

1900. American Hydroids, Part I: The Plumularidae. 

285 pages, 34 plates. United States National Museum, 

Special Bulletin, 4(1): Washington, D.C: Smithsonian Institution. 

1904. American Hydroids, Part II: The Sertularidae. 

325 pages, 41 plates. United States National Museum, 

Special Bulletin, 4(2): Washington, D.C: Smithsonian Institution. 

1915. American Hydroids, Part III: The Campanularidae 

and Bonneviellidae. United States National Museum, 

Special Bulletin, 4(3): 126 pages, 27 plates. Washington, D.C: Smithsonian Institution. 

Spracklin, B. W. 

In prep. New and Unusual Species of Hydroids from 

Carrie Bow Cay, Belize. 

Vervoort, W. 

1968. Report on a Collection of Hydroida from the 

Caribbean Region, Including an Annotated 

Checklist of Caribbean Hydroids. Zoologische Verhandelingen, 92:1-124, figures 1-41. 

Wedler, E. 

1975. Okologische Untersuchungen an Hydroiden des 

Felslitorals von Santa Marta (Kolumbien). Helgolander Wissenschaftliche Meeresuntersuchungen, 27:324-

363, figures 1-34. 
Medusae (Cnidaria) from Carrie Bow Cay, Belize 

Ronald J. Larson 

ABSTRACT 

Qualitative and quantitative data on medusae 

from reef, lagoon, and inshore areas near Carrie 

Bow Cay, Belize are provided. Of 71 species of 

medusae collected, 57 were found near the reef, 

46 in the lagoon, and 21 inshore. Medusae constituted about 10% of the biomass in the samples 

at the reef and lagoon stations, and 70% inshore. 

Liriope tetraphylla was the most abundant species 

at reef and lagoon stations. Owing to its abundance and its voracious feeding habits, it is considered to be the ecologically most important 

species in these two areas. 

Introduction 

Coral reefs are known to have an especially 

diverse plankton "community" (Wood and Johannes, 1975). Recent quantitative studies 

(Moore and Sander, 1976; Renon, 1977) have 

examined the dominant groups of mostly lower 

trophic level species (for instance, copepods), but 

not the less numerous, although nonetheless important, groups of the higher trophic level such 

as medusae. Mayer (1900, 1904) and Kramp 

(1953) have shown that a rich medusaen fauna is 

associated with coral reefs. These studies, however, lack accurate quantitative data because they 

were made prior to the development of plankton 

net flowmeters. 

The following study was undertaken to determine the species composition, abundance, and 

biomass of medusae over a coral reef. Samples 

taken from a large volume of water over a period 

Ronald J. Larson, Department of Biology, University of Victoria, 

British Columbia, Canada V8W2Y2. 

of several months improved the accuracy of biomass and species abundance data. Comparative 

samples were taken in the near-by lagoon and 

inshore waters. 

ACKNOWLEDGMENTS.—I wish to thank J. D. 

Ferraris, Mount Desert Island Biological Laboratory, for allowing me to examine plankton samples that she collected near Carrie Bow Cay, and 

K. S. Larson, Smithsonian Institution, for helpful 

suggestions and for typing the manuscript. 

Study Area and Methods 

Sixty-two quantitative plankton samples were 

taken at three stations (Figure 118) over reefs and 

in the near-reef lagoon at Carrie Bow Cay, and 

near the mainland of Belize, between January 

and April, 1978. A net having 0.5 m2 opening, 

560 /xm mesh diameter, and a flowmeter was 

used. Surface tows were made between 1730 h 

and 1830 h for 15-25 min, each filtering an 

average of 250 m3. Qualitative plankton samples 

were taken on the reef flat of Carrie Bow Cay. 

Medusae were also collected by beach seine, by 

night light and dip net, and while diving. Quantitative samples obtained by J. D. Ferraris during 

April-May 1976 and May-June 1977 (Ferraris, 

herein: 143) were also examined. 

Whole plankton samples were fixed in 10% 

formalin. Medusae were sorted later, identified 

and counted. Wet weights were obtained by 

gently pouring the preserved plankton, or medusaen sample onto a 56 jum Nitex cloth, blotting it 

from below and weighing it to the nearest 0.01 g. 

This method did not damage even very fragile 

specimens. Dry weights were not measured because the samples are intended for further taxo253 
254 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

88°10'W 

Blue 

Ground 

Range J 

, J/J) 

Stewart nl '•-•--' 

Cay :

u

l ^WeeWee 

Cay 

Tobacco Cay 

Tobacco Cay 

Entrance 

16°50'N 

South Water Cay 

Carrie Bow Cay 

Curlew Bank 

N 

5 km. 

FIGURE 118.—Map of study area near Carrie Bow Cay, indicating collecting stations for 

medusae (1 = reef; 2 = lagoon; 3 = inshore). 

nomic studies. 

The stations were located as follows: (1) reef— 

1 km S of Carrie Bow Cay, about 1.5 km from 

the shelf edge, 5-8 m depth, patch reefs and sand 

bottom; (2) lagoon—between South Water Cay 

and Twin Cays, about 3 km from the shelf edge, 

3-6 m depth, mostly seagrass bottom; (3) inshore— 1 km E of the town of Stann Creek (Dangriga), approximately 25 km from the shelf edge, 

10 m depth, sand bottom. 

Order ANTHOMEDUSAE 

Family CORYNIDAE 

Dipurena halterata (Forbes) 

Sarsia angulala (Mayer) 

Cladosarsia capitata Bouillon 

Family TUBULARIIDAE 

Euphysora gracilis (Brooks) 

Species List and Distribution 

(Letters denote unidentified or new taxa) 

Reef Lagoon Inshore 

+ + 

+ + 

+ + 

Euphysora species a 

Euphysora species b 

Hybocodon species a 

Propachycordyle species a 

Vannuccia forbesi (Mayer) 

Genus A species a 

Family HALOCORDYLIDAE 

Halocordyle disticha (Goldfuss) 

Reef Lagoon Inshore 

+ + -

+ + -

+ - -

+ - -

-I- + -
NUMBER 12 255 

Reef Lagoon Inshore Reef Lagoon Inshore 

Family CODONIDA incertae sedis 

Pachycordyle species a — + 

Cnidocodon sp. + — 

Genus B species a + — 

Genus C species a + — 

Genus D species a + — 

Family ZANCLEIDAE 

Zanclea prolifera Uchida and Sugiura + + 

Family ZANCLEOPSIDAE 

Zancleopsis dichotoma (Mayer) + + 

Family CLADONEMATIDAE 

Cladonema radiatum Dujardin + + 

Family ELEUTHERIIDAE 

Staurocladia vallentini (Browne) -I- — 

Family CYTAEIDIDAE 

Cytaeis tetrastyla Eschscholtz -I- + 

Family HYDRACTINIIDAE 

Podocoryne minuta (Mayer) 

Podocoryne ocellata (Agassiz and 

Mayer) 

Podocoryne species a 

Podocoryne species b 

Family BOUGAINVILLIIDAE 

Bougainvillia carolinensis (McCrady) 

Bougainvillia frondosa Mayer 

Koellikerina elegans (Mayer) 

Genus E species a + 

Family PANDEIDAE 

Amphinema rugosum (Mayer) + 

Amphinema turrida (Mayer) + 

Merga violacea (Agassiz and Mayer) + 

Stomoloca pterophylla Haeckel + 

Genus F species a + 

Order LEPTOMEDUSAE 

Family DIPLEUROSOMATIDAE 

Dipleurosoma collapsum (Mayer) — 

Family LAODICEIDAE 

Laodicea brevigona Allwein 

Family MITROCOMIDAE 

Tiaropsidium roseum (Maas) 

Family CAMPANULARIIDAE 

"Obelia'' sp. 

Phialidium spp. + + 

Family LOVENELLIDAE 

Eucheilota paradoxica Mayer — + 

Family PHIALUCIIDAE 

Phialucium carolinae (Mayer) + + 

+ 

+ 

+ 

+ 

+ 

— 

+ 

+ 

+ 

— 

+ 

-

+ 

+ 

-I- + 

+ + 

+ + 

+ 

Family EIRENIDAE 

Eirene lactea (Mayer) 

Helgicirrha schulzei Hartlaub 

Family AEQUOREIDAE 

Aequorea macrodaclyla (Brandt) 

Order LIMNOMEDUSAE 

Family OLINDIADIDAE 

Cubaia aphrodite Mayer 

Olindias tenuis (Fewkes) 

Family PROBOSCIDACTYLIDAE 

Proboscidactyla ornata (McCrady) 

Order TRACHYMEDUSAE 

Family GERYONIIDAE 

Linope tetraphylla (Chamisso and 

Eysenhardt) 

Family RHOPALONEMATIDAE 

Aglaura hemistoma Peron and 

Lesueur 

Amphogona apstemi (Vanhoffen) 

Persa incolorata McCrady 

Rhopalonema velatum Gegenbaur 

Order NARCOMEDUSAE 

Family AEGINIDAE 

Aegina citrea Eschscholtz 

Solmundella bitentaculata (Quoy and 

Gaimard) 

Family SOLMARISIDAE 

Pegantha rubiginosa? (Kolliker) 

Pegantha triloba Haeckel 

Solmaris corona? (Keferstein and 

Ehlers) 

Family CUNINIDAE 

Cunina globosa? Eschscholtz 

Cunina octonana McCrady 

Cunina peregrina Bigelow 

Order CUBOMEDUSAE 

Family CARYBDEIDAE 

Carybdea alata Reynaud 

Carybdea marsupialis (Linnaeus) 

Carybdea species a 

Family CHIRODROPIDAE 

Chiropsalmus quadrumanus (Muller) 

Order CORONATAE 

Family LINUCHIDAE 

Linuche unguiculata (Schwartz) 

Family NAUSITHOIDAE 

Nausilhoe punctata Kolliker 

+ 

+ 

+ 

+ 

+ 

+ 

+ 
256 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Reef Lagoon Inshore 

Order SEMAEOSTOMEAE 

Family PELAGIIDAE 

Chrysaora quinquecirrha (Desor) 

Pelagia noctiluca (Forskal) 

Family ULMARIDAE 

Amelia aurita (Linnaeus) 

+ 

+ + 

Order RHIZOSTOMEAE 

Family CASSIOPEIDAE 

Cassiopea frondosa (Pallas) 

Cassiopea xamachana R.P. Bigelow 

Totals 

Reef Lagoon Inshore 

57 46 21 

Results 

Seventy-one species of medusae were collected 

during this study. Thirteen species are sufficiently 

abundant to compare their distribution across the 

Belizean shelf (Table 19). Six species, Aglaura 

hemistoma, Amphogona apsteini, Nausithoe punctata, 

Sarsia angulata, Solmundella bitentaculata, and Zanclea 

prolifera were collected only near the reef and 

near-by lagoon. Carybdea alata juveniles appeared 

in nearly one-third of the reef and outer lagoon 

samples. Since these medusae were collected often 

with oceanic plankton, for example, gelatinous 

colonial radiolarians and the blue-green alga Oscillatoria sp., and because Bigelow (1938) noted 

that C. alata and representatives of the genera 

Amphogona, Nausithoe, Solmundella, and Zanclea are 

found in oceanic waters, the above seven species 

of medusae may prefer the oceanic conditions 

near the shelf edge. 

Species that dominated inshore samples but 

were rare or missing over the outer shelf were 

Bougainvillia carolinensis, Helgicirrha schulzei, and 

Phialucium carolinae. H. schulzei apparently is not 

restricted to inshore waters (Russell, 1953), but 

the other two species may be. Mayer (1910) reported that B. carolinensis and P. carolinae were 

both very abundant in Charleston Harbor, South 

Carolina. In her study of hydromedusae from the 

North Carolina coast, Allwein (1967) found B. 

carolinensis to be restricted to inshore waters. 

Kramp (1953) noted that P. carolinae occurred in 

the lagoon but not outside the Great Barrier Reef. 

On several occasions the cubomedusae Carybdea 

marsupialis and Chiropsalmus quadrumanus were 

taken by attracting them to a light at night, and 

TABLE 19.—Abundance of dominant medusaen species at each Carrie Bow Cay station 

(presence in at least 33% of samples) 

Species 

Aglaura hemistoma 

Amphogona apsteini 

Bougainvillia carolinensis 

Cubaia aphrodite 

Helgicirrha schulzei 

Liriope tetraphylla 

Nausithoe punctata 

Phialidium spp. 

Phialucium carolinae 

Proboscidactyla ornata 

Sarsia angulata 

Solmundella bitentaculata 

Zanclea prolifera 

Occurrence in samples 

Reef 

55 

39 

0 

34 

0 

100 

39 

82 

3 

29 

39 

47 

42 

(%) 

Lagoon 

63 

23 

10 

32 

10 

100 

41 

86 

23 

41 

59 

41 

36 

Inshore 

0 

0 

100 

25 

100 

100 

0 

100 

100 

50 

0 

0 

0 

Reef 

37 (0-340) 

7 (0-47) 

0 

4 (0-15) 

0 

583 (24-5490) 

8 (0-105) 

190 (0-2132) 

<1 (0-1) 

5 (0-57) 

32 (0-350) 

18 (0-250) 

8 (0-60) 

Number of specimens/ WOO 

(Mean and range) 

Lagoon 

20 (0-164) 

4 (0-40) 

1 (0-13) 

3 (0-13) 

3 (0-48) 

390 (13-1290) 

8 (0-40) 

368 (0-3874) 

1 (0-1) 

12 (0-124) 

19 (0-124) 

7 (0-60) 

7 (0-40) 

m

3 

Inshore 

0 

0 

4500 (1000-8310) 

<1 (0-1) 

108 (31-250) 

662 (611-690) 

0 

750 (100-1530) 

337 (103-610) 

1 (0-6) 

0 

0 

0 
NUMBER 12 257 

by seining along the beach at Stann Creek. They 

also are inshore species that the author has observed in similar locations at Puerto Rico. 

Ubiquitous species are Cubaia aphrodite, Liriope 

tetraphylla, Phialidium spp., and Proboscidactyla ornata; most of these are cosmopolitan warm-water 

forms. Cubaia aphrodite, however, is restricted to 

the Caribbean and south Florida-Bahama region. Allwein (1967) found P. ornata in both offshore and inshore samples and also noted four 

species of Phialidium, of which P. languidum had a 

similar distribution. It is not known what species 

of Phialidium occur in Belize; the various western 

Atlantic forms are incompletely described and 

need revisionary work. 

Liriope tetraphylla is the only species that occurred in every sample and is relatively abundant 

at all three stations. Workers have made similar 

observations of this species in other areas of warm 

water, including the southeastern United States 

(Allwein, 1967), Australia (Kramp, 1953), and 

Brazil (Vannucci, 1963). The importance of L. 

tetraphylla as a carnivore is demonstrated by the 

fact that many specimens of this medusa had 

their manubria filled with fish ova, chaetognaths, 

crustaceans, and other zooplankton. In several 

samples, more than 40% of L. tetraphylla specimens 

contained fish ova. In the laboratory L. tetraphylla 

is highly active, swimming about with trailing 

tentacles. This species is so transparent that generally only its shadow is visible and it can easily 

go unnoticed by prey or predators. So little is 

known about the biology of this medusa that its 

role in tropical planktonic ecosystems can only be 

surmised. 

Medusae made up a significant amount of the 

biomass (wet weight) of samples from all stations 

(Table 20). At times they constituted more than 

50% of the reef and lagoon samples. High variation in biomass and numbers of individuals occurred from day to day, apparently owing to 

considerable water movement near the outer 

shelf. Inshore samples were dominated by medusae, which formed nearly 75% of the total plankton biomass collected. Owing to the large mesh 

of the net, 560 /xm, the numerous small copepods 

TABLE 20.—Biomass (wet weight) and numbers of specimens 

of medusae in relation to total net plankton at each station 

expressed in means (ranges in parentheses) 

Station 

Reef 

(36 samples) 

Lagoon 

(22 samples) 

Inshore 

(4 samples) 

Total 

plankton 

g/1000 m

3 

23 (6-43) 

14 (2-33) 

72 (45-95) 

Medusae 

Specimens/ 

g/W00m3 1000 m

3 

2(<1-17) 925(38-5771) 

2 (<1-15) 900(34-5047) 

52 (35-61) 6329 (3532-9169) 

that occur in tropical inshore areas were obviously 

not captured. 

Seasonal variation in biomass is likely, especially inshore where nutrient input and primary 

production would increase during the wet season. 

Since this study, however, represents only a time 

period within the dry season, the variability is not 

yet known. One species, Linuche unguiculata, did 

indicate seasonal reproduction. In Janaury 1978, 

multitudes of recently released L. unguiculata 

ephyrae were found on the reef flat at Carrie Bow 

Cay. In March small numbers of this medusa, 

about 5 mm in diameter, were collected at reef 

and lagoon stations. In May vast numbers of 

mature L. unguiculata, about 15 mm, suddenly 

appeared along the barrier reef at Carrie Bow 

and South Water cays (K. Riitzler, pers. comm. 

1978). 

Conclusions 

Seventy-one species of medusae belonging to 

nine orders were collected at three stations across 

the Belizean shelf. Fifty-seven species were found 

near the reef, 46 were collected at the lagoon 

station, and 21 inshore. The most abundant medusae show three distributional patterns. Species 

occurring only in the reef and outer lagoon are 

Aglaura hemistoma, Amphogona apsteini, Nausithoe 

punctata, Sarsia angulata, Solmundella bitentaculata, 

Zanclea prolifera, and Carybdea alata. Species occurring mainly inshore are Bougainvillia carolinensis, 

Helgicirrha schulzei, and Phialucium carolinae. Ubiq-
258 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

uitous species are Cubaia aphrodite, Liriope tetraphylla, Phialidium spp., and Proboscidactyla ornata. 

Medusae constituted from 10% to 70% or more 

of the total plankton biomass retained by a 560 

/mi net at the three stations. At reef and lagoon 

stations, medusae formed about 10% of the total 

sample with a mean value of about 2 g/1000 m3, 

whereas inshore they formed 70% of the total 

plankton or 52 g/1000 m3. At reef and lagoon 

stations, a mean abundance of about 900 specimens/1000 m3

 occurred versus inshore abundance of about 6000/1000 m3

. 

Liriope tetraphylla was the only species present 

in 100% of the samples; it also was the most 

abundant species at reef and lagoon stations. This 

medusa appears to be ecologically important because of its abundance and also because of its 

voracious feeding habits. 

Literature Cited 

Allwein, J. 

1967. North American Hydromedusae from Beaufort, 

North Carolina. Videnskabelige Meddelelser Fra Dansk 

Naturhistorisk Forening, 130:117-136. 

Bigelow, H. B. 

1938. Plankton of the Bermuda Oceanographic Expeditions, VIII: Medusae Taken During the Years 

1929 and 1930. Zoologica, 23 (2): 99-189. 

Kramp, P. L. 

1953. Hydromedusae. British Museum (Natural History) 

Great Barrier Reef Expedition 1928-29, Scientific Reports, 6(4): 259-322. 

Mayer, A. G. 

1900. Some Medusae from the Tortugas, Florida. Bulletin 

of the Museum of Comparative Zoology, Harvard, 37(2): 

13-82. 

1904. Medusae of the Bahamas. The Museum of the Brooklyn Institute of Arts and Sciences, Memoirs of Natural 

Sciences, 1(1): 1 -33. 

1910. Hydromedusae. In Medusae of the World. Carnegie 

Institution of Washington, Publication, 109 (1,2): 498 

pages. 

Moore, E., and F. Sander. 

1976. Quantitative and Qualitative Aspects of the Zooplankton and Breeding Patterns of Copepods at 

two Caribbean Coral Reef Stations. Estuarine and 

Coastal Marine Science, 4:586-607. 

Renon, J. P. 

1977. Zooplancton du lagon de l'atoll de Takapota (Polynesie francaise). Annates de ITnstitul Oceanographique, Paris, 53(2):217-236. 

Russell, F. S. 

1953. The Medusae of the British Isles. 530 pages. Cambridge University Press. 

Vannucci, M. 

1963. On the Ecology of Brazilian Medusae at 25° Lat. 

S. Boletim do Instituto Oceanografico, 13(1): 143-184. 

Wood, E.J.F., and R. E. Johannes. 

1975. Tropical Marine Pollution. 192 pages. Elsevier Scientific Publishing Company. [Elsevier Oceanography Series, 12.] 
Plates 
2 6 0 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

PLATE 1.—Carrie Bow Cay in relation to the barrier reef of Belize (see also Figure 222): top left, 

discontinuous barrier reef off Ambergris Cay looking SSW; top right, barrier reef with cut east 

of Colson point, Columbus Reef and Southern Long Cay in background; center left, continuous 

shallow reef development north of South Water Cay; center right, Carrie Bow and South Water 

cays on the barrier reef, Twin Cays in the lagoon; bottom left, Carrie Bow Cay looking north, 

June 1972; bottom right, Carrie Bow Cay with partly exposed reef flat; yellow laboratory building 

left of center. 

2 6 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

PLATE 2.—Lagoon and back-reef habitats, 0-2 m depth: lop left, seagrass community, Thalassia 

testudinum and Dictyota sp.; top right, reef flat seaward of Carrie Bow Cay, Thalassia testudinum and 

Porites porites on rubble bottom; center left, Acropora cervicornis on rubble substrate in patch reef 

zone, April 1980; center right, rubble storm ridge east of Carrie Bow Cay prograding over Porites 

astreoides established on back-reef pavement, March 1975; bottom left, inner edge of reef crest 

with Porites astreoides, rubble and pavement zone in background; bottom right, community at 

inner edge of reef crest, with Halimeda opuntia, Porites astreoides, and Agaricia agaricites (Scale = 40 

cm). 

2 6 4 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

PLATE 3.—Reef crest and high-relief spur and groove zone; top left, Millepora complanata 

community, central reef crest; top right, Acropora palmata thicket, outer edge of reef crest; center 

left, coral pinnacles in transition zone between reef crest and high-relief spur and groove zone, 

with Acropora palmata, A. prolifera, Agaricia tenuifolia, and Millepora complanata, 2 m; center right, 

high-relief spur and groove development, with massive coral heads and octocorals dominating 

the sand groove, 6 m; bottom left, coral buttress formed by association oi Agaricia tenuifolia, Porites 

porites, and Millepora complanata, 3 m; bottom right, overhanging flank of coral buttress with 

prominent Gorgonia ventalina and Pseudopterogorgia sp., 8 m. 

2 6 6 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

PLATE 4.—Low-relief spur and groove zone and outer fore reef: lop left, low coral spur covered 

by coral heads, rubble, and octocorals, sand groove in background, 10 m; top right, octocoral 

community at seaward edge of low-relief spur and groove zone including Pseudopterogorgia spp., 

Gorgonia •ventalina, and Eunicea sp., 14 m; center left, top of inner reef slope, columnar growth of 

Montastrea annularis in Acropora cervicornis thicket, 17 m; center right, Acropora cervicornis community 

on top of outer ridge with corals Agaricia tenuifolia, Montastrea annularis, Madracis sp., and sponge 

Agelas sp., 14 m; bottom left, fore-reef slope community, with platy Montastrea annularis, a variety 

of sponges, including Aplysinafistularis, Mycale sp., Plakortis sp., and octocoral Iciligorgia schrammi, 

30 m; bottom right, fore-reef slope community consisting of the corals Montastrea annularis, Madracis 

decactis, Porites porites, the crinoid Nemaster rubiginosa, the octocorals Plexaura flexuosa, Eunicea sp., 

Gorgonia ventalina, and sponge Ectyaplasia ferox. 

2 6 8 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

PLATE 5.—Storm ridge and lagoon habitats: top left, intertidal rubble field on the reef crest 

looking northwest; top right, representative ophiuroid assemblage under coralline coated coral 

slab from rubble field consisting of (clockwise from center top) Ophioderma anitae, 0. rubicundum, 

0. appressum, O. guttatum, 0. guttatum lower side, and 0. phoenium; center left, transition zone 

between lagoon seagrass flat and patch reef, with seagrass Thalassia testudinum, corals Siderastrea 

siderea and Montastrea annularis, hydrocoral Millepora alcicornis, and sponges Ircinia strobilina and 

Desmapsamma anchorata; center right, patch reef community on dead coral base, with sponges 

Ircinia strobilina, Pseudoceratina crassa, Amphimedon complanata, and Niphates digitalis, hydrocoral 

Millepora alcicornis, and octocoral Gorgonia ventalina; bottom left, Twin Cays mangrove channel, 

with Thallasia testudinum covered mud bottom and Rhizophora mangle roots colonized by fire 

sponge Tedania ignis and sulphur sponge Lissodendoryx isodictyalis; bottom right, overhanging mud 

bank in Twin Cay mangrove channel covered predominately by orange sponge Ulosa ruetzleri 

and by unidentified colonial ascidian. 
j 

Mt 

Hni 

• 

y

'

r

' ffiS 4BJ L 

PJ'i u 

;W - iH^ J 

w^&^Jwt 

rkig^H0%W-. ••**' **TT| 

HI ''^inl mkrTu 

^•7 ^^HBWBs ^ 

"•"^" Jr^ 

BF^J M 

^fcLtf.' '^4^ll 

•J 

" 

l^v • «*,»& 

..^•fr ft BAJll 

^BLsk- fra! 

y * ' JH' 

Br-A- J* 

i jraEKgi 
Stony Corals (Cnidaria: Hydrozoa, Scleractinia) 

of Carrie Bow Cay, Belize 

Stephen D. Cairns 

ABSTRACT 

Three species of hydrozoan and 42 species of 

scleractinian corals were identified among 248 

specimens collected from maximum depths of 31 

m during a survey along the outer edge of the 

Belizean barrier platform near Carrie Bow Cay 

and Curlew Bank. This report includes important 

synonyms, morphological observations, and distributional data, as well as photographs of all 

species. Diversity of the Carrie Bow Cay coral 

fauna is high, as compared with adjacent areas of 

the Caribbean, although not all habitats have yet 

been equally well searched. 

Introduction 

One of the earliest reports on the Scleractinia 

off Belize (Boone, 1928) recorded two species of 

deep-water ahermatypes from off Glover's Reef 

at a depth of 885 m. More recently, Thorpe and 

Bregazzi (1960) listed 30 hermatypic species from 

Rendezvous Cay, two of which are considered to 

be only forms in this account. Stoddart (1962:19) 

listed 22 reef corals from three atolls off Belize, 

and York (1971) discussed and figured 24 hermatypes found off the southern shelf of Belize. 

These numbers have been revised to be consistent 

with the systematic concept of this paper. Specific 

references to the coral fauna and reef zonation of 

Carrie Bow Cay are found in Stoddart (1963:25-

26) and in Riitzler and Macintyre (herein: 9). 

Stephen D. Cairns, Department of Invertebrate Zoology, National 

Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. 

ACKNOWLEDGMENTS.—I thank R. J. Larson and 

B. Spracklin for helping to collect corals at Carrie 

Bow Cay. 

Materials and Methods 

Most of the corals were collected in March 

1978 during a survey of reefs at Carrie Bow Cay 

and nearby Curlew Bank to a depth of 31 m. All 

248 specimens are deposited in the United States 

National Museum collection (National Museum 

of Natural History, Smithsonian Institution). Illustrated specimens were collected at Carrie Bow 

Cay unless otherwise indicated. In some cases, 

specimens were dyed and then coated with a fine 

layer of NH4CI in order to improve contrast for 

photography; these specimens are noted in the 

figure legends. 

Because of inconsistencies in the literature, authors and references are given for all species. 

Synonymies are restricted to publications in 

which the species are correctly identified and 

usefully illustrated. Data on bathymetric ranges 

apply to the entire geographic range of the species 

and are taken largely from Goreau and Wells 

(1967). 

Species List 

(* = ahermatypic) 

Class HYDROZOA 

Order MILLEPORINA 

Family MILLEPORIDAE 

Millepora alcicornis Linnaeus 

M. complanata Lamarck 

271 
272 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Order STYLASTERINA 

Family STYLASTERIDAE 

*Stylaster roseus (Pallas) 

Class ANTHOZOA 

Order SCLERACTINIA 

SUBORDER ASTROCOENIINA 

FAMILY ASTROCOENIIDAE 

Stephanocoenia michelinii Milne Edwards and Haime 

Family POCILLOPORIDAE 

Madracis decactis (Lyman) 

M. mirabilis (Duchassaing and Michelotti) 

*M. pharensis forma pharensis (Heller) 

Family ACROPORIDAE 

Acropora palmata (Lamarck) 

A. cervicornis (Lamarck) 

A. prolifera (Lamarck) 

Suborder FUNGUNA 

Family AGARICIIDAE 

Agaricia agaricites forma agaricites (Linnaeus) 

forma carinata Wells 

forma purpurea (Lesueur) 

A. fragilis forma fragilis Dana 

A. tenuifolia Dana 

A. lamarcki Milne Edwards and Haime 

Leptoseris cucullata (Ellis and Solander) 

Family SIDERASTREIDAE 

Siderastrea siderea (Ellis and Solander) 

S. radians (Pallas) 

Family PORITIDAE 

Porites astreoides Lamarck 

P. porites forma porites (Pallas) [= P. clavaria Lamarck] 

forma divaricata Lesueur 

forma furcata Lamarck 

Suborder FAVIINA 

Family FAVIIDAE 

Favia fiagum (Esper) 

Diploria clivosa (Ellis and Solander) 

D. strigosa (Dana) 

D. labyrinthiformis (Linnaeus) 

Manicina areolata forma areolata (Linnaeus) 

forma mayori Wells 

Colpophyllia natans (Houttuyn) 

Cladocora arbuscula (Lesueur) 

Montastrea annularis (Ellis and Solander) 

M. cavernosa (Linnaeus) 

Family RHIZANGIIDAE 

*Phyllangia americana Milne Edwards and Haime 

*Astrangia solilana (Lesueur) 

*Colangia immersa Pout ales 

Family MEANDRINIDAE 

Meandrina meandriles iormxmeandrites (Linnaeus) 

forma danae (Milne Edwards and Haime) 

Dichocoenia stokesi Milne Edwards and Haime 

Dendrogyra cylindrus Ehrenberg 

Family MUSSIDAE 

Mussa angulosa (Pallas) 

Scolymia lacera (Pallas) 

S. cubensis (Milne Edwards and Haime) 

Isophyllia sinuosa (Ellis and Solander) 

Isophyllastrea rigida (Dana) 

Mycetophyllia lamarckiana Milne Edwards and 

Haime 

M. aliciae Wells 

M. danaana Milne Edwards and Haime 

M. ferox Wells 

Suborder CARYOPHYLLIINA 

Family CARYOPHYLLIIDAE 

EusmiHa fastigiata forma fastigiata (Pallas) 

Family FLABELLIDAE 

*Gardineria minor Wells 

Family MILLEPORIDAE 

Millepora alcicornis Linnaeus, 1758 

FIGURE 119a; PLATE 5: center left, center right 

Millepora alcicornis.—Vaughan, 1901:318, pis. 35, 37-38.— 

Boschma, 1948:18-19, 23-28, 79-81, 86-88, 100, fig. 6, pl. 

14: fig. 3.—Squires, 1958:259, pl. 43: figs. 1-2.—Almy 

and Carrion-Torres, 1963:144, pl. 2a.—Olivares and 

Leonard, 1971:59, pl. 5: figs, C-D.—Roos, 1971:43-44, pl. 

2.—Smith, 1971:95-96.—York, 1971:5, pl. 1: figs. 1-4.— 

Tresslar, 1974:123-124, pl. 17.—Colin, 1978:137 [color 

figs.], 146 [figs.], 150. 

Encrusting or branching colonies up to 50 cm 

in width. Branches in bushy arrangement or 

uniplanar, often anastomosing. Surface covered 

by pores; each gastropore (0.2 mm in diameter) 

surrounded by 5-7 smaller dactylopores (0.1 mm 

in diameter) in an indistinct cyclosystem. Gastrostyles and dactylostyles absent. Orange-yellow 

or brown. 

HABITAT.—Commonly found encrusting dead 

gorgonians in fore reef (4-30 m) and in lagoon 

patch reefs (3-6 m). 

Millepora complanata Lamarck, 1816 

FIGURE l\9b-c; PLATE 3: top left 

Millepora complanata.—Boschma, 1948:20, 34-35, 83-84, 94-

95, 103, figs. 2a-b, 11, pl. 7: fig. 2.—Squires, 1958:259, pl. 

43: fig. 3.—Almy and Carrion-Torres, 1963:144, pl. 2b.— 

Roos, 1971:44-45, pl. 3.—York, 1971:5, pl. 1: figs. 5-8.— 

Colin, 1978:140 [color fig.], 143 [figs.], 150. 
NUMBER 12 273 

FIGURE 119.—a, Millepora alcicornis, Bermuda; b, c, M. complanata, reef crest, 0.5 m, specimen in 

c coated with NH4 CI; d, e, Stylaster roseus, spur and groove zone. (Scale bars: a, b = 2 cm; c = 0.2 

cm; d = 1 cm; e = 0.5 cm.) 
274 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Colony forming thin, smooth, vertical plates 

united only at their bases. Otherwise, similar to 

Millepora alcicornis. 

HABITAT.—Major component of reef crest and 

fore reef spurs (0-8 m). 

DISCUSSION.—A third western Atlantic species, 

Millepora squarrosa Lamarck, 1816, is characterized 

by thin, vertical plates with lateral expansions or 

ridges, which unite with adjacent plates to form 

a honeycombed complex. 

Family STYLASTERIDAE 

Stylaster roseus (Pallas, 1766) 

FIGURE 1 \9d-e 

Stylaster roseus.—Boschma, 1965:227-247, [4 figs.], pis. 1-3.— 

Roos, 1971:45, pl. 4: figs, a-b.—Colin, 1978:141 [color 

fig.], 144 [color fig.], 151. 

Small, branching, uniplanar colonies rarely 

over 10 cm tall. Each gastropore surrounded by 

5-15 smaller, slit-like dactylopores opening on to 

the gastropore. Gastrostyles and dactylostyles present. Small, hemispherical ampullae commonly 

present, sometimes in crowded arrangement. Purple, red, or pink. 

HABITAT.—Very common in spur and groove 

zone (6-8 m) on undersides of, or within cavities 

produced by Agaricia tenuifolia. Also present on 

fore-reef slope (20-24 m). Ahermatypic. Bathymetric range: 6-30 m. 

Family ASTROCOENIIDAE 

Stephanocoenia michelinii Milne Edwards and 

Haime, 1848 

FIGURE \20a-b 

Plesiastrea goodei Verrill, 1900:553-554, pl. 67: fig. 1. 

Stephanocoenia michelinii.—Squires, 1958:246, pl. 32: figs. 1-

2.—Smith, 1971:72, pl. 2.—York, 1971:7, pl. 2: figs. 1-

4.—Laborel, 1971:175, pl. 1: fig. 4.—Colin, 1978:209 

[color fig.], 211. 

Stephanocoenia intersepta.—Roos, 1971:51, pl. 4: fig. c, pl. 5: 

figs, a-b.—Tresslar, 1974:118. pl. 1. 

Plocoid to cerioid, subhemispherical or encrusting colonies up to 30 cm in diameter. Calices 2-

3 mm in diameter. Twenty-four septa; prominent 

paliform lobes before first two cycles of septa. 

Columella large, styliform. Brown. 

HABITAT.—Only one specimen was collected 

from the ridge of the outer fore reef (14 m). 

Bathymetric range: 1-95 m. 

DISCUSSION.—This species is often mistaken in 

the field for the more common Siderastrea siderea 

or S. radians, which it superficially resembles. 

Family PociLLOPORlDAE 

Madracis decactis (Lyman, 1859) 

FIGURE I20c-d; PLATE 4: bottom right 

Madracis decactis.—Verrill, 1901a: 108-109, figs. 2, 2a, pl. 14: 

fig. 6.—Roos, 1971:52-53, pl. 8: figs, a-b.—Wells, 1973: 

19 [key].—Tresslar, 1974:120, pl. 2.—Erhardt and Meinel, 

1975, fig. 3.—Colin, 1978:211, 212 [color fig.], 214. 

Madracis decatis.—Smith, 1971:72-73, pl. 1. 

Plocoid; nodular, clavate, or thin, encrusting 

colonies rarely exceeding 15 cm in diameter. Calices about 2 mm in diameter containing 10 septa 

each. Columella styliform. Noncostate; coenosteum bears rows of short spines surrounding each 

calice. Brown, green, or purplish. 

HABITAT.—At Carrie Bow Cay found only on 

outer fore reef (15-24 m). Bathymetric range: 1-

70 m. 

Madracis mirabilis (Duchassaing and 

Michelotti, 1860) 

FIGURE 120<?-/ 

Madracis asperula.—Roos, 1971:59, pl. 9: fig. b, pl. 10. 

Madracis mirabilis.—Wells, 1973:19 [key].—Werding and Erhardt, 1976; pl. 4: fig. 1.—Colin, 1978:212 [color fig.], 

214,215 [fig.]. 

Plocoid, ramose, bushy colonies up to 2 m in 

diameter. Branches 5-9 mm in diameter with 

blunt ends. Calices 1-2 mm in diameter; 10 septa 

each. Columella styliform. Pale cream to bright 

yellow. 

HABITAT.—Found in sand trough of outer fore 

reef (17-22 m) and spur and groove zone (6-8 

m). Bathymetric range: 1-60 m. 

DISCUSSION.—This species has been confused 
NUMBER 12 275 

^ ^ 

FIGURE 120.—a, b, Stephanocoenia michelinii, outer ridge, 14 m; c, d, Madracis decactis, outer ridge, 

14 m; e, M. mirabilis, inner-reef slope, 15-18 m; fi M. mirabilis, Wee Wee Cay, 1 m, specimen 

coated with NH4C1. (Scale bars: a, c, e = 2 cm; b, d,f= 0.2 cm.) 
276 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

with Madracis asperula Milne Edwards and Haime. 

The latter is distinguished by its more slender 

branches (2-3 mm) and attenuate branch tips; it 

is ahermatypic and is usually found in deeper 

water (to 200 m). 

Madracis pharensis forma pharensis 

(Heller, 1868) 

FIGURE 121a 

Madracis pharensis forma pharensis.—Wells, 1973:18 [key]. 

Madracis pharensis.—Colin, 1978:213 [color fig.], 214. 

Thin, encrusting, stolon-like ribbons or small 

nodular projections. Calices about 1.5 mm in 

diameter, each with 20 septa: 10 large, 10 small. 

Columella styliform. Yellow with pinkish polyps. 

HABITAT.—All Carrie Bow Cay specimens were 

found on the fore-reef slope (24-28 m) attached 

to the undersides of platy Agaricia spp. 

Ahermatypic. Bathymetric range: 20-150 m. 

DISCUSSION.—Madracis pharensis forma luciphila 

Wells, 1973, forms thick, encrusting, laminar 

sheets, similar to M. decactis. It is hermatypic and 

has a shallower bathymetric range (2-95 m). 

Family ACROPORIDAE 

Acropora palmata (Lamarck, 1816) 

FIGURE \2\b,c; PLATE 3: top right, center left 

Isopora muricata forma palmata.—Vaughan, 1901:313-314, pis. 

26-27. 

Acropora palmata.—Squires, 1958:246-247, pl. 34: fig. 1.— 

Roos, 1971: 55-56, pl. 12: fig. b.—Smith, 1971: 73-74, pl. 

4.—York, 1971: 7, pl. 2: figs. 5-8.—Colin, 1978: 36, 216 

[color figs.], 218 [fig.], 219, 222. 

Broad, flat, frond-like branches, forming colony meters in diameter. Calices 1.0-1.5 mm in 

diameter, usually on highly exsert, tubular corallites. No axial corallites. Twelve septa. Wall 

porous. Yellowish brown. 

HABITAT.—Very common in turbulent shallow 

waters. At Carrie Bow Cay common directly 

seaward and shoreward of the reef crest, and at 

high spur and groove zone (1-8 m); also common 

at Curlew Bank (2-4 m). Sparse in back-reef area 

(1-2 m). Bathymetric range: 0-17 m. 

Acropora cervicornis (Lamarck, 1816) 

FIGURE \2\d; PLATE 2: center left, PLATE 4: center left, center 

right 

Isopora muricata.—Vaughan, 1901:312-313, pl. 21, pl. 22: 

fig. 2. 

Acropora cervicornis.—Roos, 1971:54-55, pl. 9: fig. a, pl. 12: 

fig. a.—Smith, 1971:73, pl. 3.—York, 1971:9, pl. 3: figs. 

1-2.—Colin, 1978:216, 217 [color figs.], 222-223. 

Cylindrical branches, forming colonies up to 3 

m in height. Calices about 1 mm in diameter; 

axial corallites slightly larger. Tubular corallites 

usually highly exsert. Twelve septa. Wall porous. 

Yellowish brown. 

HABITAT.—Common as dense thickets in sand 

trough and outer ridge of the outer fore reef (24 

m); also present in spur and groove zone, backreef area (1-2 m), and adjacent patch reefs. Bathymetric range: 0-50 m. 

Acropora prolifera (Lamarck, 1816) 

FIGURES 121e, 122a; PLATE 3: center left 

Isopora muricata forma prolifera.—Vaughan, 1901:313, pl. 22: 

fig. 1, pis. 23-25. 

Acropora prolifera.—Roos, 1971:55, pl. 11.—Smith, 1971:74.— 

York, 1971:9, pl. 3: figs. 3-4.—Colin, 1978:217 [color 

fig.], 223, 226. 

Very similar to Acropora cervicornis, but branches 

more crowded, often crossing over and fusing, 

producing flabelliform or reticulate plates. Usually smaller than A. cervicornis and not found in 

dense thickets. 

HABITAT.—Found on seaward side of reef crest 

(0.5-2.0 m) in very turbulent waters. Bathymetric 

range: 0-30 m. 

DISCUSSION.—All other characters being similar, the three species oi Acropora are differentiated 

by growth form: A. prolifera is the intermediate, 

linking the frond-like A. palmata and the ramose 

A. cervicornis. Some authors (Gregory, 1895; 

Vaughan, 1901) have considered them to be three 
NUMBER 12 277 

*.. -*> •fe ,- *, . ".'T r — 

FIGURE 121.—a, Madracis pharensis i. pharensis, fore-reef slope, 24-27 m, on underside oi Agaricia 

fragilis, specimen coated with NH4CI; b, c, Acropora palmata, off Cuba; d, A. cervicornis, exact 

locality at Carrie Bow Cay unknown; e, A. prolifera, off Belize, exact locality unknown. (Scale 

bars: a = 0.2 cm; b, d = 5 cm; c, e = 0.5 cm.) 
278 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 122.—a, Acropora prolifera, off Belize, exact locality unknown; b, Agaricia agaricites f. 

carinata, fore-reef slope, 24-27 m; c, d, A. agaricites purpurea, sand trough, 21 m; e, A. agaricites 

agaricites, patch reef area, 2-3 m; /, A. tenuifolia, exact locality at Carrie Bow Cay unknown. 

(Scale bars: a, b,f= 5 cm; c = 0.5 cm; d, e — 2 cm.) 
NUMBER 12 279 

forms of the same species, whereas others (Smith, 

1971; Roos, 1971; Wells, 1973) have retained 

them as separate species. Discussions of the problem can be found in Vaughan (1901:312-314; 

1919:482) and in Roos (1971:54). 

Family AGARICIIDAE 

Agaricia agaricites (Linnaeus, 1758) 

FIGURE I22b-e 

Agaricia agaricites.—Verrill, 1901a: 146-149 [in part, not variety e], figs. 5-7, pl. 26: figs. 2-3, pl. 27: figs 1-3, 5-7.— 

Squires, 1958:247-248, pl. 32: fig. 3, pl. 33: figs. 1-3.— 

Roos, 1971:56-58, pl. 14: figs, a-b.—Smith, 1971:74-75, 

pis. 5-6.—York, 1971:11, 13 [in part, pl. 4: figs. 3-8, 

pl. 5: figs. 1-6].—Wells, 1973:25, fig. 6a [not A. a. 

fragilis].—Tresslar, 1974:120, pl. 4.—Colin, 1978:220, 221 

[color fig.], 226, 231. 

Agaricia crassa Verrill, 1901a: 145, pl. 30: fig. 6, pl. 34: fig. 2. 

Agaricia purpurea.—Verrill, 1901a: 149-150, pl. 27: figs. 

4a-b. 

Colonies massive and encrusting; thick and 

flat, attached by pedicel; or flat with thick imbricated, vertical, bifacial lobes. Up to 1 m in 

diameter. Cerioid to meandroid. Calices 1.5-3.0 

mm in diameter, arranged in reticulate pattern 

or in rows of up to 20; 3-7 calices/cm. Columella 

small. Chocolate to purplish brown. 

HABITAT.—Agaricia agaricites forma agaricites is 

most common in back-reef and patch reef areas 

(0.5-2.0 m); A. a. forma purpurea (Lesueur, 1821) 

is common in the spur and groove zone and outer 

fore reef (7-30 m); and A. a. forma carinata Wells, 

1973, was collected only from the fore-reef slope 

(18-30 m). Bathymetric range: 1-70 m. 

DISCUSSION.—At least five growth forms have 

been described and named for this variable species and several more forms could be described 

on the basis of similar criteria. The most common 

form at Carrie Bow Cay is purpurea, a large (up to 

1 m), thick, flat colony with 5-6 calices/cm and 

long valleys having up to 20 calices. Each valley 

is bordered by prominent collines. Forma agaricites 

is smaller, hemispherical or encrusting, with 

larger calices arranged in a discontinuous or reticulate pattern and only 3-5 calices/cm. Forma 

carinata is like purpurea but has scattered, low (2-3 

cm), thick, bifacial crests. The remaining two 

forms have not been collected at Carrie Bow Cay: 

forma humilis Verrill has small (1.5 mm in diameter), pinched calices in a crowded, reticulate 

arrangement; and forma danai Milne Edwards 

and Haime is massive with tall, erect, imbricated 

lobes. 

Agaricia fragilis forma fragilis Dana, 1848 

FIGURE 123a-o 

Agaricia fragilis.—Verrill, 1901a: 142-145, pl. 26: fig. la-d.— 

Smith, 1971:75, pl. 8.—Wells, 1973:24 [not fig. 6a].— 

Colin, 1978:224 [color fig.], 234. 

Pedicelled or encrusting, very thin, unifacial 

fronds, either flat or vase-shaped. Usually less 

than 15 cm in diameter. Calices about 1.0 mm in 

diameter, 5-7/cm. Very low or no collines. Chocolate or purplish brown. 

HABITAT.—Common on fore-reef slope (18-27 

m). Bathymetric range: 3-40 m. 

DISCUSSION.—Agaricia fragilis is similar to A. 

agaricites forma purpurea, but is distinguished by 

its thinner, more delicate corallum and its lack of 

or very small collines. Roos (1971) considered A. 

fragilis a variety of A. agaricites. 

Agaricia tenuifolia Dana, 1848 

FIGURE 122/; PLATE 3: center left, bottom left 

Agaricia agaricites.—York, 1971:9, pl. 3: figs. 5-6. 

Agaricia tenuifolia.—Wells, 1973:25 [key], fig. 13.—Erhardt 

and Meinel, 1975, fig. 5.—Werding and Erhardt, 1976, 

pl. 1: fig. 1.—Colin, 1978:224 [color fig.], 227 [fig.], 231. 

Large, bushy colonies up to several meters in 

size, composed of thin, vertical, dissected fronds. 

Corallum bifacial. Valleys short, sometimes reticulate. Collines prominent. Columella small. 

Brownish. 

HABITAT.—At Carrie Bow Cay this species is 

ubiquitous: it is common in the back-reef area 

(2-4 m) and is the major component of the 

buttresses and spurs in the spur and groove zone 

of the inner and fore reef (5-8 m). It is also found 

in the sand trough and fore-reef slope (20-27 m). 

Bathymetric range: 2-27 m. 
280 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 123.—a, b, Agaricia fragilis f. fragilis, fore-reef slope, 24-27 m; c, d, Leptoseris cucullata, 

inner reef slope, 15-18 m; e,fi A. larmarcki, fore-reef slope, 24-27 m. (Scale bars: a, c — 5 cm; b, 

f= 0.5 cm; d = 1 cm; e = 2 cm.) 
NUMBER 12 281 

Agaricia lamarcki Milne Edwards and Haime, 

1851 

FIGURE I23e-f 

Agaricia lamarcki.—Wells, 1973:26-28, figs. 8-10.—Colin, 

1978:234. 

Thin, unifacial, centrally attached fronds up to 

2 m in diameter. Calices about 2 mm in diameter, 

3-5/cm along valley. Valleys long, 4-6 mm wide. 

Septa thinner than interspaces, alternating in 

height and thickness. Collines prominent and 

thick. Columella large. 

HABITAT.—Collected only on fore-reef slope 

(25-27 m). Bathymetric range: 4-46 m. 

DISCUSSION.—Distinguished from Agaricia grahamae Wells, 1973, by its thinner septa, which 

alternate in size, and its wider valleys. 

Leptoseris cucullata (Ellis and Solander, 1786) 

FIGURE I23c-d 

Agaricia nobilis Verrill, 1901a:150-151, pl. 28: figs. 1-2.— 

Smith, 1971:75-76, pl. 7. 

Agaricia agaricites.—Roos, 1971:56-58 pl. 15. 

Helioseriscucullata.—Wells, 1973:25 [key], figs. 14,33.—Colin, 

1978:229, 232 [color figs.], 234-235. 

Helioceris cucullata.—Tresslar, 1974:121, pl. 6. 

Leptoseris cucullata.—Dinesen, 1980:187-188, pl. 1: figs. 1-3. 

Very thin, unifacial, pedicelled fronds up to 40 

cm in diameter. Underside costate. Calices about 

3 mm in diameter, each of which is bordered by 

a prominent, crescent-shaped hood, which is 

strongly inclined toward the edge of the frond. 

Adjacent "hoods" eventually unite, forming a 

long colline. No columella. Green polyps on 

brown background. 

HABITAT.—Very common on fore-reef slope 

(25-28 m), particularly in slightly recessed cavities. Also present in sand trough (22 m), innerreef slope (14-16 m), and high spur and groove 

zone (6-9 m). Bathymetric range: 3-90 m. 

Family SIDERASTREIDAE 

Siderastrea siderea (Ellis and Solander, 1786) 

FIGURE 124<r; PLATE 5: center left 

Siderastrea siderea.—Vaughan, 1919:443-447, pl. 114: figs. 2-

3, pl. 122: figs. 1-3.—Roos, 1971:62-63, pl. 13.—Smith, 

1971:76-77, pl. 10.—York, 1971:15, pl. 6: figs. 1-2.— 

Tresslar, 1974:121, pl. 7.—Colin, 1978:232 [color fig.], 

235. 

Cerioid; encrusting or hemispherical boulders 

up to 2 m in diameter. Calices 4-5 mm in diameter. Usually more than 48 septa. Inner edges of 

septa slope gently (about 45°) toward columella, 

resulting in a wide fossa. Theca fenestrate. Light 

reddish brown. 

HABITAT.—Most common in the shallow backreef area (0.5-2.0 m). Also found on inner-reef 

slope and patch reefs (Curlew Bank, 6 m). Bathymetric range: 0.5-70.0 m. 

Siderastrea radians (Pallas, 1766) 

FIGURE I24a-b 

Siderastrea radians.—Vaughan, 1919:439, pl. 114: fig. 1.— 

Roos, 1971:62-63, pis. 20-21.—Smith, 1971:76, pl. 9.— 

York, 1971:15, pl. 6: figs. 3-6.—Werding and Erhardt, 

1976, pl. 3: fig. 2.—Colin, 1978:233 [color fig.], 235, 238. 

Cerioid; encrusting, hemispherical, or unattached spheroidal colonies rarely exceeding 30 cm 

in diameter. Unattached coralla with calices covering entire surface. Calices about 3 mm in diameter, 24-40 septa/calice. Inner edges of septa 

almost vertical, producing a narrow fossa. Theca 

fenestrate. Greenish or greyish brown. 

HABITAT.—Most common in the back-reef area 

(0-20 m), but also collected from the outer fore 

reef (28 m). Bathymetric range: 0-33 m. 

DISCUSSION.—Distinguished from S. siderea by 

its smaller size, fewer septa, and vertical, inner 

septal edges. S. radians forma stellata Verrill, a 

form with a tendency toward deeper calices in a 

meandriform arrangement, was not collected at 

Carrie Bow Cay. 

Family PORITIDAE 

Porites astreoides Lamarck, 1816 

FIGURE 124a7; PLATE 2: center right, bottom left, 

bottom right 

Porites astreoides.—Vaughan, 1901:317-318, pis. 32-34.— 

Verrill, 1901a: 160-161, pl. 31: fig. 4.—Squires, 1958:250-
282 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

*#. ifc # 

*A^- - ^r" *^lfe 1$ 4k 

FIGURE 124.—a, b, Siderastrea radians, exact locality at Carrie Bow Cay unknown; c, S. siderea, 

patch reef area, 1 m, specimen coated with NH4CI; d, Pontes astreoides, fore-reef slope, 18 m; e, 

P. porites i. porites, exact locality at Carrie Bow Cay unknown; /, P. porites i. porites, patch reef 

area, 2-3 m. (Scale bars: a = 1 cm; b, c = 0.2 cm; d,f= 2 cm; e = 0.5 cm). 
NUMBER 12 283 

251, pl. 39: figs. 2-3.—Laborel, 1971:187-188, pl. 3: figs. 

4-6.—Roos, 1971:61-62, pl. 18: fig. a, pl. 19: fig. a.— 

Smith, 1971:77, pl. 15.—York, 1971:21, pl. 9: figs. 1-6.— 

Tresslar, 1974:121, pl. 8.—Erhardt and Meinel, 1975, 

fig. 1. 

Porites asteroides.—Colin, 1978:237, 240 [color figs], 242, 243 

[fig-]-

Cerioid; encrusting, flat, or hemispherical colonies rarely exceeding 60 cm in diameter. Surface 

of colony usually bumpy in shallow-water specimens, flat in deeper-water specimens. Calices 

1.25-1.50 mm in diameter, each with 12 septa. 

Columella small. Bright green or yellowish 

brown. 

HABITAT.—Found in all reef environments: 

back reef (1-2 m), spur and groove zone (6-8 m), 

sand trough (22 m), and fore-reef slope (18-28+ 

m). Also known from muddy back reefs and 

mangrove areas (Roos, 1971). Bathymetric range: 

0.2-70.0 m. 

Porites porites (Pallas, 1766) 

FIGURES 124*-/, \25a-c; PLATE 2: top right 

Porites porites.—Vaughan, 1901:314-316, pl. 2: fig. 4, pis. 28-

31.—Squires, 1958:251-253, pl. 38: figs. 1-3, pl. 39: fig. 

1.—Roos, 1971:58-60, pis. 16-17.—Smith, 1971:78-79, 

pis. 13-14.—York, 1971:17, 19, pl. 7: figs. 1-6, pl. 8: figs. 

1-5.—Erhardt and Meinel, 1975, fig. 2.—Colin, 1978:236, 

237, 239, 243 [color figs.]. 

Porites polymorpha.—Verrill, 1901a: 158-159, pl. 31: fig. 3. 

Porites.—Bernard, 1906:5-12, pis. 9-16. 

Porites divaricata.—Smith, 1971:78.—Colin, 1978:236 [color 

fig.], 239. 

Porites furcata.—Smith, 1971:78, pl. 12.—Tresslar, 1974:121, 

pl. 9.—Colin, 1978:233 [color fig.], 238. 

Cerioid, ramose colonies up to 50 cm tall and 

several meters wide. Branch diameters range from 

a delicate 5 mm to a massive 25+ mm. Branch 

tips clubbed or cylindrical, bifurcated or unbranched. Theca porous. Calices 1.5-2.0 mm in 

diameter, 12 septa/calice. Five-six small pali. Yellow, grey, or buff brown. 

HABITAT.—Porites porites iormafurcata Lamarck, 

1816, is the most common form at Carrie Bow 

Cay, being found in all reef environments, including the lagoon and back reef (1-3 m), the spur 

and groove zone (7 m), the sand trough (21 m), 

and the fore-reef slope (27 m). Forma porites was 

observed only in shallow (1-2 m) back-reef areas. 

Forma divaricata Lesueur, 1821, was collected unattached at nearby Twin Cays in a shallow backreef environment, but not at Carrie Bow Cay. 

Bathymetric range: 0.5-50 m. 

DISCUSSION.—There has been continued disagreement on whether the three phenotypes of 

Porites porites are distinct species or simply forms 

of the same species. Discussions of the problem 

are found in Vaughan (1901), Bernard (1906), 

and Roos (1971). This study treats them as 

formae, owing to the almost continuous range of 

growth forms shown by the Carrie Bow Cay and 

other USNM specimens. Traditionally the typical 

form (sometimes called P. clavaria Lamarck, 1816) 

has thick branches (over 10 mm in diameter) and 

clavate branch tips; forma furcata has branches 

about 10 mm in diameter, without swollen 

branch tips; and forma divaricata has thin 

branches (5-6 mm in diameter), which often 

bifurcate near the tip. 

Family FAVIIDAE 

Favia fragum (Esper, 1797) 

FIGURE I25d 

Favia fragum.—Vaughan, 1901:303-304, pl. 8.—Squires, 

1958:253, pl. 42: fig. 2.—Roos, 1971:68-69, pis. 26-27, pl. 

28: fig. a.—Smith, 1971:79-80, pis. 16-17.—York, 1971: 

21, pl. 9: figs. 7-8.—Colin, 1978:240, 241 [color fig.], 242, 

247. 

Small, plocoid, encrusting or hemispherical colonies usually less than 10 cm in diameter. Calices 

round or irregular in outline; mono-, di-, or tricentric; 5-6 mm in diameter; 36-40 septa/calice. 

Costae dentate. Columella spongy. Brown or yellow. 

HABITAT.—Common in shallow lagoonal and 

back-reef areas (0.5-1 m). Bathymetric range: 

0.2-30.0 m. 
284 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

m 

FIGURE 125.—a, Porites pontes i. divaricata, Twin Cays, 0.7 m; b, P. porites porites, same specimen 

as Figure 124e; c, P. pontes furcata, exact locality at Carrie Bow Cay unknown; d, Favia fragum, 

reef crest, 0.5 m; e,f Diploria clivosa, exact locality at Carrie Bow Cay unknown. (Scale bars: a, 

d,f= 0.5 cm; b, c, e = 2 cm.) 
NUMBER 12 285 

Diploria clivosa (Ellis and Solander, 1786) 

FIGURE 125*-/ 

Maeandrina clivosa.—Matthai, 1928:71-76. 

Diploria clivosa.—Squires, 1958:253, pl. 42: fig. 2.—Roos, 

1971:69-70, pl. 28: fig. b, pis. 30-31.—Smith, 1971:80-81, 

pis. 19-20.—York, 1971:23, pl. 10: figs. 1-2.—Olivares 

and Leonard, 1971:56, pl. 3: figs, A-B.—Colin, 1978:244 

[color figs.], 250. 

Meandroid; encrusting or hemispherical, 

knobby colonies up to 1 m in diameter. Valleys 

narrow (3.5-6.0 mm across) and variable in 

length: usually long but sometimes very short, 

even monocentric. Septa thin, in two alternate 

series; 25-38 septa/cm along valley. Principal 

septa have paliform lobes. Columella spongy. 

Color variable: yellow, green, bluish, or grey. 

HABITAT.—Common in lagoon and back-reef 

areas (0.5-2.0 m). Also present in spur and groove 

zone (6 m). Bathymetric range: 0-15 m. 

Diploria strigosa (Dana, 1848) 

FIGURE \26a-b 

Platygyra viridis.—Vaughan, 1901:306-308, pis. 9-13. 

Maeandrina cerebrum.—Verrill, 190la:74-78, pl. 10: fig. 1, pl. 

12: fig. 4, pl. 14: figs. 4-5.—Matthai, 1928:55-63. 

Diploria strigosa.—Squires, 1958:253-254, pl. 42: fig. 1.— 

Roos, 1971:70-71, pl. 29.—York, 1971:23, pl. 23: figs. 7-

8.—Olivares and Leonard, 1971:56, pl. 3: figs. C-D.— 

Erhardt, 1974, fig. Ic—Tresslar, 1974:122, pl. 10.—Colin, 

1978:241 [color fig.], 246 [figs.], 247, 250. 

Meandroid; large, hemispherical, evenly convex colonies up to 2 m in diameter. Valleys 6-9 

mm across and usually continuous, not short; 15-

20 septa/cm, most of which reach columella 

through paliform lobes. Columella spongy. Dull 

yellow or greenish brown. 

HABITAT.—Common in back-reef area (0-2 m) 

and found once in sand trough of outer fore reef 

(28 m). Bathymetric range: 0-40 m. 

DISCUSSION.—Diploria strigosa is often confused 

with D. clivosa. The former is distinguished by its 

even convexity of corallum shape; longer, deeper, 

and wider valleys; and less numerous septa/cm. 

Diploria labyrinthiformis (Linnaeus, 1758) 

FIGURE \26c-d 

Meandra labyrinthiformis.—Verrill, 1901a:70-73, pl. 10: figs. 

1,3. 

Maeandrina labyrinthiformis.—Matthai, 1928:63-71. 

Diploria labyrinthiformis: Roos, 1971:71, pl. 32.—Smith, 1971: 

81, pl. 21.—York, 1971:23, pl. 10: figs. 3-6.—Erhardt and 

Meinel, 1975, fig. 6.—Werding and Erhardt, 1976, pl. 2: 

fig. 1.—Colin, 1978:245 [color figs.]. 250-251. 

Meandroid; large, hemispherical, evenly convex colonies up to 2 m in diameter. Valleys 5-8 

mm across and continuous. Collines grooved by 

distinct ambulacra. 14-17 septa/cm, most of 

which reach columella through paliform lobes. 

Columella spongy. Orange-yellow or brownish 

yellow. 

HABITAT.—Common in back-reef area (0.5-2.0 

m). Bathymetric range: 0-43 m. 

Manicina areolata (Linnaeus, 1758) 

FIGURE 126*-/ 

Manicina areolata.—Vaughan, 1901:305, pl. 4: figs. 2-3.— 

Matthai, 1928:80-91.—Squires, 1958:254-255, pl. 37: figs. 

1-3.—Almy and Carrion-Torres, 1963:152-153, pl. 1 lb.— 

Roos, 1971:72 pl. 35.—Smith, 1971:83, pis. 25-27.—York, 

1971:27, pl. 12: figs. 1-3.—Werding and Erhardt, 1976, 

pl. 4: fig. 2.—Colin, 1978:248 [color figs.], 251, 254. 

Meandra areolata.—Verrill, 1901a:81-84, pl. 11: figs. 1-2, pl. 

12: figs. 1-3. 

Manicina gyrosa.—Matthai, 1928:91-95. 

Manicina mayori.—Smithr 1971:83-84.—York, 1971:27, pl. 

12: figs. 4-5. 

Meandroid colonies consisting of one continuous central valley with numerous side valleys, or 

a discontinuous series of sinuous valleys. Corallum usually unattached, with a short pedicel or 

flat base. Collines bear 12-18 septa/cm. Principal 

septa have large paliform lobes. Columella large 

and spongy. Yellow to brown, grey, or green. 

HABITAT.—Forma areolata is common in shallow back and patch reefs (York, 1971); however, 

it was most abundant at Carrie Bow Cay on the 

outer fore reef (16-27 m). One specimen of forma 

mayori Wells, 1936, was collected from the spur 

and groove zone (10 m). Bathymetric range: 0-

65 m. 

DISCUSSION.—Two forms of this species are rec-
286 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

I f ME&&*8 & 

FIGURE 126.—a, b, Diploria strigosa, exact locality at Carrie Bow Cay unknown; c, d, D. 

labyrinthiformis, off Andros, Bahamas; e, Manicina areolata i. areolata, Twin Cays, 1 m; / M. areolata 

mayori, spur and groove zone. (Scale bars: a, c, e = 2 cm; b, d,f= 1 cm.) 
NUMBER 12 287 

ognized (Wells, 1973). Manicina areolata forma 

areolata is the smaller form, rarely longer than 15 

cm, with one continuous central valley, several 

side valleys, and generally a short central stalk. 

Forma mayori occurs as large (up to 30 cm in 

diameter), hemispherical boulders, with many 

sinuous, disconnected valleys and a flat base. 

Colpophyllia natans (Houttuyn, 1772) 

FIGURE 127a-b 

Colpophyllia nalans.—Matthai, 1928:101-107 [not pl. 71: fig. 

9, = C. breviserialis].—Roos, 1971:73, pl. 33.—Smith, 1971: 

82-83, pl. 23.—Wells, 1973, fig. 15.—Colin, 1978:249 

[color figs.], 254. [Not York, 1971:25, pl. 11, = C. breviserialis. ] 

Large but light-weight, meandroid colonies up 

to 1 m in diameter. Valleys long and sinuous, 15-

20 mm wide, 10-12 mm deep. Collines bear 8-9 

septa/cm. Collines always have two thin parallel 

ridges. Rudimentary lamellar columella. Green, 

brown, or yellow. 

HABITAT.—Common at Carrie Bow Cay: found 

in back reef (1-2 m), sand trough (22 m), and 

fore-reef slope (18-27 m). Bathymetric range: 

0.5-55.0 m. 

DISCUSSION.—Colpophyllia amaranthus (Houttuyn, 1772) may be a form of C. natans. It has 

been characterized as a smaller, stalked colony 

with shorter, straight valleys and more septa/cm. 

Cladocora arbuscula (Lesueur, 1821) 

FIGURE 128a 

Cladocora arbuscula.—Roos, 1971:65, pl. 22: figs. a-b.—Smith, 

1971:84, pl. 28.—York, 1971:27 pl. 12: figs. 6-8.—Olivares, 1971:75, pl. 1: figs, A-B.—Colin, 1978:252 [color 

fig-], 255. 

Ramose, bushy, unattached colonies up to 30 

cm in diameter. Calices only at ends of branches. 

Corallites cylindrical, branch and calicular diameters 3-4 mm. Branches distinctly costate. 

About 36 septa/calice. Columella small and papillose. Brown. 

HABITAT.—Found only in shallow back-reef 

Thalassia zone (0.5-2.0 m). Bathymetric range: 

0.5-21.0 m. 

Montastrea annularis (Ellis and Solander, 

1786) 

FIGURE 127c-d; PLATE 4: center left, bottom left, bottom right 

Orbicella acropora.—Vaughan, 1901:301-302, pis. 6-7. 

Orbicella annularis.—Verrill, 1901a:94-96, pl. 15: figs. 1-2.— 

Vaughan, 1919:364-375. 

Orbicella hispidula Verrill, 1901a:96, pl. 15: figs. 3a-b. 

Montastrea annularis.—Squires, 1958:256, pl. 40: fig. 3, pl. 41: 

figs. 1-2.—Roos, 1971:65-66, pl. 24: fig. b, pl. 25: fig. b.— 

Smith, 1971:85-86, pis. 31-32.—York, 1971:29, pl. 13: 

figs. 1-8.—Tresslar, 1974:122 pl. 13.—Colin, 1978:230, 

243, 253 [color figs.], 258-259. 

Massive, plocoid colonies up to 2 m in height 

or diameter. Shape variable: hemispherical, flat 

encrusting, platy, or columnar (see Graus and 

Macintyre, herein: 441). Calices 2-4 mm in diameter; 24 septa/calice, 12 of which reach the 

well-developed, trabecular columella. Yellowish 

brown or green. 

HABITAT.—Very common in all reef environments: back reef (0.3-2.0 m), spur and groove 

zone (10-18 m), and outer fore reef (16-25 m). 

Bathymetric range: 0.3-80.0 m. 

Montastrea cavernosa (Linnaeus, 1767) 

FIGURE 127*-/ 

Orbicella cavernosa.—Verrill, 1901a:102-103.—Vaughan, 

1919:380-384. 

Montastrea cavernosa.—Squires, 1958:255-256, pl. 40: figs. 1-

2.—Roos, 1971:66-67, pl. 22: fig. c, pl. 23.—Smith, 1971: 

86-87, pl. 33.—York, 1971:31, pl. 14: figs. 1-4.—Laborel, 

1971:198-200, pl. 5: figs. 2-4.—Tresslar, 1974:122, pl. 

14.—Colin, 1978:8 [fig.], 256 [color figs.], 259, 262. 

Montastrea braziliana.—Smith, 1971:86. 

Massive, plocoid colonies up to 2 m in diameter. Hemispherical, flat, or encrusting in shape; 

often with small, irregular knobs. Calices 5-11 

mm in diameter; 48 septa/calice, 24 of which 

reach the well-developed, trabecular columella. 

Columellar trabeculae usually swirled in clockwise direction. Brown, green, olive, or red. 

HABITAT.—Common in spur and groove zone 

(16-20 m) and entire outer fore reef (16-26+ m). 

Bathymetric range: 0.5-95 m. 
288 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 127.—a, b, Colpophyllia natans, exact locality at Carrie Bow Cay unknown; *, Montastrea 

annularis, exact locality at Carrie Bow Cay unknown; d, M. annularis, fore-reef slope, 24-27 m; 

* , / M. cavernosa, exact locality at Carrie Bow Cay unknown. (Scale bars: a, c, * = 2 cm; b = 1 

cm; d,f= 0.5 cm.) 
NUMBER 12 289 

FIGURE 128.—a, Cladocora arbuscula, Thalassia area, 1 m; b-d, Astrangia solitaria, Curlew Bank, 6 

m; *, Phyllangia americana, Twin Cays, on a mangrove root, 0.5 m, specimen coated with NH4C1; 

/ Colangia immersa, inner reef slope, 16 m. (Scale bars: a, b = 1 cm; c-f = 0.2 cm.) 
290 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Family RHIZANGIIDAE 

Phyllangia americana Milne Edwards and 

Haime, 1849b 

FIGURE 128* 

Phyllangia americana.—Roos, 1971:74, pl. 36.—Smith, 1971: 

87-88.—Olivares and Leonard, 1971:64, pl. 8: figs. C-D.— 

Calin, 1978:257 [color fig.], 262-263. 

Small, encrusting colonies that increase by extratentacular budding from edge zone. Corallites 

cylindrical, about 10 mm in calicular diameter 

and 10 mm tall. Calices have 36-48 septa with 

6-7 highly exsert primaries. Paliform lobes often 

present on S3. Columella rudimentary, spongy. 

Color of corallum brown. 

HABITAT.—Not found at Carrie Bow Cay, but 

present at Twin Cays (4 km NW of Carrie Bow 

Cay), encrusting a living Strombus gastropod and 

a mangrove root (0.5-1.0 m). Ahermatypic. Bathymetric range: 0.3-17 m. 

Astrangia solitaria (Lesueur, 1817) 

FIGURE I28b-d 

Astrangia solitaria.—Roos, 1971:74, pl. 34.—Smith, 1971: 

87.—Wells, 1972:2-4, figs. 1-5.—Colin, 1978:257 [color 

fig.], 262. 

Encrusting colonies that increase by extratentacular budding from basal stolons. Stolons often 

subsequently obscured, giving the appearance of 

solitary corallites. Corallites cylindrical, measuring 3-6 mm in calicular diameter, up to 20 mm 

tall. Calices have 36-48 septa; the fourth cycle is 

rarely complete. Narrow, multiple paliform teeth 

on inner edges of all but last cycle of septa. 

Columella rudimentary, papillose. Corallum 

brown. 

HABITAT.—Cryptic. Found in crevices and on 

undersides of platy corals or overhangs. Common 

at Curlew Bank (6 m) and buttress zone (6 m). 

Ahermatypic. Bathymetric range: 0.2-43 m. 

Colangia immersa Pourtales, 1871 

FIGURE 128/ 

Encrusting colonies that increase by extratentacular budding from edge zone. Coenosteum 

between corallites often obscured, which gives 

them the appearance of solitary corallites. Corallites cylindrical, measuring 6-8 mm in calicular 

diameter and up to 10 mm tall. Calices have 36-

48 septa with six highly exsert primaries. Welldeveloped paliform lobes on S3. Columella papillose or lamellar. Polyp light green with colorless 

tentacles. Corallum white, speckled with brown. 

HABITAT.—Cryptic. Found rarely on undersides of platy corals on inner reef slope (17 m) 

and fore-reef slope (26 m). Ahermatypic. Bathymetric range: 3-95 m. 

Family MEANDRINIDAE 

Meandrina meandrites (Linnaeus, 1758) 

FIGURE 129a-c 

Meandrina braziliensis.—Verrill, 1901b: 190-192, fig. 14, pl. 34: 

fig. 1.—Smith, 1971:89-90, pl. 36. 

Meandrina meandrites.—Vaughan, 1901:296-297, pl. 3, pl. 4: 

fig. 1, pl. 5.—Roos, 1971:76, pl. 40.—Smith, 1971:89, pl. 

35.—Erhardt, 1974, figs. 2-3.—Colin, 1978:261 [color 

figs.], 263, 264 [color fig.], 266. 

Pectinia maeandrites.—Matthai, 1928:161 -166. 

Pectinia brasiliensis.—Matthai, 1928:167-169. 

Meandrina braziliensis.—Laborel, 1971:201-207, figs. 2-5, pl. 

6: figs. 3-4. 

Meandrina danae.—Smith, 1971:90. 

This species occurs in three forms. Forma meandrites produces massive, meandroid, flat or convex 

colonies up to 1 m in diameter. The sinuous 

valleys are long, about 10 mm wide, and continuous. Forma danae (Milne Edwards and Haime, 

1848), is much smaller (up to 15 cm long), and 

usually unattached. It has one long, continuous 

valley, which usually bifurcates at both ends, and 

smaller, paired valleys projecting perpendicular 

to the main valley. Forma brasiliensis (Milne Edwards and Haime, 1848), is similar to the former 

but does not have bifurcated valley tips. It is most 
NUMBER 12 291 

FIGURE 129.—a, b, Meandrina meandrites i. meandrites, exact locality at Carrie Bow Cay unknown; 

*, M. meandrites f. danae, Wee Wee Cay, 5-6 m; d, e, Dichocoenia stokesi, patch reef area, 7 m;fi D. 

stokesi, exact locality at Carrie Bow Cay unknown, protuberant short calices, "stellans" form; g, 

D. stokesi, fore-reef slope, 30 m, "stellaris" form. (Scale bars: a = 5 cm; b, * = 0.5 cm; c-g = 2 
292 SMITHSONIAN CONTRIBUTIONS TO THE MARTNE SCIENCES 

common off Brazil. All forms have 6-8 wide 

primary septa per cm alternating with smaller 

septa. Columella lamellar. Brown to yellow. 

HABITAT.—Forma meandrites is very common 

on the outer fore reef, especially the fore-reef slope 

(22-40 m). It also occurs in the spur and groove 

zone (6-8 m) and at adjacent patch reefs (7 m). 

Forma danae was not seen at Carrie Bow Cay but 

was collected at Wee Wee Cay (11 km SW of 

Carrie Bow Cay) in a Thalassia area (4-6 m). 

Forma brasiliensis was not collected at Carrie Bow 

Cay. Bathymetric range 0.5-80 m. 

DISCUSSION.—Meandrites meandrites forma danae 

bears the same morphological relationship to the 

typical form as Manicina areolata forma areolata 

bears to M. areolata forma mayori. The three forms 

of M. meandrites are often considered as separate 

species. 

Dichocoenia stokesi Milne Edwards and 

Haime, 1848 

FIGURE \29d-g 

Dichocoenia stokesii.—Mathai, 1928:198-201.—Squires, 1958: 

257, pl. 34: fig. 4.—Roos, 1971:77-78, pis. 41-43. 

Dichocoenia stellaris.—Matthai, 1928:201, pl. 63: fig. 2.— 

Wells, 1973:45-47, figs. 31-33.—Colin, 1978:267, 268 

[color fig.]. 

Dichocoenia stokesi.—York, 1971:33, pl. 15: figs. 1-4.—Wells, 

1973:44-45. figs. 29-30.—Colin, 1978; 265 [color figs.], 

266-267, 268 [color fig.]. 

Plocoid to meandroid, flat or convex colonies 

up to 50 cm in diameter. Calices monocentric or 

polycentric in valleys up to 50 mm long, 3-5 mm 

wide. Costae absent or very short; intercalicular 

surface spinose. Small paliform lobes commonly 

present on principal septa. Columella rudimentary, varying from spongy to lamellar. Yellow, 

brown, or green. 

HABITAT.—Common in most reef environments: patch reefs (5-7 m), spur and groove zone 

(9-10 m), and outer fore reef (18-31 m). Bathymetric range: 2-72 m. 

DISCUSSION.—Dichocoenia stellaris Milne Edwards and Haime, 1848, is considered a distinct 

species by some authors on the basis of its flat 

growth form and short, nonmeandrine calices. 

My examination of the Carrie Bow Cay specimens and additional specimens in the USNM 

collection and at the University of Miami revealed all intergrades of corallum shape (flat to 

spherical) and valley length. I see no reason for 

maintaining them as separate species based on 

skeletal characters. 

Dendrogyra cylindrus Ehrenberg, 1834 

FIGURE I30a-b 

Dendrogyra cylindrus.—Matthai, 1928:170-171.—Roos, 1971: 

76-77, pis. 38-39.—Smith, 1971:90-91, pis. 38-39.—York, 

1971:33, pl. 15: figs. 5-8.—Colin, 1978: 269 [color figs.], 

270, 271 [fig.], 274 [fig.]. 

Meandroid colonies forming vertical pillars up 

to 3 m tall. Valleys sinuous, narrow (3-4 mm), 

and discontinuous. Collines have 7-10 thick, nondentate septa/cm. Collines broad with grooved 

ambulacra; ambulacrum rudimentary or absent 

toward top of pillar. Columella lamellar and 

discontinuous. Dark brown. 

HABITAT.—Rare at Carrie Bow Cay. Collected 

only in spur and groove zone (14 m) and back 

reef (1 m). Bathymetric range: 1-20 m. 

Family MUSSIDAE 

Mussa angulosa (Pallas, 1766) 

FIGURE I30c-d 

Mussa angulosa.—Matthai, 1928:204-208 [not pl. 64: fig. 2, 

pl. 68: fig. 2].—Roos, 1971:79-80, pis. 45-46.—Smith, 

1971:92, pl. 42 [not pl. 41].—Tresslar, 1974:123, pl. 15.— 

Colin, 1978:270, 272 [color figs.], 275. 

Large, phaceloid to meandroid-phaceloid colonies with branch diameters up to 5 cm. Calices 

occur at ends of branches; mono- to polycentric 

(up to four centers) with trabecular linkage. Costae dentate near calice, often forming long, thin, 

raised ridges toward base. Septa coarsely dentate. 

Trabecular columella well developed. Large, 

fleshy polyps; pink, purple, brown, or green. 

HABITAT.—Rare at Carrie Bow Cay. Found on 

inner-reef slope (21 m) and fore-reef slope (17-26 

m). Bathymetric range: 1.5-59 m. 
NUMBER 12 293 

FIGURE 130.—a, b, Dendrogyra cylindrus, 14 m; c, d, Mussa angulosa, inner reef slope, 21 m; *, / 

Scolymia lacera, fore-reef slope, 20-28 m. (Scale bars: a, c, e = 2 cm; b, d = 1 cm;/ = 0.5 cm.) 
294 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Scolymia lacera (Pallas, 1766) 

FIGURE 130*-/ 

Scolymia lacera.—Wells, 1964:381-382, pl. 22: figs. 5-6.— 

Roos, 1971:78-79, pl. 44.—Smith, 1971:92, pl. 41.—Lang, 

1971:952-957, figs. 1-2.—Wells, 1971:960-963, figs. 1, 2, 

4, 6.—Laborel, 1971:220, pl. 7: fig. 6.—Colin, 1978:273-

274 [figs.], 275, 278. 

Solitary, subcylindrical, firmly attached coralla 

u p to 15 cm in diameter (but more often 5-6 cm). 

Calice round to elliptical in outline. U p to six 

cycles of septa; last cycle usually incomplete. 

Septa of first three cycles larger and more exsert 

than others. Septa coarsely dentate, less than five 

triangular teeth/cm. Costae dentat e like septa. 

Columella large, elongate, trabecular. Polyps 

large and fleshy; green, brown, or red. 

HABITAT.—Commo n on the fore-reef slope (14-

40 m) and sand trough of outer fore-reef (21 m). 

Bathymetric range: 14-80 m. 

Scolymia cubensis (Milne Edwards and Haime, 

1849a) 

FIGURE \3\a-b 

Scolymia cubensis.—Lang, 1971:952-957, 2 figs.—Wells, 1971: 

960-963, figs. 1, 3, 5, 7.—Tresslar, 1974:123, pl. 16.— 

Colin, 1978:273, 276 [color figs.], 278. 

Solitary, subcylindrical, firmly attached coralla 

u p to 10 cm in diameter. U p to six cycles of septa, 

last cycle incomplete. Septa finely dentat e with 

at least five tall, attenuated teeth/cm. Costae also 

dentate. Columella large, trabecular. Dark 

brown. 

HABITAT.—Foun d only on fore-reef slope (30-

40 m). Bathymetric range: 20-80 m. 

DISCUSSION.—Scolymia cubensis is distinguished 

from the very similar S. lacera by its finer septal 

dentition and its less exsert and thinner S1-3. A 

third similar species, S. wellsii Laborel, 1967, is 

distinguished by its very tall, laciniate septal dentition. Small coralla (smaller than 3 cm) of all 

three species are difficult to distinguish. 

Isophyllia sinuosa (Ellis and Solander, 1786) 

FIGURE \3\c-d 

Isophyllia dipsacea.—Verrill, 1901a: 118-121, pl. 18: fig. 2, pl. 

19: figs. 2, 3, pl. 20: fig. 2. 

Isophyllia fragilis.—Verrill, 1901a: 121-125, pl. 16: figs. 1, 2, 

pl. 17: figs. 1-7, pl. 18: fig. 1, pl. 19: figs. 1, 4, 5. 

Isophyllia sinuosa.—Matthai, 1928:237-247.—Squires, 1958: 

257-258, pl. 40: fig. 4.—Roos, 1971:81, pis. 48, 50.— 

Smith, 1971:94, pl. 46.—York, 1971:39, pl. 18: figs. 1-3.— 

Colin, 1978:276-277 [color figs.], 279. 

Medium size, meandroid colonies u p to 20 cm 

in diameter. Valleys short (monocentric) or long 

and sinuous; 20-25 m m across. Collines tall and 

narrow, often ridged. About 7-9 septa/cm, the 4 -

5 principal septa reaching the columella. Septa 

bear sharp, attenuated teeth. Columella and calicular linkages trabecular. Variegated, lavender, 

brown, green, yellow. 

HABITAT.—Ver y common in back reef (1-2 m); 

also found in sand trough of outer fore reef (28 

m). Previously known bathymetri c range: 1-10 

m. 

DISCUSSION.—Isophyllia multiflora Verrill, 1901a, 

has been considered a distinct species by some 

authors because of its (1) smaller corallum, (2) 

narrower valleys, an d (3) larger numbe r of septa / 

cm. I consider these characteristics to be within 

the range of variation of /. sinuosa, therefore /. 

multiflora was not distinguished in this report. 

Isophyllastrea rigida (Dana, 1848) 

FIGURE 131*-/ 

Mussa rigida.—Verrill, 1901a: 127-128, pl. 25: figs. 2-3, pl. 

33: fig. 4. 

Isophyllastrea rigida.—Matthai, 1928:263-268, pl. 54: fig. 3, 

pl. 57, figs: 2-3.—Squires, 1958:258, pl. 41: fig. 3.—Roos, 

1971:80-81, pl. 47.—Smith, 1971:92-93, pl. 43.—York, 

1971:35, pl. 16: figs. 1-6.—Colin, 1978:279, 280 [color 

fig-], 282. 

Cerioid, attached, flat to hemispherical colonies up to 15 cm in diameter. Calices mono-, di-, 

or tricentric; polygonal or irregular in shape. A 

monocentric calice about 10 m m in diamete r 

contains 25-30 septa. Coarsely dentat e septa. Ru-
NUMBER 12 295 

FIGURE 131.—a, b, Scolymia cubensis, fore-reef slope, 39 m; *, d, Isophyllia sinuosa, 2 m; *, / 

Isophyllastrea rigida, upper spur and groove zone, 6 m. (Scale bars: a, c, e = 2 cm; b, d,f= 0.5 

cm.) 
296 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

dimentary columella, composed of loose trabeculae. Green, pinkish, or purple. 

HABITAT.—Common in spur and groove zone 

(6-8 m). Also found in back reef (2 m) and forereef slope (20 m). Bathymetric range: 1-20 m. 

Mycetophyllia lamarckiana Milne Edwards 

and Haime, 1848 

FIGURE 132* 

Mycetophyllia lamarckiana.—Matthai, 1928:250-255 [in 

part].—Smith, 1971:93-94, pl. 44 [not pl. 45 = M. 

firox].—Wells, 1973:38, fig. 16.—Colin, 1978:280-281 

[color figs.], 282. 

Not Mycetophyllia lamarckiana.—Roos, 1971:82, pis. 49, 51 [= 

M. ferox].—York, 1971:37, pl. 17: figs. 1-7 [= M.ferox and 

IM. danaand). 

Medium size (up to 12 cm in diameter), weakly 

attached, often turbinate colonies with a nearly 

circular calicular surface. The central, founder 

polyp is surrounded by a ring of 7-10 polyps 

budded circumorally. Subsequently each marginal polyp forms a chain of polyps intramurally, 

a colline forming between each chain. A chain 

may split, in which case a new colline is added. 

Continuous valleys, 10-20 mm wide, all radiate 

from the central polyp. Septa coarsely dentate. 

Direct lamellar linkage. Rudimentary columella. 

Green, brown, or grey. 

HABITAT.—Not common at Carrie Bow Cay: 

found in spur and groove zone (6-8 m) and forereef slope (27 m). Bathymetric range: 3-60 m. 

DISCUSSION.—Several specimens kept alive for 

several days revealed a complete absence of polyp 

tentacles but a distinct submarginal row of tentacles, one corresponding to each septum. These 

short tentacles, recessed in a continuous groove 

3-5 mm from the edge of the corallum, immobilized and transferred prey to the oral surface 

(coenosarc) where it was then transported to an 

adjacent stomodeum. Lack of polyp tentacles has 

been reported for Mycetphyllia reesi Wells, 1973, 

but this is the first report of a ring of submarginal 

tentacles for Scleractinia. 

Mycetophyllia aliciae Wells, 1973 

FIGURES 132/ 133a-* 

Mycetophyllia aliciae Wells, 1973:41-43, figs. 25-28.—Colin, 

1978:283 [figs.], 285 [color figs.], 286 [fig.], 287, 290. 

Thin, platy, weakly attached colonies forming 

nearly circular calicular surfaces up to 30 cm in 

diameter. Circumoral budding followed by intramural budding. Radiating, low collines, often 

discontinuous, flanking wide valleys (20-80 mm) 

of 1-5 rows of calices. Collines may be absent. 

Thick, coarsely dentate septa; 7-12/calice. Inner 

edges of septa usually elevated adjacent to calicular fossa. Direct lamellar linkage. Columella 

absent. Greyish brown, greenish brown, or green. 

HABITAT.—Collected only on fore-reef slope (25 

m). Bathymetric range: 18-75 m. 

DISCUSSION.—Mycetophyllia aliciae resembles M. 

lamarckiana in its similar mode of intratentacular 

budding. The former is distinguished by its 

thicker septa, wider valleys, larger corallum, and 

lack of columella. 

Mycetophyllia danaana Milne Edwards and 

Haime, 1849a 

FIGURE 132a-* 

?Mycetophyllia lamarckana.—York, 1971:37, pl. 17: figs. 1-4. 

Mycetophyllia danaana.—Wells, 1973:38, figs. 17-18.—Colin, 

1978:281 [color fig.], 287. 

Massive, weakly attached, irregularly-shaped, 

meandroid colonies up to 35 cm in diameter. 

Founder polyp (calice) usually not apparent. Sinuous valleys long and continuous, 8-17 mm wide. 

Valleys usually deep (10-15 mm) but may be 

shallow (4 mm). Coarsely dentate septa. Collines 

sometimes discontinuous or hydnophoroid. Direct 

lamellar linkage. Columella rudimentary or absent. Variable in color; often dark green valleys 

with reddish brown collines. Also pale green, grey, 

and brownish yellow. 

HABITAT.—Very common in all areas of outer 

fore reef (16-27 m); also collected from spur and 

groove zone (6-8 m). Bathymetric range: 3-30 m. 

DISCUSSION.—Mycetophyllia danaana can easily be 
NUMBER 12 297 

FIGURE 132.—a, b, Mycetophyllia danaana, fore-reef slope, 24-27 m; c, M. danaana, fore-reef slope, 

16 m, hydnophoroid specimen; d, Gardineria minor, fore-reef slope; *, M. lamarckiana, fore-reef 

slope, 24-27 m; / M. aliciae, fore-reef slope, 20-28 m. (Scale bars: a, c = 2 cm; b, e = 1 cm; d,f 

= 0.2 cm.) 
298 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 133.—a, b, Mycetophyllia aliciae, same specimen as Figure 132 / *, d, M. ferox, fore-reef 

slope, 17 m; *, / Eusmilia fastagiata, spur and groove zone, 10 m. (Scale bars: a, c = 2 cm; b, d, 

e = 1 cm;/ = 0.5 cm.) 
NUMBER 12 299 

confused with M. ferox, especially the shallow 

valley forms of M. danaana. It is distinguished by 

its continuous, wider valleys and its more widely 

spaced calices (usually more than 10 mm apart). 

The characters of valley depth and number of 

septa/calice (see Wells, 1973:38) are not reliable 

to distinguish the species. 

Mycetophyllia ferox Wells, 1973 

FIGURE 133*-a" 

Mycetophyllia lamarckana.—Roos, 1971:82, pis. 49, 51.—York, 

1971: 37, pl. 17, figs. 5-7. 

Mycetophyllia lamarckiana.—Smith, 1971: 93-94, pl. 45. 

Mycetophyllia ferox Wells, 1973:40-41, figs. 22-24.—Werding 

and Erhardt, 1976, pl. 3: fig. 1.—Colin, 1978:284 [color 

fig.], 286 [fig.], 287. 

Meandroid, weakly attached colonies nearly 

circular in outline and up to 35 cm in width. 

Founder polyp (calice) usually not evident. Valleys shallow, usually less than 10 mm in width, 

and sometimes discontinuous: short segments 

completely enclosed by collines. Septa coarsely 

dentate. Direct lamellar linkage. Columella rudimentary or absent. Brown or green. 

HABITAT.—Rare at Carrie Bow Cay: one specimen collected from fore-reef slope (17-18 m). 

Bathymetric range: 10-35 m. 

DISCUSSION.—See discussion of M. danaana. 

Small colonies of Mycetophyllia are generally 

difficult to identify because the pattern of their 

intratentacular budding is not yet established. 

More specimens and analyses are necessary in 

order to distinguish the valid species and/or drop 

the current species to the status of forma. Characteristics of the soft parts, such as tentacle arrangement, may be of taxonomic value. Color 

does not seem to be a reliable character. 

19: figs. 1-6.—Wells, 1973:49, fig. 34a.—Colin, 1978:289 

[color fig.], 291, 292 [color figs.]. 

Phaceloid to phaceloflabellate, bushy colonies 

up to 50 cm in diameter. Corallites usually mono, di-, or tricentric but may produce even longer, 

polycentric series up to 15 cm long. Calices terminal. An average monocentric calice measures 

13 X 19 mm in calicular diameter with 72 septa, 

14 much larger and more exsert than the others. 

Ridged costae correspond to the exsert septa. 

Septal edges smooth. Columella large, composed 

of twisted trabeculae. Brown, green, or yellow. 

HABITAT.—Most common in spur and groove 

zone (6-10 m) but also found on sand trough of 

outer fore reef (21 m). Bathymetric range: 1-

65 m. 

DISCUSSION.—Forma flabellata Wells, 1973, distinguished by its long, polycentric series, which 

produce flabellate branches, was not found at 

Carrie Bow Cay. 

Family FLABELLIDAE 

Gardineria minor Wells, 1973 

FIGURE 132a" 

Gardineria minor Wells, 1973:49-53, fig. 36a-g. 

Solitary, subcylindrical, firmly attached. Calices round, 4-7 mm in diameter. Theca smooth, 

no costae. Four complete cycles of septa. Inner 

edges of Si smooth; higher cycle septa dentate to 

laciniate. Paliform lobes usually present before 

S2. Columella papillose. 

HABITAT.—Cryptic, but common on undersides 

of platy corals (for example, Agaricia) on fore-reef 

slope (18-27 m). Ahermatypic. Bathymetric 

range: 2-241 m. 

Family CARYOPHYLLIIDAE 

Eusmilia fastigiata forma fastigiata (Pallas, 

1766) 

FIGURE 133*-/ 

Eusmilia fastigiata.—Matthai, 1928:190-196.—Roos, 1971: 

83, pl. 52.—Smith, 1971:95, pl. 48.—York, 1971:41, pl. 

Conclusions 

Thirty-seven species of hermatypic Scleractinia 

are reported in this paper. In addition, five ahermatypic species found at Carrie Bow Cay bring 

the total number of Scleractinia for this cay to 

42. In comparison, species records of hermatypic 

corals from other areas of the Caribbean include 
300 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

46 from Jamaica (Wells, 1973), 41 from Panama 

(Porter, 1972), 40 from Bonaire (Scatterday, 

1974), 38 from Ceycen Island, Colombia (Erhardt 

and Meinel, 1975), and 37 from Bahia Concha, 

Colombia (Erhardt, 1974). (These totals have 

been revised in accord with the systematic approach of this paper.) 

The known distribution of corals in western 

Atlantic reefs suggests that habitats not sampled 

in the study area may support other species. For 

example, Mycetophyllia reesi Wells, 1973, and at 

least two species of Agaricia may exist in the 

deeper fore-reef slope (30-70 m), while more intensive collecting in shallow depths may disclose 

the presence oi Porites branneri Rathbun, 1888. In 

fact, two species of hermatypes known previously 

from Belize were not collected in this study: 

Colpophyllia breviserialis Milne Edwards and 

Haime, 1849a (reported as C. natans by York, 

1971, pl. 11: figs. 4-6) and Oculina diffusa Lamarck, 1816 (unreported record in USNM collections) , bringing the total to 39 hermatypes known 

from Belize. In addition to the five ahermatypes 

known from Carrie Bow Cay, five more have 

been reported from Belize: Oculina valenciennesi 

Milne Edwards and Haime (reported by York, 

1971); Caryophyllia ambrosia Alcock and Deltocyathus 

agassizii Pourtales (= Deltocyathus italicus Mitchelotti) (reported by Boone, 1928); Deltocyathus moseleyi Cairns andjavania cailleti (Duchassaing and 

Michelotti) (reported by Cairns, 1979). Thus, the 

scleractinian fauna of Belize totals 49 species, of 

which 39 are hermatypic and 10 are ahermatypic. 

Calcified hydrozoans, commonly listed as part 

of a stony coral fauna, were also found during 

this study: two species of Milleporina and one 

representative of Stylasterina. 

Literature Cited 

Almy, C. C , Jr., and C. Carrion-Torres 

1963. Shallow-Water Stony Corals of Puerto Rico. Caribbean Journal of Science, 3:133-162, 21 plates. 

Bernard, H. M. 

1906. The Family Poritidae, II: The Genus Porites, II: 

Porites of the Atlantic and West Indies. Catalogue 

of the Madreporarian Corals in the British Museum 

(Natural History), 6: 73 pages, 17 plates. 

Boone, L. 

1928. Coelenterata from the Tropical East American 

Seas: Scientific Results of the First Oceanographic 

Expedition of the Pawnee, 1925. Bulletin of the 

Bingham Oceanographic College, 1(5): 1-8, plates 1-3. 

Boschma, H. 

1948. The Species Problem in Millepora. Zoologische Verhandelingen, 1: 115 pages, 15 plates. 

1965. Further Notes on Stylaster roseus (Pallas), Parts 1-2. 

Proceedings, Koninklijke Nederlandsche Akademie van 

Wetenschappen, series C, 68:227-250, plates 1-4. 

Cairns, S. D. 

1979. The Deep-Water Scleractinia of the Caribbean 

Sea and Adjacent Waters. Studies on the Fauna of 

Curaqao, 57(180): 341 pages, 40 plates. 

Colin, P. I. 

1978. Caribbean Reef Invertebrates and Plants. 512 pages. 

Hong Kong: T. F. H. Publications, Inc., Ltd. 

Dana, J. D. 

1848. Zoophytes. In United States Exploring Expedition during the Years 1838-1842 under the Command of Charles 

Wilkes, 7:121-708. 

Dinesen, Z. D. 

1980. A Revision of the Coral Genus Leptoseris (Scleractinia: Fungiina: Agariciidae). Memoirs of the Queensland Museum, 20(1): 181-235, 16 plates. 

Duchassaing, P., and J. Michelotti 

1860. Memoire sur les coralliaires des Antilles. Memoires 

de l'Academie des Sciences de Turin, series 2, 19:279-

365, 10 plates. 

Ehrenberg, C. G. 

1834. Beitrage zur physiologischen Kenntniss der Corallenthiere . . . zur physiologischen Systematik 

derselben. Abhandlungen der Preussischen Akademie der 

Wissenschaften, 1:225-380. 

Ellis, J., and D. Solander 

1786. The Natural History of Many Curious and Uncommon 

Zoophytes, Collected. . .by the late John Ellis, Systemat-
NUMBER 12 301 

ically Arranged and Described by the Late Daniel Solander. xii -I- 208 pages, 63 plates. London. 

Erhardt, H. 

1974. Liste der scleractinen Korallen der Bahia Concha 

bei Santa Marta, Atlantikkiiste Kolumbien. Senckenbergiana Biologica, 55:399-407, 5 figures. 

Erhardt, H., and W. Meinel 

1975. Die scleractinen Korallen der Insel Ceycen, Islas 

San Bernardo, vor der Kolumbianischen Atlantikkiiste. Philippia, 2:236-247, 7 figures. 

Esper, J. C. 

1797. Fortsetzungen der Pflanzenthiere. 230 pages. Niirnberg. 

Goreau, T. F., and J. W. Wells 

1967. The Shallow-Water Scleractinia of Jamaica: Revised List of Species and Their Vertical Distribution Range. Bulletin of Marine Science, 17:442-453. 

Gregory, J. W. 

1895. Contributions to the Paleontology and Physical 

Geology of the West Indies. Quarterly Journal of the 

Geological Society of London, 51:255-312, plate 11. 

Heller, C. 

1868. Die Zoophyten und Echinodermen des Adriatischen Meeres. Zoologisch-Botanische Verhandlungen, 

18:1-88, 3 plates. 

Houttuyn, M. 

1772. Natuurlyke Historic of uitvoerige Beschryving ver Dieren, 

Planten en Mineraalen, volgens het Samenstel van den 

Heer Linnaeus. Deel 1, 614 pages, plates 126-138. 

Amsterdam. 

Laborel, J. 

1967. A Revised List of Brazilian Scleractinian Corals 

and Description of a New Species. Postilla, 107:1-

14, figures 1-4. 

1971. Madreporaires et Hydrocoralliaires recifaux des 

cotes bresilliennes. Resultats Scientifiques des Campagnes de la Calypso, 9(36)25:171-229, 8 plates. 

Lamarck, J.B.P.A. de 

1816. Histoire des polypes. In Histoire naturelle des animaux 

sans vertebres, 2: 568 pages. 

Lang, J. C. 

1971. Interspecific Aggression by Scleractinian Corals, 

1: The Rediscovery of Scolymia cubensis. Bulletin of 

Marine Science, 21:952-959, 2 figures. 

Lesueur, C. A. 

1817. Observations on Several Species of the Genus 

Actinia, Illustrated by Figures./owrna/ of the Academy 

of Natural Sciences of Philadelphia, 1:149-154, 169-

189, plate 8. 

1821. Description de plusieurs animaux appartenant 

aux polypiers lamelliferes de M. le Chev. de Lamarck. Memoires du Museum d'Histoire Naturelle, 6: 

271-298, plates 15-17. 

Linnaeus, C. 

1758. Systema Naturae. Edition 10, volume 1, 824 pages. 

1767. Systema Naturae. Edition 12, volume 1, part 2, pages 

533-1327. 

Lyman, T. 

1859. On a New Species of Coral (Astrea decactis). Proceedings of the Boston Society of Natural History, 6:274-

278. 

Matthai, G. 

1928. A Monograph of the Recent Meandroid Astreidae. In Catalogue of the Madreporarian Corals in the 

British Museum (Natural History), 7: 288 pages, 72 

plates. 

Milne Edwards, H., and J. Haime 

1848. Recherches sur les polypiers, memoire 4: Monographic des Astreides. Annates des Sciences Naturelles, 

series 3, 10:209-320, plates 5-9. 

1849a. Recherches sur les polypiers, memoire 4: Monographic des Astreides. Annates des Sciences Naturelles, 

series 3, 11:235-312. 

1849b. Recherches sur les polypiers, memoire 4: Monographic des Astreides. Annates des Sciences Naturelles, 

series 3, 12:95-197. 

1851. Recherches sur les polypiers, memoire 6: Monographic des Fongides. Annates des Sciences Naturelles, 

series 3, 15:73-144. 

Olivares, M. A. 

1971. Estudio taxonomico de algunos madreporarios del 

Golfo de Cariaco, Sucre, Venezuela. Boletin del 

Instituto Oceanografico de la Universidad de Oriente, 10: 

73-78, 2 plates. 

Olivares, M. A., and A. B. Leonard 

1971. Algunos corales petreos de la Bahia de Mochima, 

Venezuela. Boletin del Instituto Oceanografico de la 

Universidad de Oriente, 10:49-70, 11 plates. 

Pallas, P. S. 

1766. Elenchus Zoophytorum. xvi + 28 + 451 pages. HagueComitum. 

Porter, J. W. 

1972. Ecology and Species Diversity of Coral Reefs on 

Opposite Sides of the Isthumus of Panama. In M. 

L. Jones, editor, The Panamaic Biota: Some Observations Prior to a Sea-Level Canal. Bulletin of 

the Biological Society of Washington, 2:89-116. 

Pourtales, L. F. 

1871. Deep-Sea Corals. In Illustrated Catalogue of the Museum of Comparative Zoology, 4:93 pages, 8 plates. 

Rathbun, R. 

1888. Annotated Catalogue of the Species of Porites and 

Synaraea in the United States National Museum, 

with a Description of a New Species of Porites. 

Proceedings of the United States National Museum, 10: 

354-366. 

Roos, P. J. 

1971. The Shallow-Water Stony Corals of the Nether-
302 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

lands Antilles. Studies on the Fauna of Curaqao, 

37(130): 108 pages, 53 plates. 

Scatterday, J. W. 

1974. Reefs and Associated Coral Assemblages off Bonaire, Netherlands Antilles, and Their Bearing on 

Pleistocene and Recent Reef Models. Proceedings of 

the Second International Coral Reef Symposium, 2:85-

106. 

Smith, F.G.W. 

1971. Atlantic Reef Corals. 2nd edition, 164 pages, 48 

plates. Coral Gables, Florida: University of Miami 

Press. 

Squires, D. F. 

1958. Stony Corals from the Vicinity of Bimini, Bahamas, B.W.I. Bulletin of the American Museum of Natural History, 115:215-262, plates 28-43. 

Stoddart, D. R. 

1962. Three Caribbean Atolls: Turneffe Islands, Lighthouse Reef, and Glover's Reef, British Honduras. 

Atoll Research Bulletin, 87: 151 pages. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays, October 30-31, 1961. Atoll 

Research Bulletin, 95: 142 pages. 

Thorpe, J. E., and P. K. Bregazzi 

1960. Experiments and Observations on the Corals at 

Rendezvous Cay. In Cambridge Expedition to British 

Honduras, General Report, pages 22-28. 

Tresslar, R. C. 

1974. Corals. In T. J. Bright and L. H. Pequegnat, 

editors, Biota of the West Flower Garden Bank, pages 

116-139, plates 1-17. Houston, Texas: Gulf Publishing Co. 

Vaughan, T. W. 

1901. The Stony Corals of the Porto Rican Waters. 

Bulletin of the United States Fish Commission for 1900, 

20(2):290-320, 38 plates. 

1919. Fossil Corals from Central America, Cuba, and 

Puerto Rico, with an Account of the American 

Tertiary, Pleistocene, and Recent Coral Reefs. 

Bulletin of the United States National Museum, 103: 

189-524, plates 68-152. 

Verrill, A. E. 

1900. Additions to the Anthozoa and Hydrozoa of the 

Bermudas. Transactions of the Connecticut Academy of 

Arts and Sciences, 10:551-572, plates 67-69. 

1901 a. Variations and Nomenclature of Bermudian, West 

Indian, and Brazilian Reef Corals with Notes on 

Various Indo-Pacific Corals. Transactions of the Connecticut Academy of Arts and Sciences, 11:63-168, 

plates 10-35. 

1901b. Comparisons of the Bermudian, West Indian, and 

Brazilian Coral Faunae. Transactions of the Connecticut Academy of Arts and Sciences, 11:169-206, plates 

10-35. 

Wells, J. W. 

1936. The Nomenclature and Type Species of Some 

Genera of Recent and Fossil Corals. American Journal of Science, 5(31):97—135. 

1964. The Recent Solitary Mussid Scleractinian Corals. 

Zoblogische Mededeelingen, 39:375-384, plates 20-23. 

1971. Note on the Scleractinian Corals Scolymia lacera 

and Scolymia cubensis in Jamaica. Bulletin of Marine 

Science, 21:960-963, 7 figures. 

1972. Some Shallow-Water Ahermatypic Corals from 

Bermuda. Postilla, 156:10 pages, 3 plates. 

1973. New and Old Scleractinian corals from Jamaica. 

Bulletin of Marine Science, 23(1): 16-58, 36 figures. 

Werding, B., and H. Erhardt 

1976. Los Corales (Anthozoa e Hidrozoa) de la Bahia 

Chengue en el Parque Nacional "Tairona'' (Colombia). Mitteilungen aus dem Instituto Colombo-Aleman de Investigaciones Cientificas, 8:45-57, 4 plates. 

York, M. 

1971. Patch Reef Coral Communities of Southern British Honduras and Illustrated Catalogue of Common British Honduras Corals. In A Guidebook for 

the Field Trip to the Southern Shelf of British Honduras, 

appendix I, 50 pages, 23 plates. [21st Annual 

Meeting of the Gulf Coast Association of Geological Societies and the Gulf Coast Section, Society 

of Economic Paleontologists and Mineralogists, 

sponsored by the New Orleans Geological Society.] 
Octocorallia (Cnidaria) from Carrie Bow Cay, Belize 

Katherine Muzik 

ABSTRACT 

At least 36 species of Octocorallia (in 13 genera 

from four families, two orders) were found growing off Carrie Bow Cay, Belize. The distribution 

and abundance of common species were investigated in the main zones of the transect, the 

lagoon, patch reefs, and "gorgonian city" in 

South Water Cut. Observations on natural history, including competition, feeding, and storm 

damage, are recorded. 

Introduction 

This study is the first ever undertaken of the 

Octocorallia of Carrie Bow Cay. Although the 

octocorals, in particular those of the taxonomic 

order Gorgonacea (gorgonians), are a conspicuous 

and characteristic component of Caribbean coral 

reefs, work on their ecology and natural history 

in scarce (Cary, 1914; Kinzie, 1970, 1973; Goldberg, 1973; Opresko, 1973). Since the early monographs of Duchassaing and Michelotti (1860, 

1864) only two workers (Deichmann, 1936; Bayer, 

1961) have made substantial contributions to the 

systematics of the group in the western Atlantic. 

There are, however, several taxonomic lists from 

various Caribbean locations (Hargitt and Rogers, 

1901; Gordon, 1925; Guitart-Manday, 1959; 

Gonzales-Brito, 1970) and one field guide (Cairns, 

1977). Only one species, Plexaura homomalla (Esper) has received,considerable attention from investigators, owing to the commercially important 

prostaglandins in its tissues (Bayer and Weinheimer, 1974). 

Katherine Muzik, Okinawa Expo Aquarium, Motobu-cho, Okinawaken, 905-03, Japan. 

ACKNOWLEDGMENTS.—I thank A. B. Rath for 

diving assistance and fruitful discussion during 

my visit to Carrie Bow Cay, and S. D. Cairns for 

help in revising my manuscript. 

Material and Methods 

Observations were made during 25 SCUBA 

and 12 snorkel dives, 12-26 April 1979, at habitats near Carrie Bow Cay, including all zones of 

the transect (Riitzler and Macintyre, herein: 9), 

the lagoon, patch reefs 0.1-2.0 km due southwest, 

and "gorgonian city," an area 0.6 km north of 

Carrie Bow Cay having abundant octocorals (Figures 134, 135, 136). At two sites, one on the outer 

ridge, the other a patch reef, colonies of Octocorallia were labeled with small numbered plastic 

tags, photographed, identified, and observed during several dives. Photographs were made with a 

Nikonos camera. Abundance was visually estimated. Identifications were made using field 

characters combined with standard techniques of 

sclerite preparations (Bayer, 1961) from branches 

cut from living colonies. All samples collected are 

deposited in the Department of Invertebrate Zoology, National History, Smithsonian Institution. 

Species List 

(* = Collected by K. Riitzler in 1975, identified by F. M. 

Bayer; f = collected by R. J. Larson in 1972, identified by 

R. J. Larson and F. M. Bayer) 

Order ALCYONACEA 

Family ANTHOTHELIDAE 

Erythropodium caribaeorum (Duchassaing and Michelotti) 

Iciligorgia schrammi Duchassaing (Figures 134a", 137a) 

Family BRIAREIDAE 

Briareum asbestinum (Pallas) (Figure 134a) 

303 
304 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Order GORGONACEA 

Family GORGONIIDAE 

Gorgonia mariae Bayer 

G. ventalina Linnaeus (Figure 135; Plate 3: bottom right, 

Plate 4: top left, top right, bottom right) 

Pseudopterogorgia acerosa (Pallas) (Figure 135) 

P. americana (Gmelin) 

P. bipinnata (Verrill)* 

P. kallos (Bielschowsky)f 

P. rigida (Bielschowsky) 

Pseudopterogorgia sp. (Figure 1346) 

Pterogorgia anceps (Pallas) (Figure 135) 

P. citrina (Esper) 

Family PLEXAURIDAE 

Eunicea calyculata (Ellis and Solander) 

E. calyculata forma coronata Bayer 

E. clavigera Bayer 

E. laciniata (Pallas) 

E. mammosa Lamouroux (Figure 1376) 

E. succinea (Pallas) 

E. succinea forma plantaginea (Lamarck) 

Eunicea spp. (4+) 

Muricea atlantica (Kiikenthal) 

Muriceopsis flavida (Lam arck) * f 

M. petila Bayer 

Plexaura flexuosa Lamouroux (Figure 135; Plate 4: bottom 

right) 

P. homomalla (Esper) 

Plexaura sp. 

Plexaurella dichotoma (Esper) 

P. grisea Kunze*f 

Plexaurella sp. 

Pseudoplexaura flagellosa (Houttuyn) (Figure 134*) 

P. porosa (Houttuyn) 

P. wagenaari (Stiasny)f 

Pseudoplexaura sp. 

Thesea plana Deichmann 

FIGURE 134—Underwater photographs of octocoral species characteristic for some habitats 

along the barrier reef study transect near Carrie Bow Cay: a, Briareum asbestinum (0.4 m tall) 

lagoon, sand and turtle grass bottom, 4 m deep; b, Pseudopterogorgia sp. (1.5 m tall), sand trough 

habitat, 24m; *, Pseudoplexaura flagellosa (over 2 m tall), outer ridge near drop-off, 12 m; d, 

Iciligorgia schrammi (1.5 m tall), fore-reef slope, 20 m. 
NUMBER 12 305 

FIGURE 135—"Gorgonian City" in South Water Cut, 5-10 m deep, characterized by strong 

currents and a smooth pavement with abundant gorgonians: Pterogorgia anceps, lower center; 

Plexaura flexuosa, lower left; Gorgonia ventalina, upper left; Pseudopterogorgia acerosa, upper center 

(picture width, 3 m). 

Results 

SPECIES DISTRIBUTION AND ABUNDANCE.—The 

octocorals of Carrie Bow Cay are represented by 

at least 36 species in 13 genera, 4 families, and 

two orders. Traditionally, the Octocorallia have 

been subdivided into seven orders (Bayer, 1956) 

but division into fewer orders has at times been 

accepted (Kiikenthal, 1921; Stiasny, 1939). The 

orders here recognized, distinguished on the basis 

of skeletal morphology, are Protoalcyonaria, Alcyonacea, Gorgonacea, Helioporacea, and Pennatulacea. The two orders that occur at Carrie 

Bow Cay are Alcyonacea and Gorgonacea. The 

order Alcyonacea now includes the former orders 

Stolonifera, Telestacea, and Alcyonacea, and the 

suborder Scleraxonia, all with skeletons composed 

of sclerites. The order Gorgonacea as now defined 

excludes the Scleraxonia and is characterized by 

octocorals with axial skeletons of "gorgonin," a 

collagenous protein. 

The main factor limiting the distribution of 

gorgonians and alcyonaceans is the availability of 

firm substrate; other factors are water movement 

and light (Kinzie, 1970, 1973). Figures 134 and 

135 show octocoral populations and suitable substrates in different habitats. Table 21 lists the 

distribution of selected octocorals near Carrie 

Bow Cay by habitat. Figure 136 illustrates the 

location of taxa within the transect area. The 

highest number of individuals was observed on 

the pavement of "gorgonian city," a habitat approximately midway between Carrie Bow and 

South Water cays. Gorgonian city is characterized by a strong unidirectional tidal current, a 

smooth, hard substrate, and significant light 
306 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Lagoon Back Reef Crest 

High 

Spur and Groove 

Low 

Spur and Groove Trough 

Fore 

Outer Reef 

Ridge | Slope 

1100 1200 1300 1400 1500 1600 meters I 

Gorgonia ventalina 

30

 J 

meters 

Briareum asbestinum 

Erythropodium caribaeorum 

Gorgonia mariae 

Eunicea spp. and Pseudopterogorgia spp. 

Muricea atlantica 

FIGURE 136—Distribution of principal octocoral species 

along barrier reef transect near Carrie Bow Cay. Species are 

arranged in order of appearance and of continuity with 

increasing depth, not taxonomically. 

Plexaurella dichotoma 

Iciligorgia schrammi 

Pseudopterogorgia bipinnata 

Pseudopterogorgia acerosa 

Pseudopterogorgia americana 

Pseudoplexaura flagellosa 

Pseudoplexaura porosa 

Muriceopsis petila 

Thesea plana 

(depth, 5-10 m). Although octocorals are less 

dense in patch reefs than in gorgonian city, almost 

the same species occur in both areas. The lowrelief spur and groove zone and the outer ridge 

are next in abundance and have similar species 

composition. Octocorals in very calm zones (the 

trough and its slopes and the deep fore-reef slope) 

are depauperate and largely restricted to species 

of Pseudopterogorgia, Eunicea, and Pseudoplexaura. 

Only a few species and a few colonies grow in the 

calm lagoon, where substrate is a limiting factor. 

The occurrence oi Erythropodium caribaeorum at Carrie Bow Cay is conspicuously rare as compared to 

its common occurrence reported by Kinzie (1973) 

for Jamaica, who found it often in the rear zone, 

reef flat, crest, and Acropora cervicornis (Lamarck) 

zones. Although present in the lagoon and crest 

at Carrie Bow Cay, E. caribaeorum is more common 

in the low-relief spur and groove, outer ridge and, 

to a depth of 25 m, on the fore-reef slope (Table 

21). The only known species in the genus Erythropodium, it forms low incrusting rather cryptic colonies so that it is easily overlooked. 

NATURAL HISTORY OBSERVATIONS.—The flamingo-tongue snail, Cyphoma gibbosum Linnaeus, was 

frequently observed browsing on Gorgonia ventalina 

and occasionally on species of other genera. It has 

been reported (Birkeland and Gregory, 1975) to 

prefer gorgoniid octocorals. Fish predation, although observed on California species of Muricea 

(Clarke, 1970) and reported by Randall (1967) 

from gut contents of a few West Indian species of 

fishes, was not observed at Carrie Bow Cay. Some 

colonies of Gorgonia ventalina consist of just a main 

axial skeleton with stubby branchlets, 1-3 cm 

long. The cause of this unusual shape is unknown 

and not previously reported but it could be explained by fish or turtle grazing. The axial skeleton appears to be well-covered by living gorgonian tissue. The role that sclerites in the superficial 

soft tissue play in minimizing predation is unknown. Noxious chemistry may also play an important anti-predator role (Norris and Fenical, 

herein: 417). 

Complete overgrowth by competing Millepora 

alcicornis Linnaeus is fairly common in several 
NUMBER 12 307 

TABLE 21.—Occurrence of selected Octocorallia at various Carrie Bow Cay habitats, confirming 

presence but not necessarily absence of species from listed habitats; total records of genera 

Pseudopterogorgia, Eunicea, and Pseudoplexaura indicated as separate categories due to difficulty of 

field identification to species level; where species identification was possible, records for species 

in these genera are given also (+ = common; (+) = uncommon) 

Species 

Erythropodium caribaeorum 

Iciligorgia schrammi 

Briareum asbestinum 

Gorgonia mariae 

G. ventalina 

Pseudopterogorgia spp. 

P. acerosa 

P. americana 

P. bipinnata 

Pterogorgia anceps 

P. citrina 

Eunicea spp. 

E. calyculata 

E. calyculata i. coronata 

E. clavigera 

E. laciniata 

E. mammosa 

E. succinea f. plantaginea 

Muricea atlantica 

Muriceopsis petila 

Plexaura flexuosa 

P. homomalla 

Plexaurella dichotoma 

Pseudoplexaura spp. 

P. flagellosa 

P. porosa 

Thesea plana 

Lagoon 

and 

back 

reef 

(+) 

(+) 

(+) 

(+) 

(+) 

(+) 

(+) 

(+) 

(+) 

Crest 

(+) 

(+) 

+ 

+ 

High 

spur 

and 

groove 

+ 

(+) 

+ 

+ 

+ 

+ 

+ 

+ 

Low 

spur 

and 

groove 

+ 

(+) 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

Transect 

Inner 

slope 

(+) 

+ 

(+) 

+ 

+ 

+ 

Trough 

(+) 

(+) 

(+) 

+ 

(+) 

Outer 

slope 

+ 

+ 

+ 

+ 

Outer 

ridge 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

Forereef 

slope 

+ 

+ 

+ 

(+) 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

Patch 

reef 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

Gorgonian city 

+ 

+ 

+ 

+ 

+ 

4-

+ 

+ 

+ 

+ 

+ 

+ 

+ 

species of octocorals, particularly in Gorgonia and 

Pseudopterogorgia, where the original shape of the 

colony is commonly maintained by the millepore 

and is easily recognizable. Overgrowth by sponges 

is also common. The pinkish Desmapsamma anchorata (Carter), is especially noticeable at the southern edge of gorgonian city, where both octocorals 

and sponges are growing on the rubble and live 

coral in the sandy zone. 

Night dives revealed that octocoral polyps of 

most genera are expanded at night (presumably 

to feed) as well as during the day (Figure 137). 

Gorgonia ventalina, however, was not observed expanded at night. It may feed only during the day, 

or it may rely exclusively on photosynthesis by its 

zooxanthellae for its energy source. Further observations are needed to determine whether Gorgonia does expand its polyps at night. The extent 

of the dependence of octocorals on food capture 

is unknown; all species reported here except Iciligorgia schrammi have abundant zooxanthellae in 

their tissues. This alcyonacean occurs deeper than 
308 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 137—Photomacrographs of expanded polyps: a, Iciligorgia schrammi; b, Eunicea mammosa. 

(Scale: a = 5 mm; b = 2 mm.) 

15-20 m on the fore-reef slope but occasionally 

also on coral mounds in the trough (20-24 m). 

Although long-term studies are necessary to 

determine growth rates, repair of branches harvested with clippers occurred rapidly. In all 20 

colonies (18 species) clipped and revisited at the 

two sites, the remaining living branch was healing 

satisfactorily within four days of being cut. 

In April 1979, loose and abraded toppled colonies of octocorals lay scattered in the spur and 

groove zone and in the lagoon. These dying colonies, certainly more common that I have ever 

observed elsewhere, were apparently dislodged 

from their substrates during hurricane Greta in 

September 1978. The extent of the damage can, 

of course, only be estimated since there are no 

pre-storm abundance data. Similar observations, 

however, were made in Puerto Rico after hurricane Edith (Glynn et al., 1964). Weakening of the 

substratum around a holdfast by boring oiganisms has been postulated as the main cause of 

toppling of the colonies, hence of mortality during 

storms (Kinzie, 1973). 

Discussion and Conclusions 

The Octocorallia of Carrie Bow Cay are abundant and diverse; the species found in this preliminary investigation (at least 36) are comparable 

both in density and number of species with areas 
NUMBER 12 309 

in the Caribbean known to have abundant gorgonians. Kinzie (1973) lists 43 species in Jamaica, 

but seven of them live deeper than the typical 

dives (25 m maximum) of this study. Opresko 

(1973) reports 29 species from three patch reefs in 

the Florida Keys. Although most of the species 

collected at Carrie Bow Cay are known from 

elsewhere in the Caribbean, at least four appear 

to be new, for example, species oi Pseudopterogorgia, 

Plexaurella, and Eunicea. The distribution of octocorals is limited by availability of firm substrate 

as well as by requirements for light and currents. 

Some species exhibit preference for certain habitats. Four areas, the low-relief spur and groove, 

the outer ridge, the patch reefs, and gorgonian 

city are similar in both species composition and 

abundance. The species of the turbulent crest and 

high-relief spur and groove zones differ from species of the deep, calm waters of the trough. Some 

species, such as Pterogorgia citrina, abundant only 

at gorgonian city, exhibit a distribution that is so 

far unexplained. Colonies of Erythropodium caribaeorum, abundant in Jamaica, appear to be less 

common at Carrie Bow Cay. These and other 

findings need further investigation for confirmation and explanation. 

Literature Cited 

Bayer, F. M. 

1956. Octocorallia. In R. C. Moore, editor, Treatise on 

Invertebrate Paleontology, Part F: Coelenterata, pages 

166-231. New York: Geological Society of America and University of Kansas Press. 

1961. The Shallow-Water Octocorallia of the West Indian 

Region. 373 pages. The Hague: Martinus Nijhoff. 

Bayer, F. M., and A. J. Weinheimer, editors 

1974. Prostaglandins from Plexaura homomalla: Ecology, 

Utilization and Conservation of a Major Medical 

Marine Resource, A Symposium. Studies in Tropical 

Oceanography, 12: xii + 165 pages. Miami: University of Miami Press. 

Birkeland, C , and B. Gregory 

1975. Foraging Behavior and Rates of Feeding of the 

Gastropod, Cyphoma gibbosum (Linnaeus). In S. A. 

Earle and R. J. Lavenberg, editors, Results of the 

Tektite Program: Coral Reef Invertebrates and 

Plants. Natural History Museum of Los Angeles County, 

Science Bulletin, 20:57-67. 

Cairns, S. D. 

1977. Guide to the Commoner Shallow-Water Gorgonians (Sea Whips, Sea Feathers and Sea Fans) of 

Florida, the Gulf of Mexico, and the Caribbean 

Region. Sea Grant Field Guide Series, 6: 74 pages. 

Miami: University of Miami Sea Grant Program. 

Cary, L. R. 

1914. Observations upon the Growth-Rate and Oecology of Gorgonians. Carnegie Institute of Washington 

Publication, 182:79-90. 

Clarke, T. A. 

1970. Territorial Behavior and Population Dynamics of 

a Pomacentrid Fish, the Garibaldi Hypsypops rubicunda. Ecological Monographs, 40(2): 189-212. 

Deichmann, E. 

1936. The Alcyonaria of the Western Part of the Atlantic 

Ocean. Memoirs Museum of Comparative Zoology, 53: 

1-317. 

Duchassaing, P., and J. Michelotti. 

1860. Memoire sur les coralliaires des Antilles. Memoires 

de l'Academie des Sciences de Turin, series 2, 19:279-

365. 

1864. Supplement au memoire sur les coralliaires des 

Antilles. Memoires de l'Academie des Sciences de Turin, 

series 2, 23:97-206. 

Glynn, P. W., L. R. Almodovar, and J. G. Gonzalez. 

1964. Effects of Hurricane Edith on Marine Life in La 

Parguera, Puerto Rico. Caribbean Journal of Science, 

4:335-345. 

Goldberg, W. 

1973. The Ecology of the Coral-Octocoral Communities 

off the Southeast Florida Coast: Geomorphology, 

Species Composition and Zonation. Bulletin of Marine Science, 23:465-488. 

Gonzalez-Brito, P. 

1970. Una lista de los Octocorales de Puerto Rico. Caribbean Journal of Science, 10:63-69. 

Gordon, I. 

1925. Gorgonids from Curacao Island. Bijdragen tot der 

Dierkunde, 24:15-24. 

Guitart-Manday, D. 

1959. Gorgonias del litoral de la costa norte de Cuba. 

Acuario Nacional Sibarimar Centro Experimental, 1:1-

24. 
310 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Hargitt, C , and C. Rogers. 

1901. The Alcyonaria of Puerto Rico. Bulletin of the U.S. 

Fish Commission, 20:265-287. 

Kinzie, R. 

1970. The Ecology of the Gorgonians (Cnidaria, Octocorallia) of Discovery Bay, Jamaica. 107 pages. 

Ph.D. dissertation, Yale University. 

1973. The Zonation of West Indian Gorgonians. Bulletin 

of Marine Science, 23:93-155. 

Kiikenthal, W. 

1921. Versuch eines naturlichen Systems der Oktokorallen. Sitzungsberichte der Preussischen Akademie der 

Wissenschaften 4:82-102. 

Opresko, D. M. 

1973. Abundance and Distribution of Shallow-Water 

Gorgonians in the Area of Miami, Florida. Bulletin 

of Marine Science, 23:535-558. 

Randall, J. 

1967 Food Habits of Reef Fishes of the West Indies. 

Studies in Tropical Oceanography, 5:665-847. Miami: 

University of Miami Press. 

Stiasny, G. 

1939. System der Octocorallia. Zoologische Mededeelingen, 

21:367-368. 
Distribution of Sipuncula in the Coral Reef Community. 

Carrie Bow Cay, Belize 

Mary E. Rice 

and Ian G. Macintyre 

ABSTRACT 

A survey of sipunculan fauna associated with 

coral rock and rubble along a transect across the 

coral reef at Carrie Bow Cay, Belize, revealed 

eight species, six of which inhabited burrows 

within rock samples and two of which occurred 

in crevices and crannies. Comprising 95 percent 

of the collection, the four most abundant species 

were, in order of abundance: Lithacrosiphon alticonus Ten Broeke; Aspidosiphon brocki Augener; 

Phascolosoma perlucens Baird; and Paraspidosiphon 

steenstrupi (Diesing). The greatest concentration of 

sipunculans, measured as number per square meter of surface area of rock, occurred in reef-crest 

material. Sipunculans were least abundant in 

material from the patch-reef zone of the back reef 

and from the fore-reef slope. The four most abundant species showed a distinct distributional pattern along the transect. Generally, sipunculans 

from the same reef habitat were densest in fresh, 

relatively unaltered coral rock having little secondary infill of calcite cement and a rather uniform skeletal framework characteristic of species 

of Porites and Acropora. Samples of more highly 

eroded, commonly well-cemented, and presumably older rocks contained few, if any sipunculans. Aspects of sipunculan distribution suggested 

for further study include reproductive strategy 

and variations in substrate characteristics. 

Introduction 

Marine worms of the phylum Sipuncula are 

common inhabitants of coral reef communities 

Mary E. Rice, Department of Invertebrate Zoology, and Ian G. 

Macintyre, Department of Paleobiology, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C. 20560. 

throughout the world. They occupy a variety of 

habitats within the community: sand burrowing 

species occur in sand flats of lagoons and in 

sediments surrounding the bases of coral growths; 

other species inhabit crevices in rubble or live 

within algal mats on surfaces of coral boulders; 

still others dwell in burrows that they excavate in 

coral rubble and in coral-rock framework. These 

rock-boring species, often occurring in great densities, contribute to the erosion of reefs by weakening the supporting structures of the corals and 

thus increasing their susceptibility to breakage 

and destruction by physical stresses related to 

currents and wave action. Numerous papers have 

referred to the boring activities of sipunculans in 

coral limestone and to their possible role in reef 

destruction (Gardiner, 1903; Otter, 1937; Rice, 

1969, 1975a, 1976; Rice and Macintyre, 1972). 

This study is a preliminary survey of the sipunculan fauna associated with the reef at Carrie 

Bow Cay, Belize. From the distributions of sipunculans in different zones of this coral reef community and in different types of coral rock, habitat preferences of the rock-boring species are 

suggested. Reproductive patterns that may have 

a bearing on distribution are also noted. 

ACKNOWLEDGMENTS.—We express our appreciation to D. S. Putnam for his technical contributions to this project and to J. Piraino for photographic assistance. 

Materials and Methods 

Observations and collections were made during 

three 10-day trips to Carrie Bow Cay in 1972, 

311 
312 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Lagoon Back Reef 

Tholassia 

Zone 

0 

Sand & 

Rubble 

Zone 

Patch Reef Zone 

I100 

Rubble & Pavement Zone 

1200 

Reef 

Crest 

Inner Fore Reef 

High Spur & Groove Zone Low Spur & Groove Zone 

i300 l40 0 I500 

Outer Fore Reef 

Inner 

Reef 

Slope 

Sand 

Trough 

Outer 

Ridge 

I600 

Fore Reef 

Slope 

Meters 

Q. 

in 

40 

30 -

y 20 -

-2 10 -

o 

Z 

ZZ 

Lithacrosiphon alticonus 

[;-.. j Aspidosiphon brocki 

| ^ j Phascolosoma perlucens 

Y/A Paraspidosiphon steenstrupi 

~\ Ofhers 

1 2 3 4 5 

Station No. 

FIGURE 138.—Density of sipunculans in coral rock along the transect of the reef, Carrie Bow 

Cay, 1973, 1974. Density is expressed as numbers of sipunculans per kilogram of rock (1973, 

1974) and as numbers of sipunculans per square meter of rock surface (1974), for each of six 

stations. 
NUMBER 12 313 

1973, and 1974. Species of sipunculans on the 

reef were surveyed in 1972, and types of coral 

rock inhabited as well as numbers of sipunculans 

per kilogram of rock were determined in 1973. 

These observations were repeated in 1974 and, in 

addition, numbers of sipunculans per unit of 

surface area of rock were determined. 

COLLECTING SITES.—Sipunculans were collected at numerous sites along a transect across 

different zones of the reef to the north of Carrie 

Bow Cay (Figure 138). The transect extends from 

the Thalassia beds of the lagoon across the back 

reef, reef crest, inner fore reef and outer fore reef 

to a depth of 35 m on the fore-reef slope. This 

transect, which has been the site of many other 

studies, is described in detail by Riitzler and 

Macintyre (herein: 9). Density of sipunculan species was examined at six stations ranging in depth 

from intertidal waters at the reef crest to 35 m on 

the fore-reef slope (Figure 138). 

COLLECTION OF ROCK.—On average, four rocks 

were collected at each station but this number 

varied from two to 10, depending on their availability and size; the greatest number was collected at station 6 on the outer fore-reef slope. 

These were usually free boulders of coral rock 

found on the surface of the sediment. Samples of 

limited size (0.4 to 4.6 kg, averaging 1 to 2 kg) 

were selected at random at each site. 

DESCRIPTION OF ROCK SAMPLES.—Length, 

width, thickness, and general shape of each rock 

were recorded along with associated flora and 

fauna. Rocks were weighed to the nearest gram, 

and surface area was calculated by covering the 

rocks with aluminum foil and converting the 

weight of the foil to square meters, after the 

method of Marsh (1970). 

EXTRACTION OF SIPUNCULANS.—Except for 

small fragments saved for petrographic analysis, 

each rock was completely fractured with a rock 

pick and chisel and all sipunculans were extracted. Species were identified, counted and, in 

most cases, preserved in 70 percent ethyl alcohol. 

A few specimens were kept alive for laboratory 

observations on spawning and developmental 

patterns. 

IDENTIFICATION OF CORAL ROCK.—Component 

coral species of each rock fragment were identified. Other factors examined included the character and distribution of encrusting biota, the 

destruction of coral skeleton by bioerosion, and 

secondary infilling by submarine lithification. 

The extent of secondary infilling by magnesium 

calcite cement is said to be directly related to 

both the length of time and degree of water 

agitation to which the substrata have been exposed (Macintyre, 1977). Borings of known species were saved for a separate study on boring 

mechanisms of rock-dwelling sipunculans (Rice, 

1976). 

Results 

AEREAL DISTRIBUTION.—Of the eight species of 

sipunculans collected from coral rock and rubble 

at Carrie Bow Cay (Table 22), six species were 

found in burrows within the rocks and two in 

crevices or crannies. The four most abundant 

species, which comprised 95 percent of the specimens collected over the three-year period, were 

rock-boring species. In order of abundance, they 

were: Lithacrosiphon alticonus Ten Broeke, Aspidosiphon brocki Augener, Phascolosoma perlucens Baird, 

and Paraspidosiphon steenstrupi (Diesing). 

The most common species (47 percent of specimens collected) was Lithacrosiphon alticonus, which 

is also referred to in the literature by its synonym, 

L. gurjanovae. It was found at all depths along the 

transect, including intertidal sites, but was most 

common in depths of 3 to 35 m. This small 

species, which averages about 10 mm in retracted 

length, possesses an anterior calcareous cone, typical of the genus, that functions to close the 

opening of the boring when the anterior retractable portion of the body—the introvert—is withdrawn (Figure 139a). Endemic to the Caribbean, 

L. alticonus has been reported from beachrock, 

recent unaltered coral rock, and highly eroded 

coral rock (Ten Broeke, 1925; Murina, 1967; 

Rice, 1975a). 

Twenty-three percent of the specimens were 

Aspidosiphon brocki, found in small numbers at 
314 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 22.—Sipunculans collected from coral rock at Carrie Bow Cay, 1972, 1973, 1974 

Sipunculan species 

Aspidosiphon brocki 

Golfingia sp. 

Lithacrosiphon alticonus 

Paraspidosiphon fischeri 

P. speciosus 

P. steenstrupi 

Phascolosoma perlucens 

P. varians 

Number collected 

(3-year total) 

261 

6 

531 

9 

8 

107 

171 

40 

Percent of 

total sample 

23.0 

0.5 

47.0 

0.8 

0.7 

9.4 

15.1 

3.5 

Habitats 

Borings in recent coral limestone; commonly intertidal to 1 m depth; rubble of Porites porites, P. 

astreoides, Acropora palmata 

Crevices and holes of coral rock; scattered throughout coral reef community 

Borings in all types of coral rock from fresh, dense 

coral to extensively bored and eroded limestone; 

most common in deeper water, 3-35 m 

Borings in all types of coral rock; most common in 

rubble of Porites porites in shallow water 

Borings usually in eroded coral rock, with secondary 

infill; most frequent in deeper waters of outer reef 

slope 

Borings in all types of coral rock throughout reef; 

at all depths, most common at reef crest; Porites 

astreoides, Acropora palmata 

Borings in recent coral limestone; shallow waters, 

intertidal to 3 m; abundant in rocks of Acropora 

palmata and Agaricia sp. 

Crevices and holes of coral rock; algal mats covering 

rocks; most abundant in intertidal and shallow 

waters 

various depths along the transect and in great 

densities at the reef crest. The smallest species 

collected, it ranges from 3 to 10 mm in length of 

the trunk. The thickened anterior and posterior 

shields of the trunk are characteristic of the genus 

(Figure 1396), although the posterior shield may 

be weakly developed in specimens that are regenerating after undergoing asexual reproduction. 

Unlike species that reproduce sexually, A. brocki 

reproduces asexually (Figure 139<r) by constricting 

the posterior end to form a juvenile individual; 

thereafter, the posterior end of the adult regenerates (Rice, 1970). This species was first described from the Philippines and has been reported since from calcareous rock throughout the 

Caribbean (Murina, 1967; Rice, 1970, 1975a). 

Phascolosoma perlucens, which comprised 15 percent of the sipunculans collected, was found only 

in shallow waters of the reef crest and the highrelief spur and groove zone. Averaging 30 to 40 

mm in extended length, this long, slender species 

has concentrations of conical papillae at the posterior end and at the base of the introvert. The 

introvert is marked dorsally by bands of reddishbrown pigment (Figure 139a7

). Frequently referred to in the literature by its junior synonym, 

Phascolosoma dentigerum, this circumtropical species 

has been reported as the most common rockboring species in the Caribbean (Rice, 1975a). 

Paraspidosiphon steenstrupi made up only 9 percent of the collection. Although found in small 

numbers, it occurred along most of the transect. 

This species, which has characteristic anterior 

and posterior shields, averages 20 mm in length 

when the introvert is retracted (Figure 139<?). 

Circumtropical in distribution, it is especially 

common throughout the Indo-Pacific and is reported from a number of localities in the Caribbean (Shipley, 1903; Fischer, 1922a, 1922b; Ten 

Broeke, 1925; Murina, 1967; Rice, 1975a). 

The size and shape of the borings of sipunculans in coral rock reflect the size, shape, and 

activity of the inhabiting species (Rice, 1969, 

1975a). Borings of Lithacrosiphon alticonus, a rela-
NUMBER 12 315 

tively inactive species, are generally straight and 

cylindrical, and the closed end of the passage is 

distinctly rounded. The small species, Aspidosiphon 

brocki, has a short, narrow boring, usually straight 

and always near the surface, extending at any 

angle from the surface into the rock. The borings 

of Phascolosoma perlucens, a larger, more active 

species capable of considerable extension and contraction, are long and winding, sometimes extending deep into the interior of the rock. The 

unpredictable course of these borings makes it 

difficult to extract specimens without injury. The 

borings of Paraspidosiphon steenstrupi are also long 

and penetrate far into the rock, but they are 

usually less sinuous than those of P. perlucens. 

DISTRIBUTION ALONG THE TRANSECT.—Available surface area of substrate rock is an important 

limiting factor for sipunculan habitation; among 

rocks of different thickness, shape, and size, a 

comparison of densities of the indwelling sipunculan fauna is most meaningful when numbers of 

sipunculans are related to surface area. In the 

1974 data considered below, densities of species 

at the six transect stations are calculated as numbers per square meter of rock surface (Figure 138). 

The greatest concentration of sipunculans was 

found on the reef crest (station 2). The substratum 

of this zone, partially exposed at low tide and 

subject to strong wave action, consists of in-place 

scattered live coral heads on a coral rock pavement, coral boulders, and coral rubble. The coral 

rock, predominantly Porites astreoides Lamarck and 

Acropora palmata (Lamarck), contains traces of submicrosucrosic magnesium calcite (Macintyre, 

1977) and consists of a relatively unaltered aragonite coral skeleton. All four rocks (totaling 4 

kg) from this station consisted of dead Porites 

astreoides. Growths of coralline and filamentous 

FIGURE 139.—Common sipunculans from Carrie Bow Cay: a, Lithacrosiphon alticonus; b, c, 

Aspidosiphon brocki (specimen in c undergoing asexual reproduction, division will occur at the 

point of the constriction, the posterior portion forming a juvenile individual); d, Phascolosoma 

perlucens; e, Paraspidosiphon steenstrupi. (Scale = 2 mm.) 
316 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

algae, some encrusting sponges and foraminifers 

{Homotrema sp.) were scattered on the rock surfaces. Other than sipunculans, the few living 

organisms within the rocks consisted of a few 

polychaetes and boring barnacles {Lithotrypa sp.), 

and some vermetids and boring sponges near the 

surface. 

All four common rock-boring sipunculan species at Carrie Bow Cay occurred at the reef crest; 

the dominant species, here accounting for 71 

percent of the individuals, was Aspidosiphon brocki. 

The order of abundance of the other three species 

was Paraspidosiphon steenstrupi, and equal numbers 

oi Lithacrosiphon alticonus and Phascolosoma perlucens. 

Other species associated with the rocks, either in 

old cavities or in crevices but showing no evidence 

of forming their own burrows, were Phascolosoma 

varians and an unidentified species of Golfingia. 

The second largest concentration of sipunculans was found in depths of 2 to 3 m in an area 

of large coral buttresses seaward of the reef crest 

(station 3). Known as the high-relief spur and 

groove zone, it is dominated by a substratum of 

living corals, coral rubble, and boulders interspersed with sand patches and loose coral that 

cover a smooth rock pavement. Whereas the reef 

crest is in part intertidal, the high-relief spur and 

groove zone is entirely subtidal. Of the four rocks 

examined (totaling 5.6 kg), one was fresh Porites 

astreoides with no infill. It was difficult to break 

and contained only one sipunculan. Two rocks 

were fresh Acropora palmata and the fourth fresh 

Agaricia sp. One had a trace of submicrosucrosic 

magnesium calcite infill. 

The most abundant species at this station were 

Phascolosoma perlucens and Lithacrosiphon alticonus. 

Only small numbers of Phascolosoma varians, Paraspidosiphon steenstrupi, and Aspidosiphon brocki were 

present. 

Next in abundance of sipunculans was station 

5 on the outer ridge in 15 m of water, where the 

bottom is primarily living coral with patches of 

sand and coral rubble. Two rocks totaling more 

than 5 kg from this area consisted of fresh Porites 

astreoides having little or no infill. One contained 

numerous Lithacrosiphon alticonus and a few Aspidosiphon brocki; its upper surface was covered with 

red boring sponges and encrusting and filamentous algae, whereas the lower surface was almost 

barren, with a lesser amount of crustose coralline 

algae and red sponges. This second rock contained only a few polychaetes and one specimen 

of L. alticonus; around its edges were several 

patches of live coral, also encrusting byrozoans, 

crustose coralline algae, and a white sponge. Its 

internal composition was similar to that of the 

first rock, except that one surface contained more 

submicrosucrosic magnesium calcite infill. 

Next in abundance of sipunculans was station 

4 located in 23 m of water at the base of the inner 

reef slope; here the substratum is mainly sandy 

sediment with small amounts of scattered rubble 

and a few heads of live Montastrea annularis (Ellis 

and Solander), dominated by flattened growth 

forms. Of the three rock samples (6.7 kg) collected, two were M. annularis and one Manicina 

areolata (L.). One Montastrea annularis rock had 

considerable crustose coralline algal cover and 

extensive borings at the surface and contained 

numerous sipunculans of the genus Lithacrosiphon. 

The other M. annularis rock had very little covering growth and was nearly devoid of sipunculans. 

The third rock had an open porous meandroid 

skeleton and large masses of boring sponges near 

the surface. It contained many sipunculans, 

mainly Lithacrosiphon, but they were less concentrated than in the first rock. The sipunculan 

burrows commonly ran parallel to the dissepiments of the coral skeleton. 

The remaining two stations at either end of the 

transect had the fewest sipunculans. Station 1 in 

the patch-reef zone of the back reef (depth about 

1 m) had substrata of scattered rubble and boulders in a sand matrix between live coral patches 

(dominantly Montastrea annularis). The four rocks 

from this station were composed of M. annularis 

(total weight, 3.5 kg). Surfaces adjacent to the 

sand were almost bare, with scattered foraminifers {Homotrema sp.) and some crustose coralline 

algae. On the upper surfaces, crustose coralline 

algae and red and brown filamentous algae were 

common. The degree of infiltration and erosion 
NUMBER 12 317 

and the amoun t of freshly preserved coral skeleton 

varied from a highly eroded skeleton with extensive sediment-rich submicrosucrosic magnesium 

calcite to an almost freshly preserved skeleton. 

T h e highest density of sipunculans was found in 

the rock having the most intact skeleton and the 

least calcite infill. Species of sipunculans at station 1 were Lithacrosiphon alticonus, Aspidosiphon 

brocki, Paraspidosiphon steenstrupi, and Phascolosoma 

varians. 

Station 6 was located on the fore-reef slope in 

depths of 25 to 35 m. Th e substratum exposed 

between the rich cover of octocorals and dominantly platy coral colonies was in-place coral 

framework with a thin cover of Halimeda-rich sand 

and scattered rubble. Th e coral rock from this 

zone was highly eroded and infiltrated with 

sponge growth. Th e 10 rocks collected (total 

weight, 14 kg) were covered with numerous 

sponges, some compoun d tunicates, tunicates, 

coralline algae, "leafy" brown and green algae, 

bryozoans, an d small serpulid worm tubes. These 

rocks were lying on, or were partially embedded 

in the sediment. Thre e were so extensively bored, 

infilled, and lithified with dense microcrystalline 

and porous submicrosucrosic magnesium calcite 

that the coral skeleton could not be recognized. 

Five corals were identified in the other seven 

rocks: Porites astreoides, Meandrina sp. Stephanocoenia 

sp., Agaricia sp., an d Siderastrea siderea (Ellis and 

Solander). Tw o rocks were fresh coral with their 

skeletal structure intact an d with little secondary 

infill. Th e others were water worn and extensively 

bored an d they contained some submarine lithification. Boring sponges ha d infiltrated the rocks. 

Empty borings and other large holes within the 

rocks were filled with sand an d mu d an d patches 

of submicrosucrosic magnesium calcite. Few sipunculans were found in any of these rocks, the 

most occurring in a sample of fresh Porites astreoides 

that contained five Lithacrosiphon alticonus and one 

Paraspidosiphon steenstrupi. Paraspidosiphon speciosus 

was found only in highly eroded samples. A few 

polychaetes, alphaeid shrimp, an d an echiuran 

occurred within the sand-filled holes in the rock. 

Whe n density of sipunculans was measured as 

numbe r per kilogram of rock, the distribution of 

species and relative densities were almost the 

same as when measured as numbe r per square 

meter of surface area. Dat a from 1973 an d 1974 

are compared in Figure 138. 

DISTRIBUTION IN DIFFERING ROC K TYPES.—O f 

the nine types of coral rock in the 1974 collections, 

Pontes, Agaricia, and Acropora specimens contained 

the greatest density (Table 23) and diversity of 

sipunculans. These rocks were not distributed 

equally along the transect; only Porites astreoides 

and Agaricia sp. were collected at more than one 

station. Porites rock, the most common, was found 

at four stations: reef crest, high-relief spur and 

groove, outer ridge, and fore-reef slope. Porites 

rock at the reef crest contained, in order of abundance, Aspidosiphon brocki, Paraspidosiphon steenstrupi, and equal numbers oi Lithacrosiphon alticonus 

and Phascolosoma perlucens. Th e sipunculan fauna 

in Pontes rocks from other stations consisted almost entirely of Lithacrosiphon alticonus (Figure 

140). All Porites samples were recent, unaltered 

coral rock having little secondary calcite infill. 

T h e density of sipunculans varied considerably, 

even amon g rocks of the same composition at the 

same locality (Figure 140). 

OBSERVATIONS ON REPRODUCTIVE ACTIVITY.— 

After removal from coral rock, specimens oi Phascolosoma perlucens, Phascolosoma varians, Paraspidosiphon steenstrupi, and Lithacrosiphon alticonus were 

TABLE 23.—Density of sipunculans in different types of coral 

rocks 

Coral species 

Acropora palmata 

Agaricia sp. 

Manicina sp. 

Meandrina sp. 

Montastrea annularis 

Porites astreoides 

Porites sp. 

Siderastrea siderea 

Stephanocoenia sp. 

Rocks 

Number 

4 

2 

-

-

7 

6 

4 

7 

-

Weight 

(kg) 

10.78 

3.34 

-

-

8.09 

11.10 

5.37 

10.84 

-

Sipunculans 

Total 

number 

75 

27 

-

-

50 

72 

22 

61 

-

Number/ 

kg rock 

6.96 

8.08 

-

-

6.18 

6.49 

4.10 

5.63 

-
318 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

1200 

1000 

800 -

600 -

400 

200 

H I Lithacrosiphon alticonus 

Wl Aspidosiphon brocki 

i H Phascolosoma perlucens 

vfA Paraspidosiphon steenstrupi 

O Others 

Fore - Reef 

Inner Outer 

Outer Ridge 

High Spur & 

Groove Zone 

F©re=R«§f 

ileee 

No, Rocks examined 

FIGURE 140.—Density of sipunculans along the transect in 

Porites astreoides rock, Carrie Bow Cay, 1974. 

kept alive in the laboratory for one to two months 

for observations on spawning. The first two species were observed to spawn in April and Paraspidosiphon steenstrupi in June. Lithacrosiphon alticonus 

did not spawn even though oocytes in the body 

cavity were visible through the body wall of many 

specimens. A planktotrophic larval stage occurs 

in the development of Phascolosoma perlucens and 

P. varians. Although development of Paraspidosiphon steenstrupi has not been observed, the small 

size of the egg and the relatively sparse yolk 

indicate that development of this species is also 

planktotrophic. 

Specimens of Aspidosiphon brocki, known to reproduce asexually by constricting the posterior 

end to form a new individual (Rice, 1970), were 

examined for evidence of reproductive activity. 

Nineteen percent of the April 1974 collections 

had posterior constrictions. An additional eight 

percent had juveniles in their burrows. 

Discussion and Conclusions 

The rock-boring sipunculans at Carrie Bow 

Cay showed specific distribution within the reef 

community, the greatest diversity occurring at 

the reef crest where six species (of which four were 

common rock borers) were found. The four rock 

borers, found together only in the reef crest and 

high-relief spur and groove zones, each had distinctive distribution on the transect. Lithacrosiphon 

alticonus was found all along the transect in both 

deep and shallow waters but it was most abundant from the high-relief spur and groove zone to 

the outer ridge. Aspidosiphon brocki was dominant 

at the reef crest but occurred only sporadically 

and in small numbers in other areas. Paraspidosiphon steenstrupi was most abundant at the reef crest 

but occurred in small numbers along most of the 

transect. Phascolosoma perlucens was limited to the 

reef crest and to the shallow waters of the forereef zone. In being related to reef zonation, the 

overall distribution of these four sipunculan species must also be related directly or indirectly to 

physical parameters that control this zonation, 

the most important of which are water agitation 

and light intensity, both depth-dependent factors. 

The substratum most densely inhabited by sipunculans was generally fresh coral rock with 

almost uniform skeletal framework, as is characteristic of Porites and Acropora species, and with 

relatively little secondary calcite cement infill. 

The more highly eroded, water-worn, older rocks 

on the transect generally contained few, if any, 

sipunculans. Internally, such rocks showed considerable infilling of calcite cement and were 

heavily infiltrated by boring sponges. The presence of numerous tunnels and cavities suggested 

previous habitation by boring organisms such as 

sipunculans. The absence of extant sipunculans 

might be explained by natural death of specimens 

that had formed the borings, by natural succession of boring organisms, or by death due to a 

catastrophic event (for example, rocks having 

been buried in sediment). The lack of new boring 

by sipunculans might be due to unavailability of 

sufficient solid substratum in rocks that are heavily bored and infiltrated with sponges. 

Typically, the rocks containing sipunculans 

were covered by sponges and algal growths (filamentous, leafy and crustose coralline). Encrusting 

organisms such as bryozoans and the foramini-
NUMBER 12 319 

feran Homotrema sp. occurred most commonly but 

not exclusively on the underside of the rock samples. Sponges, along with a host of other invertebrates such as crabs, stomatopods, alpheid 

shrimp, isopods, barnacles, certain polychaetes, 

and bivalves, were found within the rocks—either 

in interstices and cavities or in burrows that they 

had formed in the rocks. Organisms in burrows 

included the boring sponges, polychaetes of the 

families Spionidae, Sabellidae, and Serpulidae as 

well as certain boring bivalves {Lithophaga sp. and 

Gastrochaena sp.) and boring barnacles {Lithotrya 

sp.). Sipunculans were rarely found in rocks with 

barren surfaces and were not observed in live 

portions of coral colonies. Sipunculan borings 

open most commonly on the upper surfaces and 

sides of rocks, but also on the lower surface lying 

on the sand. Boring sipunculans are known to 

feed on the surfaces of rocks they inhabit, ingesting bits of debris and sand entrapped in the 

surface cover and possibly pieces of algae (Rice, 

1969, 1975a); thus the epibiota are probably important to the survival of the animals within the 

rock, although details of this association remain 

to be investigated. 

The present study points out pertinent factors 

for future evaluation of the relative influence of 

various physical and biotic factors on distribution 

of rock-dwelling sipunculans in the coral community. The greater density of animals in recent 

unaltered coral rock from Carrie Bow Cay than 

in highly eroded rock indicates the probable importance of internal structure of the rock to infiltration by boring sipunculans. On the other hand, 

the observation that different species may inhabit 

the same type of coral rock in different areas of 

the reef community suggests that physical factors 

associated with locality—such as agitation and 

depth of water—may be even more significant 

than coral skeleton in determining specific distribution. These data, however, must be considered 

preliminary because the same rock types could 

not be collected in each reef locality. Moreover, 

the variation in density of sipunculans in similar 

rocks from the same locality indicates the complexity of the problem and the necessity for more 

rigidly controlled quantitative procedures. Further studies should compare the relative significance to rock-boring sipunculans of the type of 

coral skeleton and degree of secondary infilling. 

Tests of habitat preference could be made by 

transplanting rocks of known composition to different parts of the reef community and by examining these rocks over a period of years. 

Many other questions concerning colonization 

of coral rock by boring sipunculans remain to be 

explored. Factors affecting larval settlement are 

unknown, yet they undoubtedly play an important role in determining distribution of those 

species having planktotrophic larvae, such as 

Phascolosoma perlucens (Rice, 1975b). Distribution 

of species lacking planktotrophic larvae would be 

regulated by different factors. For example, an 

explanation for the localized concentrations of 

Aspidosiphon brocki might be found in the pattern 

of asexual reproduction of this species. Young, 

crawling juveniles, unable to traverse great distances, probably would colonize either the same 

rock as the parent or other rocks in the vicinity. 

Recent studies on infaunal communities of 

coral rock have suggested that surface area, porosity of rock, percentage of live coral and epibiota may influence infaunal distribution and colonization (Hutchings, 1978). Therefore, reliable 

quantitative data and improved methods of approach are obviously necessary. Brock and Brock 

(1977) proposed the use of acid dissolution of rock 

for quantitative removal of specimens previously 

preserved in formalin. Hutchings (1977) measured exposed surface area of substrata by coating 

surfaces with Playtex rubber which, after drying, 

can be peeled off, traced and measured with a 

polar planimeter. Various other techniques for 

measurements and sampling have been listed in 

a recent review of research methods for studies of 

coral reef cryptofauna (Hutchings, 1978). To 

date, most research efforts on specific groups of 

rock-dwelling fauna in coral reefs have focussed 

on polychaetes, although sipunculans have been 

recognized as a significant component of the 

coral-rock infauna (Kohn and Lloyd, 1973; Peyrot-Clausade, 1974; Hutchings, 1974, 1977). It is 

hoped that future studies will apply the more 

refined current techniques to the rock-boring si-
320 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

punculans and thus may provide a better understanding of their distribution in the reef community and of their significance to the total reef 

ecosystem. 

Literature Cited 

Brock, R. E., and J. H. Brock 

1977. A Method for Quantitatively Assessing the Infaunal Community in Coral Rock. Limnology and 

Oceanography, 22:948-951. 

Fischer, W. 

1922a. Westindische Gephyreen. Zoologischer Anzeiger, 55: 

10-18. 

1922b. Gephyreen des Reichmuseums zu Stockholm. Arkiv 

forZoologi, 15:1-39. 

Gardiner, J. S. 

1903. The Maldive and Laccadive Groups, with Notes 

on Other Coral Formations in the Indian Ocean. 

In J. S. Gardiner, editor, The Fauna and Geography 

of the Maldive and Laccadive Archipelagoes, 1:333-341. 

Cambridge: Cambridge University Press. 

Hutchings, P. A. 

1974. A Preliminary Report on the Density and Distribution of Invertebrates Living on Coral Reefs. In 

A. M. Cameron, B. M. Campbell, A. B. Cribb, R. 

Endean, J. S. Jell, O. A. Jones, P. Mather, and F. 

H. Talbot, editors, Proceedings of the Second International Coral Reef Symposium, 1:285-296. Brisbane, 

Australia: The Great Barrier Reef Committee. 

1977. Distribution and Abundance of Cryptofauna from 

Lizard Island, Great Barrier Reef. Marine Research 

in Indonesia, 19:99-112. 

1978. Non-Colonial Cryptofauna. In D. R. Stoddart and 

R. E. Johannes, editors, Coral Reefs: Research 

Methods. Monographs on Oceanographic Methodology, 

5:251-262. Paris: UNESCO. 

Kohn, A. J., and M. C. Lloyd 

1973. Polychaetes of Truncated Reef Limestone Substrates on Eastern Indian Ocean Coral Reefs: 

Diversity, Abundance, and Taxonomy. Internationale Revue der gesamlen Hydrobiologie und Hydrographic, 

58:369-399. 

Macintyre, I. G. 

1977. Distribution of Submarine Cements in a Modern 

Caribbean Fringing Reef. Journal of Sedimentary 

Petrology, 47:503-516. 

Marsh, J. A., Jr. 

1970. Primary Productivity of Reef-building Calcareous 

Red Algae. Ecology, 51:255-263. 

Murina, V. V. 

1967. [On the Sipunculid Fauna of the Littoral of Cuba.] 

Zoologicheskii Zhurnal, 46:35-47. [In Russian.] 

Otter, G. W. 

1937. Rock-destroying Organisms in Relation to Coral 

Reefs. In British Museum (Natural History) Great 

Barrier Reef Expedition 1928-29, 1:323-352. 

Peyrot-Clausade, M. 

1974. Ecological Study of Coral Reef Cryptobiotic Communities: An Analysis of the Polychaete Cryptofauna. In A. M. Cameron, B. M. Campbell, A. B. 

Cribb, R. Endean, J. S. Jell, O. A. Jones, P. 

Mather, and F. H. Talbot, editors, Proceedings of 

the Second International Coral Reef Symposium, 1:269-

283. Brisbane, Australia: The Great Barrier Reef 

Committee. 

Rice, M. E. 

1969. Possible Boring Structures of Sipunculids. American 

Zoologist, 9:803-812. 

1970. Asexual Reproduction in a Sipunculan Worm. 

Science, 167:1618-1620. 

1975a. Survey of the Sipuncula of the Coral and Beachrock Communities of the Caribbean Sea. In M. E. 

Rice and M. Todorovic, editors, Proceedings of the 

International Symposium on the Biology of the Sipuncula 

andEchiura, 1:35-49. Belgrade: Naucno Delo Press. 

1975b. Observations on the Development of Six Species 

of Caribbean Sipuncula with a Review of Development in the Phylum. In M. E. Rice and M. 

Todorovic, editors, Proceedings of the International 

Symposium on the Biology of Sipuncula and Echiura, 1: 

141-160. Belgrade: Naucno Delo Press. 

1976. Sipunculans Associated with Coral Communities. 

Micronesica, 12:119-132. 

Rice, M. E., and I. G Macintyre 

1972. A Preliminary Study of Sipunculan Burrows in 

Rock Thin-Sections. Caribbean Journal of Science, 12: 

41-44. 

Shipley, A. E. 

1903. Report on the Gephyrea Collected by Professor 

Herdman at Ceylon in 1902. In W. A. Herdman, 

Report to the Government of Ceylon on the Pearl Oyster 

Fisheries of the Gulf of Manaar, part 1, supplement 3, 

pages 169-176. 

Ten Broeke, A. 

1925. Westindische Sipunculiden und Echiuriden. Bijdragen tot de Dierkunde, 24:81-96. 
Anthuridea (Crustacea: Isopoda) 

of Carrie Bow Cay, Belize 

Brian Kensley 

ABSTRACT 

Fourteen species of anthurid and paranthurid 

isopods from Carrie Bow Cay, Belize, are considered. These include two new genera, Belizanthura 

and Minyanthura, and seven new species, Apanthura 

geminsula, Belizanthura imswe, Mesanthura fasciata, 

M. punctillata, M. reticulata, Minyanthura corallicola, 

and Paranthura caribbiensis. These species are relatively abundant in shallow water (1.5 m or less) 

habitats such as coral rubble and coarse sandy 

sediments, but scant in deeper (6-24 m) sediments. Few species were observed in the entire 

range from the intertidal zone to depths of 40 m. 

Male anthurids found in the plankton show morphological adaptations suggesting that they had 

left their substrates for reproductive rather than 

feeding purposes. 

Introduction 

During January-February 1978 the author carried out two weeks of collecting at Carrie Bow 

Cay, Belize, from a variety of habitats, ranging 

from very shallow water to depths of about 40 m. 

Seventeen species of anthurids and paranthurids 

were collected (three represented by single specimens, and not treated in this paper). This is the 

highest number of anthuridean species taken in 

any single locality, and probably indicates that 

many species await discovery upon more careful 

investigation of different substrate and habitat 

types. This high number of species, and in some 

Brian Kensley, Department of Invertebrate Zoology, National Museum 

of Natural History, Smithsonian Institution, Washington, D.C. 

20560. 

cases, high numbers of specimens, is a strong 

indication of the important role the group often 

plays in shallow-water ecosystems. Very little is 

known of the biology, expecially the feeding of 

these animals. It is hoped that future living specimen studies in the laboratory will provide such 

data. 

Material is deposited in the National Museum 

of Natural History, Smithsonian Institution. In 

the accompanying figures a scale (in millimeters) 

is given only for the entire animal in dorsal view. 

ACKNOWLEDGMENTS.—My thanks are due the 

following Smithsonian personnel: R. J. Larson 

for information and for specimens from horizontal plankton sampling, C. A. Child for help in 

collecting and sorting, J. N. Norris for algal identifications, A. C. Cohen for material collected, 

and F. D. Ferrari for making earlier Belize material available. I am very grateful to T. E. Bowman for reading the manuscript and for his many 

useful comments and criticisms. 

Species List 

Suborder ANTHURIDEA 

Family ANTHURIDAE 

Apanthura Stebbing 

Apanthura geminsula, new species 

Apanthura signata Menzies and Glynn 

Apanthuroides Menzies and Glynn 

Apanlhuroides millae Menzies and Glynn 

Belizanthura, new genus 

Belizanthura imswe, new species 

Mesanthura Barnard 

Mesanthura fasciata, new species 

Mesanthura paucidens Menzies and Glynn 

Mesanthura pulchra Barnard 

Mesanthura punctillata, new species 

321 
322 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 141.—Apanthura geminsula, new species, holotype, 9: a, complete specimen; b, antennule; 

c, antenna; d, mandible; e, maxilla;/, maxilliped; g, telson; h, pleopod 1; i, pereopod 1; j , 

pereopod 7; k, pleon in lateral view. 
NUMBER 12 323 

FIGURE 142.—Apanthura geminsula, new species, 6: a, cephalon; b, pereopod 1; c, mandible; d, 

pleopod 2. 

Mesanthura reticulata, new species 

Minyanthura, new genus 

Minyanthura corallicola, new species 

Pendanthura Menzies and Glynn 

Pendanthura tanaiformis Menzies and Glynn 

Family PARANTHURIDAE 

Accalathura Barnard 

Accalathura crenulata (Richardson) 

Paranlhura Bate and Westwood 

Paranthura caribbiensis, new species 

Paranthura infundibulata Richardson 

Suborder ANTHURIDEA 

Family ANTHURIDAE 

Apanthura, Stebbing 

Apanthura geminsula, new species 

FIGURES 141, 142 

DESCRIPTION OF FEMALE.—Bod y moderately indurate, semitranslucent when alive, lacking any 

pigmentation. Cephalon with low triangular 

rostrum, not extending beyond anterolateral 

corners. Eyes dorsolateral. Body proportions: 

C < 1 = 2>3< 4 = 5 = 6>7 . Pereonites 4, 5, 

and 6 each with middorsal shallow pit. Pleonites 

1-5 fused, but indicated ventrolaterally, with 

shallow dorsal grooves indicating lines of fusion; 

pleonite 6 free. Telson thin, hardly indurate, with 

hyaline border, widest at midlength, distodorsally 

slightly concave, with few distal setae; 2 large 

basal statocysts present. 

Antennula r peduncle 3-segmented, basal segment equal in length to 3 distal segments; flagellum of 3 articles. Antenna l peduncle 5-segmented, 

second segment grooved to accommodat e antennule; flagellum of 2 articles. Mandibula r palp 3-

segmented, terminal segment short, bearing 2 

setae, middle segment longest, with elongate distal seta; incisor of 2 blunt cusps; lamina dentat a 

with 4 marginal serrations; molar distally bluntly 

bilobed. Maxilla slender with one strong spine 
324 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 24.—Distribution and morphological comparison of four species of Apanthura 

Distribution 

Pigment 

Dorsal pits 

Antennular flagellum 

basal article $ 

Antennal flagellum 

articles 

Maxillipedal endite 

Uropodal endite 

Size? 

A. magnifica 

Georgia to Florida 

absent 

slight depression on 

pereonites 4-6 

elongate 

4 

present 

notched 

8.5 mm 

A. signata 

Puerto Rico; Belize 

present 

slight depression on 

pereonites 4-6 

short 

3 

present 

notched 

4.5 mm 

A. significa 

Venezuela 

absent 

slight depression on 

pereonites 4-6 

elongate 

3 

absent 

unnotched 

5.0 mm 

A. geminsula 

Belize 

absent 

marked pits on 

pereonites 4-6 

short 

2 

present 

notched 

8.1 mm 

and 5 smaller distally curved spines. Maxilliped 

5-segmented, terminal segment situated on outer 

distal angle of fourth segment, with 5 setae; third 

segment narrowed, less than half length of fourth 

segment; slender endite on inner surface with 

single terminal seta. Pereopod 1 unguis almost 

half length of dactylus; propodal palm with few 

simple spines, rounded hyaline lobe at about 

midpoint: carpus triangular, distally narrowed. 

Pereopods 2 and 3 considerably smaller than 

pereopod 1. Posterior pereopods with propodus 

having strong posterodistal spine and several 

small fringed scales on posterior margin; carpus 

small, triangular, underriding propodus, with 

strong posterodistal spine. Pleopod 1 exopod operculiform, distal margin bearing several plumose 

setae; endopod slightly shorter and about half 

width of exopod, with few distal plumose setae. 

Uropodal exopod distally notched, extending to 

distal end of basis; endopod extending beyond 

telsonic apex, distally rounded, bearing numerous 

simple setae. 

DESCRIPTION OF MALE.—Eyes considerably 

larger than those in female. Antennular flagellum 

of 8 or 9 articles, each bearing whorl of filiform 

aesthetascs. Mandibular palp as in female, but 

incisor, lamina dentata, and molar reduced to 

blunt nonsclerotised lobes. Pereopod 1 with hyaline process at about midpoint of propodal palm 

narrower than in female; numerous simple spines 

on medial margin; carpus with narrowly rounded 

hyaline posterodistal lobe. Pleopod 2 endopod 

with cylindrical, apically narrowly rounded copulatory stylet extending beyond apices of rami. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble, coarse sediments, 0.2-1.5 m; Twin Cays, 

under mangroves, 0.2 m. 

Holotype: 2 (TL 8.1 mm), Twin Cays, USNM 

171166. 

Allotype: 6 (TL 4.8 mm), Twin Cays, USNM 

171167. 

Paratypes: 38, 3$, Twin Cays, USNM 171168. 

Additional Material: 120$, 3 sub<5, 18, 50 juveniles. 

REMARKS.—Superficially the species of the genus Apanthura are all remarkably similar, but 

subtle differences are to be found in the mandibular palp spination-setation, telsonic shape, maxillipedal segment proportions, and in the structure of the first maxillipeds. Apanthura magnifica 

Menzies and Frankenberg (1966), A. signata, and 

A. significa Paul and Menzies (1971) have been 

recorded either from the Caribbean or from adjoining areas, and are all figured as having pleonites 1-3 free, and pleonites 4 and 5 dorsally fused. 

Clearing specimens of A. magnifica and A. geminsula 

in lactic acid and Chlorozol Black shows that 

pleonites 1-5 are dorsally fused, but that pleonites 

1-3 have a groove or fold over the dorsum which 

appears as an articulation. Careful examination 

of the type material of A. significa and A. signata 

shows a similar fusion. The main distinguishing 

features of the four species of Apanthura discussed 

here are summarized in Table 24. 
NUMBER 12 325 

ETYMOLOGY.—The specific name geminsula is a 

Latinized form of Twin Cays, the locality where 

the species was abundant. 

Apanthura signata Menzies and Glynn 

FIGURE 143 

Apanthura signata Menzies and Glynn, 1968:28, fig. 10.—Paul 

and Menzies, 1971:42. 

DESCRIPTION OF FEMALE.—Body hardly indurate. Body proportions: C< 1 = 2>3<4 < 

5 > 6 > 7. Pereonites 4, 5, and 6 with shallow 

middorsal depression. Pleonites 1-5 dorsally 

fused, laterally distinct; pleonite 6 free. Telson 

not indurate, gently convex dorsally with relatively broad hyaline border, widest at midpoint; 

with 2 proximal statocysts. 

Antennular peduncle 3-segmented, basal segment slightly shorter than 2 distal segments together; flagellum of 3 articles, second article relatively elongate. Antennal peduncle 5-segmented, 

flagellum of 3 articles. Mandibular palp 3-segmented, terminal segment with 3 terminal setae, 

first and second segments each with single elongate seta; incisor of 3 blunt cusps, lamina dentata 

with 5 marginal serrations, molar blunt. Maxilliped 5-segmented, with thin-walled endite on 

inner surface. Pereopod 1 unguis almost half the 

length of dactylus; propodal palm with rounded 

distal lobe and convex hyaline flange, few simple 

setae; carpus triangular, posterodistal angle produced, narrowly rounded. Posterior pereopods 

with triangular carpus bearing strong posterodistal spine. Pleopod 1 exopod operculiform, endopod more than half as wide and almost as long 

as exopod. Uropodal exopod notched, extending 

beyond basis. 

DESCRIPTION OF SUBMALE.—Eyes larger than in 

female, but not as large as in mature male. Antennular flagellum lacking filiform aesthetascs. 

DESCRIPTION OF MALE.—Cephalon with eyes 

much larger than in female. Antennular flagellum of 9 articles bearing filiform aesthetascs. 

Mandible with palp as in female, but incisor, 

lamina dentata, and molar somewhat reduced. 

Pereopod 1 with propodal palm having low 

rounded process at about midpoint, hyaline convex flange on inner surface shorter than in female, 

but with more setae than in female. Pleopod 2 

endopod with cylindrical, distally rounded copulatory stylet extending well beyond rami. 

COLOR NOTES.—Pigment pattern fairly regular, especially the dark band between eyes and 

narrow bands on anterior and posterior dorsal 

parts of pereonites, and on pleonite 6 in female. 

Pigmentation in male lacking regularity found in 

female; chromatophores scattered over dorsal and 

ventral surfaces. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble and coarse sediments, intertidal to 24 m: 

~ 1209, 2 subo\ 16, 25 juveniles. 

PREVIOUS RECORDS.—Puerto Rico. 

Apanthuroides Menzies and Glynn 

Apanthuroides millae Menzies and Glynn 

FIGURES 144, 145 

Apanthuroides millae Menzies and Glynn, 1968:30, fig. 11. 

DESCRIPTION OF FEMALE.—Integument hardly 

indurate, with numerous pits on cephalon, pereon, pleon, and telson, and fine scales (seen with 

difficulty). Body proportions: C < 1 > 2 = 

3 = 4 = 5>6>7<P . Cephalon with short triangular rostrum not extending as far as anterolateral corners. Eyes dorsolateral. Pereonite 7 very 

short, one-third length of 6. Pleonites 1-5 fused, 

with shallow grooves indicating lines of fusion; 

pleonite 6 fused with telson. Latter proximally 

broad, distal two-thirds narrowed, apically 

rounded, with strong middorsal longitudinal 

ridge. 

Antennular peduncle 3-segmented, basal segment broad, equal in length to 2 distal segments 

plus 2 basal flagellar article; flagellum of 4 articles, two distal articles each with single aesthetasc. 

Antennal peduncle 5-segmented, second segment 

grooved to accommodate antennule; flagellum of 

7 articles. Mandibular palp 3-segmented, middle 

segment broadest and longest, terminal segment 

with 3 distal finely fringed spines; incisor of 2 

blunt cusps, narrow lamina dentata with 6 or 7 
326 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 143.—Apanthura signata Menzies and Glynn: a, complete specimen, 9; b, cephalon, 8; c, 

antennule 9; d, mandible; e, maxilliped;/, pereopod 1, 9; g, pereopod 1, 8; h, pereopod 2, 8; i, 

telson. 
NUMBER 12 327 

FIGURE 144.—Apanthuroides millae Menzies and Glynn, holotype, 2: a, complete specimen; b, 

antennule; c, antenna; d, left mandible; e, right mandible;/, maxilla; g, pereopod I; h, pleopod 

1; i, maxilliped; ^, uropodal exopod; k, pereopod 6. 
328 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 145.—Apanthuroides millae Menzies and Glynn, 8: a, 

pleopod 2; b, cephalon; c, pereopod 1. 

serrations; molar of right mandible elongate, slender, finely ridged, absent on left mandible. Maxilla elongate, slender, apically with 3 or 4 broadened cusps, no spines distinguishable. Maxilliped 

5-segmented, terminal segment semicircular, setose, second segment elongate, with well-developed thin-walled endite on inner face. Pereopod 

1 not markedly larger than following pereopods; 

unguis one-third length of dactylus, with short 

basal spine; propodus not very broad, palm 

straight, with 3 spines. Posterior pereopods, propodus with 2 distal finely-fringed spines; carpus 

slightly underriding propodus, anterior margin 

somewhat shorter than posterior margin; latter 

with strong posterodistal spine. Pereopod 7 absent. Pleopod 1 exopod twice width and subequal 

in length to endopod, rami overlapping slightly, 

together forming opercular surface closing off 

branchial chamber. Uropodal exopod extending 

well beyond basis, twice as long as wide, distally 

rounded, margin entire, sparsely setose; endopod 

almost circular, sparsely setose. 

DESCRIPTION OF MALE.—Integument somewhat less pitted than in female. Eyes enlarged. 

Antennular flagellum with 6 or 7 articles bearing 

filiform aesthetascs. Propodus of pereopod 1 with 

thin convex flange along palm, latter with 5 finely 

fringed spines in distal half. Pleopod 2 endopod 

with cylindrical copulatory stylet, apically 

rounded. 

MATERIAL EXAMINED.—Carrie Bow Cay, coarse 

sediments, 6-24 m: 3? (TL 2.3-2.8 mm), \6 (TL 

2.8 mm), USNM 171154. 

REMARKS.—The type material of this species, 

consisting of two males from Puerto Rico, agrees 

well with the male of the present material. Menzies and Glynn (1968), in describing A. millae, 

neither figured nor mentioned the characteristically pitted integument, neither did they figure 

the unusual mandible. 

Several features of this species agree with Natalanthura foveolata Kensley, 1978a, described from 

the southwest Indian Ocean. These similarities 

include the pitted integument, the 5-segmented 

maxilliped with endite, the elongate mandible 

with a slender molar present only on one side, 

pelopod 1 with the exopod and endopod together 

forming an operculum over the branchial chamber, and pleonites 1-5 fused, pleonite 6 fused with 

the telson. (With regard to the latter feature, 

Kensley (1978a) described the pleon of N. foveolata 

as having pleonites 1-3 free, 4 and 5 fused, 6 

fused with the telson. Clearing of a specimen with 

lactic acid and Chlorozol Black has shown that 

pleonites 1-5 are completely fused, as in the 

present species). The aforementioned features, 

especially the unique mandibular structure, leave 

no doubt that Natalanthura should be regarded as 

a junior synonym oi Apanthuroides. 

Belizanthura, new genus 

DIAGNOSIS.—Eyes present. Antennal flagellum 

of 7 articles. Mandibular palp 3-segmented; incisor, lamina dentata, and molar reduced in 8, 

normal in 9. Maxilliped 7-segmented with endite 

in 9; 5-segmented and lacking endite in 8. Pereopods 1-3 similar, subchelate. Pereopods 4-7 

with triangular carpus underriding propodus. 

Pleopod 1 not operculiform. Pleonites 1-6 free. 

Telson lacking statocysts. 
NUMBER 12 329 

TABLE 25.—Comparison of anthurid genera possessing a seven-segmented maxilliped 

Character Neohyssura Ocsanthura Minyanthura Belizanthura 

Eyes 

8 mouthparts 

Carpus of pereopods 

4-7 

Pleopod 1 

Pleonites 

Telson 

Statocysts 

absent 

p 

triangular 

non-operculiform 

1-5 free, 6 fused 

with telson 

spiniform 

7> 

absent 

similar to $ 

rectangular 

non-operculiform 

1-6 free 

flattened 

2 

present 

similar to 2 

rectangular 

operculiform 

1-5 fused, 6 fused 

with telson 

flattened 

2 

present 

very reduced 

triangular 

non-operculiform 

1-6 free 

flattened 

absent 

TYPE-SPECIES.—Belizanthura imswe, new species. 

ETYMOLOGY.—The generic name is derived 

from the country of Belize, plus anthura, the suffix 

often used for anthurid genera. 

REMARKS.—A 7-segmented maxilliped as in the 

female of Belizanthura also occurs in Neohyssura 

Amar (1952), Ocsanthura Kensley (1978b), and 

Minyanthura, new genus. The features separating 

these genera are summarized in Table 25. 

Belizanthura imswe, new species 

FIGURES 146, 147 

DESCRIPTION OF FEMALE.—Body very slender, 

semi-transparent when alive, integument thin, 

not indurate. Body proportions: C > 1 < 2 = 

3 < 4 = 5>6>7 . Cephalon with low triangular 

rostrum, tiny dorsal eyes. Pleonites free, 1-5 subequal, 6 with notch in posterodorsal margin. Telson widest at midlength, tapering to rounded 

setose apex, lateral margins distally with 7 or 8 

shallow serrations; statocysts absent. 

Antennular peduncle 3-segmented, flagellum 

of 4 articles, terminal article tiny. Antennal peduncle 5-segmented; flagellum of 7 articles. Mandibular palp 3-segmented, middle segment twice 

length of first, third segment bearing 3 distal 

fringed setae; incisor of 3 cusps; lamina dentata 

with 4 serrations and several tiny spinules; molar 

bluntly rounded. Maxilla with one large and 4 

small distal spines. Maxilliped 7-segmented, terminal segment small, with 3 setae; third segment 

short; endite reaching to distal margin of fourth 

segment, thin-walled, with single terminal seta. 

Pereopods 1-3 similar, unguis half length of dactylus; propodal palm very slightly sinuous, with 

few scattered setae; carpus triangular, with posterodistal point. Posterior pereopods with unguis 

about one-quarter length of dactylus, propodus 

with 3 posterodistal sensory spines; carpus almost 

triangular, with short free anterior margin. Pleopod 1 not operculiform, subequal to following 

pleopods. Uropodal exopod pyriform, basally 

broad, outer margin with 4 or 5 serrations, distally 

rounded, setose; endopod longer than basis, extending slightly beyond telson, distally rounded, 

setose. 

DESCRIPTION OF MALE.—Antennular flagellum 

of 8 articles, each with whorl of filiform aesthetascs. Eyes enormously enlarged, almost meeting 

middorsally and midventrally, with just sufficient 

space between to accommodate mandibular 

palps; remainder of mandible reduced to short 

nonmasticatory segment. Maxillae absent. Maxilliped 5-segmented, only terminal segment bearing 3 setae; endite absent. Pereopod 1 unguis 

about one-third length of dactylus; propodal 

palm straight, armed with about 8 short, fringed 

spines; carpus triangular, with apical fringed 

spine and short, acute process. Pereopod 2 propodus slightly shorter than pereopod 1, palm 

armed with 3 strong ventrodistal sensory spines. 

Posterior pereopods as in female. Pleopod 1 basis 

with 2 slender retinaculae, exopod and endopod 

subequal in length, broadly rounded distally both 

bearing elongate plumose setae. Pleopod 2 basis 
330 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 146.—Belizanthura imswe, new species, holotype, 9: a, complete specimen; b, antenna; c, 

antennule; d, mandible; e, maxilla;/, maxilliped; g, pereopod 1; h, pleopod 1; i, uropod; ;', 

telson. 
NUMBER 12 331 

FIGURE 147.—Belizanthura imswe, new species, 8: a, cephalon; b, mandible; c, maxilliped; d, 

pereopod 1; e, pereopod 2; / pereopod 7; g, pleopod 2. 

with 3 retinaculae; endopod bearing stout, api- rubble and shallow coarse sediments, 0.1-0.3 m; 

cally rounded copulatory stylet extending well Twin Cays, in algal mat under mangroves, 0.1-

beyond ramus; exopod with distinct transverse 0.3 m. 

suture at midlength. Holotype: 9 (TL 3.4 mm), Twin Cays, USNM 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 171172. 
332 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Allotype: 6 (TL 2.5 mm), USNM 171173. 

Paratypes: 49 (TL 3.0-3.7 mm), 46 (TL 2.3 

mm-2.7 mm), Twin Cays, USNM 171174. 

Additional Material: 279, 28. 

ETYMOLOGY.—The specific name is the acronym for "Investigations of Marine Shallow Water 

Ecosystems." 

FIGURE 148.—Mesanthura fasciata new species, holotype, 9: a, complete specimen; b, antennule; 

c, antenna; d, mandible; e, maxilla;/, maxilliped; g, pleon in lateral view. 
NUMBER 12 333 

FIGURE 149.—Mesanthura fasciata new species, 6": a, pereopod 1; 6, pereopod 7; c, pleopod 1. 

Mesanthura, Barnard 

Mesanthura fasciata, new species 

FIGURES 148, 149 

DESCRIPTION OF FEMALE.—Integument not indurate. Body proportions: C < 1 < 2 = 3 < 

4 = 5 > 6 > 7. Cephalon with low rostrum, not 

extending beyond anterolateral corners; eyes dorsolateral. Pereonites 2 and 3 each with anterodorsal hollowed area. Pleonites 1-5 fused, indicated 

laterally, and with dorsal grooves marking fusion 

lines; pleonite 6 free, with middorsal notch in 

posterior margin. Telson dorsally slightly convex, 

distal margin rounded. 

Antennular peduncle 3-segmented, flagellum 

3-segmented, distal segment one-quarter length 

of second segment, with 3 aesthetascs. Antennal 

peduncle 5-segmented, second segment grooved 

to accommodate antennule; flagellum 3-articulate. Mandibular palp 3-segmented, third segment one-third length of second, with 4 finely 

fringed spines; incisor of 3 cusps; lamina dentata 

with 4 serrations; molar bluntly rounded. Maxilla 

with 1 strong spine and 5 smaller distally curved 

spines. Maxilliped 5-segmented, terminal segment small, set obliquely at outer distal angle of 

fourth segment; tiny thin-walled endite present 

at base of third segment. Pereopod 1 dactylus 

with small spine at base of unguis; propodal palm 

with rounded lobe at about midlength; 5 simple 

setae on inner face of propodus; carpus triangular, 

distally somewhat produced and rounded. Pereopods 2-7 similar; posterior pereopods with triangular carpus bearing short posterodistal spine, 

underriding propodus; latter with 2 serrate posterodistal spines; posterior margin bearing tiny 

fringed spines. Pleopod 1 exopod operculiform, 

extending somewhat beyond endopod; both rami 

with distal plumose setae; basis with 3 retinaculae. Uropodal exopod with strong distal notch, 

fringed with plumose setae, extending beyond 

base of endopod; latter oval, extending slightly 

beyond telsonic apex. 

COLOR NOTES.—Red-brown pigment pattern 

constant, characterized by irregular patch on 

cephalon, delicate reticulation on pereonites 1-3; 

thin anterior submedian lines and posterior solid 

transverse bars on pereonites 4-7; pleon with 5 

short transverse bars on fused pleonites 1-5, 
334 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

pleonite 6 with fine middorsal tracery; patches of 

pigment on telson and uropodal bases and endopods. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble and coarse sediments, intertidal to 24 m. 

Holotype: 9 (TL 4.5 mm), Carrie Bow Cay, 

USNM 171162. 

Paratypes: 69 (TL 3.7-4.5 mm), USNM 171163. 

Additional Material: 289, 25 juveniles. 

REMARKS.—On the basis of the color pattern, 

the present species bears some resemblance to M. 

occidentalis Menzies and Barnard, 1959, from 

southern California, especially in the delicate tracery on pereonites 1-3. The Californian species, 

however, lacks the strong posterior pigment bars 

on pereonites 4-7 and the 5 shorter bars on the 

pleon. The 2 species can further be separated on 

the form of the maxilliped. 

ETYMOLOGY.—The specific name, derived from 

the Latin fasciata (striped), refers to the transverse 

bands of pigment on the posterior pereonites and 

on the pleon. 

Mesanthura paucidens Menzies and Glynn 

FIGURES 150, 151 

Mesanthura paucidens Menzies and Glynn, 1968:27, fig. 9A-G. 

DESCRIPTION OF FEMALE.—Anterolateral lobes 

of cephalon rounded, not extending beyond rostrum. Cephalon with marked dorsolateral ridge; 

midventral margin posterior to maxilliped evenly 

convex. Pereon and pleon not indurate. Pleonites 

1-5 fused, sutures barely discernible along ventrolateral margin; pleonite 6 free, with narrow 

middorsal notch in posterior margin. Telson dorsally gently convex, distal margin evenly rounded. 

Antennular peduncle 3-segmented, distal segment very short, flagellum of 3 articles, terminal 

article bearing several setae and 2 aesthetascs. 

Antennal peduncle 5-segmented, second segment 

grooved to accommodate antennule; flagellum of 

3 (?4) short setose articles. Mandibular palp 3-

segmented, terminal segment shortest and narrowest, with 6 stout spines; incisor of 3 blunt 

cusps, linked to somewhat reduced and rounded 

molar by 6-serrate lamina dentata. Maxilla slender, with 6 terminal spines. Maxilliped 5-segmented, second segment with reduced endite on 

inner face. Pereopod 1 unguis half length of 

dactylus; propodal palm finely crenulate, with 

small convex transparent process at midpoint 

bearing simple setae. Pereopod 2 unguis one-third 

length of dactylus; propodus cylindrical, with 

strong serrate distoventral spine; carpus triangular. Pereopods 5-7 with carpus and propodus 

each with distoventral spine. Pleopod 1 exopod 

operculiform, about 3 times width of endopod; 

basis with 4 retinaculae. Uropodal exopod (Figure 150^) distally sinuous rather than notched. 

DESCRIPTION OF MALE.—Eyes larger than in 

female. Antennular flagellum of 7 articles, each 

with whorl of filiform aesthetascs. Pereopod 1 

with lobe at about midpoint of propodal palm, 

inner surface of propodus with many finely serrate spines. Pleopod 2 copulatory stylet cylindrical, extending beyond rami, distally spinulose, 

apically rounded and slightly sclerotised. 

COLOR NOTES.—Female with roughly rectangular dorsal patches of chromatophores on cephalon and pereonites. Pleon with 5 laterally 

linked transverse bars. Telson and uropodal endopod and basis with proximal chromatophores. 

Male with pigmentation of pereon heavier and 

less defined than in female, but with 5 pleonal 

bars as in female. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble and shallow sediments: 259 (TL 6.6 mm), 

16* (TL 6.4 mm), 9 juveniles. Twin Cays, under 

mangroves: 19. 

PREVIOUS RECORDS.—Puerto Rico. 

REMARKS.—The palm of pereopod 1 has a 

definite rounded process at midlength, but as this 

is transparent, it may have been overlooked and 

therefore not figured by Menzies and Glynn 

(1968). The presence of a small maxillipedal endite is unusual in Mesanthura, but may be related 

to the relatively small body size, and the almost 

interstitial habit. The fact that the holotype is 

only about one-third the length of the present 

mature male and females may account for the 

omissions in the original description. 
NUMBER 12 335 

w 

»••»•: 

y«' 

m»&s 

lex 

FIGURE 150.—Mesanthura paucidens Menzies and Glynn, 9: a, complete specimen; b, antenna; c, 

antennule; d, cephalon in lateral view; e, pleon in lateral view; / mandible; g, maxilla; h, 

maxilliped; z, pereopod 1;/, pereopod 2; k, pereopod 7; /, pleopod 1. 
336 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 151.—Mesanthura paucidens Menzies and Glynn, 8: a, pereopod 1; b, antennule; c, 

pleopod 2. 

Mesanthura pulchra Barnard 

FIGURES 152, 153 

Mesanthura pulchra Barnard, 1925:145, fig. 9e.—Schultz, 

1969:109, fig. 151. 

Mesanthura decorata Menzies and Glynn, 1968:26, fig. 8A-I. 

DESCRIPTION OF FEMALE.—Cephalon with rostrum reaching as far forward as anterolateral 

angles. Body proportions: C < 1 = 2 > 3 < 

4 = 5 = 6 > 7. Anterior 5 fused pleonites equal in 

length to pereonite 7; individual pleonites indicated by very short ventrolateral incisions; pleonite 6 free, with middorsal slit in posterior margin. 

Telson broadly rounded distally. 

Antennular peduncle 3-segmented, basal segment longest and broadest, flagellum of 3 articles. 

Antennal peduncle 5-segmented, second segment 

broadest, hollowed dorsally to accommodate antennule; flagellum of 3 short articles. Mandibular 

palp 3-segmented, middle segment longest, distal 

segment shortest, with row of 10 spines; incisor of 

4 rounded cusps, linked to blunt rounded molar 

by lamina dentata having 5 serrations. Maxilla 

slender, with 5 or 6 distal spines. Lower lip bilobed, ending in narrowly rounded process, with 

fine lateral setae. Maxilliped 5-segmented, second 

segment longest; third segment slightly narrower 

than fourth; latter with 4 short setae on median 

margin; distal segment triangular, with 2 stout, 

fringed setae and few simple setae. Pereopod 1 

unguis one-third length of dactylus; propodal 

palm with rounded process at about midpoint 

bearing row of 6 setae, remainder of propodus 

and carpus with few scattered setae. Pereopod 2 

unguis one-quarter length of dactylus; propodus 

with distoventral spine and few setae; carpus very 

short, triangular. Pereopods 5-7, propodus with 

distoventral simple spine plus fringed spine and 

several short spinules; carpus triangular, underriding propodus, with distoventral simple spine. 
NUMBER 12 337 

FIGURE 152.—Mesanthura pulchra Barnard, 9: a, complete specimen; b, antenna; c, antennule; d, 

pereopod 1; e, mandible;/ maxilla; g, maxilliped; h, lower lip; i, pleopod \;j, pereopod 2; k, 

pereopod 7; /, pleon in lateral view. 
338 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 153.—Mesanthura pulchra Barnard, 8: a, antennule; b, pereopod 1; c, pleopod 2. 

Pleopod 1 exopod operculiform, 3 times width of 

endopod, with numerous plumose setae on distal 

margin; endopod with 9 plumose setae on distal 

margin; basis with 6 retinaculae. Uropods typical 

of genus. 

DESCRIPTION OF MALE.—Antennule with 3-segmented peduncle, flagellum of 7 articles, each 

bearing whorl of filiform aesthetascs. Pereopod 1 

similar to female, but with unguis half length of 

dactylus, propodus with dense band of spines on 

inner face of paim. Pleopod 2 copulatory stylet of 

endopod simple, cylindrical, extending well beyond rami. 

COLOR PATTERN.—Broad band of chromatophores on cephalon posterior to eyes; hollow, 

roughly rectangular dorsal patch on pereonites 

1-6, pereonite 7 and pleon with pigment band 

laterally broad, narrowed middorsally. Scattering 

of chromatophores on telson and uropods. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble and shallow sediments: 129 (TL 9.3 mm), 

4 juveniles. 

PREVIOUS RECORDS.—St. Thomas and St. John, 

U.S. Virgin Islands; Puerto Rico; Dry Tortugas, 

Florida. 

TYPE MATERIAL.—Barnard's type series from 

the Copenhagen Museum consists of 28 and 1 

ovigerous 9 syntypes, St. Thomas and St. John, 

10-18 fathoms (18-33 m); ovigerous 9 lectotype 

(TL 6.5 mm); 2d (TL 5.4 mm, 4.7 mm) paralectotypes. 

REMARKS.—Although Barnard described this 

species in 1925, examination of the 3 type specimens still reveals the pigment pattern quite 

clearly. Menzies and Glynn (1968) based their 

species M. decorata on differences in the pigment 

patterns but made no comparison of appendages. 

Comparison of the Carrie Bow Cay material and 

that from the Dry Tortugas, Barnard's types, and 

Menzies and Glynn's types shows that despite 

slight differences (for example, in the presence or 

absence of a clear area on the cephalon), the basic 

pigment pattern is the same. Comparison of the 

appendages, especially the first pereopods in both 

the male and female shows no differences in the 

4 groups of specimens available. Schultz's figure 
NUMBER 12 339 

(1969, fig. 151) misrepresents the pigment pattern. 

Mesanthura punctillata, new species 

FIGURES 154, 155 

DESCRIPTION OF FEMALE.—Integument moderately indurate. Cephalon with large dorsolateral 

eyes, low triangular rostrum extending slightly 

beyond anterolateral corners. Body proportions: 

C < 1 = 2>3< 4 = 5>6>7 . Pleonites 1-5 

fused, only pleonite 1 indicated laterally; pleonite 

6 free, with middorsal slit in posterior margin. 

Telson broadly rounded distally. Antennular peduncle 3-segmented, flagellum of 3 articles. Antennal flagellum 3-articulate. Mandibular palp 

3-segmented, middle segment twice length of distal segment, latter armed with 7 finely fringed 

spines; incisor with 3 cusps; lamina dentata with 

5 blunt marginal serrations; molar acute, slightly 

sclerotised. Maxilla with 1 strong and 4 smaller 

spines. Maxilliped 5-segmented, distal segment 

semicircular, with 4 setae; penultimate segment 

with 2 medial distal setae. Pereopod 1 unguis half 

length of dactylus, with tiny spine at base; propodus proximally broad, palm with hyaline serrate lobe at midlength; carpus narrowly triangular, distal rounded part bearing 5 serrations. Posterior pereopods with short triangular carpus underriding propodus; posterior margin of propodus 

bearing short, fringed scales; posterodistal corner 

with 1 simple and one serrate spine. Pleopod 1 

exopod operculiform, about 3 times wider than 

endopod; both rami with distal plumose setae; 

basis with 4 retinaculae. Uropodal exopod with 

outer sinuous margin, but not notched; endopod 

almost circular. 

DESCRIPTION OF SUBMALE.—Antennular flagellum elongate and swollen, but lacking aesthetascs. 

Pereopod 1 as in female. Pleopod 2 not yet differentiated. 

DESCRIPTION OF MALE.—Eyes larger than in 

female, extending dorsally and ventrally. Antennule with flagellum of 10 articles bearing whorls 

of filiform aesthetascs. Mandible with palp as in 

female; incisor and lamina dentata reduced and 

not sclerotised; molar absent. Pereopod 1 as in 

female but with dense band of simple spines on 

inner face of propodus. Pleopod 2 endopod with 

copulatory stylet extending well beyond rami, 

apically rounded. 

COLOR NOTES.—Female with pigment pattern 

consisting of almost solid red-brown bar between 

eyes and extending in lobes posteriorly; pereon, 

pleon, telson, and uropods bearing scattered and 

separate pigment spots. (When chromatophores 

are expanded, pigment pattern is still scattered 

and does not become reticulate.) Male with chromatophores scattered dorsally and ventrally over 

entire body, denser than in female and with no 

discernible pattern. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble, coarse sediments, intertidal to 12 m. 

Holotype: Ovigerous 9 (TL 6.4 mm), Carrie Bow 

Cay, USNM 171157. 

Allotype: 8 (TL 4.5 mm), Carrie Bow Cay, 

USNM 171158. 

Paratypes: 8 (TL 4.5 mm), Carrie Bow Cay, 

USNM 171159; 39 (TL 5.7 mm, 5.2 mm, 5.2 

mm), Carrie Bow Cay, USNM 171160. 

Additional Specimens: 59, 2 sub<5, 8 juveniles. 

REMARKS.—The distinctive pigment pattern 

distinguishes this species from its congeners from 

the same area. Other differences may be seen in 

the number of mandibular palp spines, shape of 

the uropodal exopod, the degree of lateral indication of the fusion of pleonites 1-5, and the 

armature of the first pereopod. 

ETYMOLOGY.—The specific name punctillata, derived from the Latin word for small spots, refers 

to the overall scattered spots of pigment. 

Mesanthura reticulata, new species 

FIGURE 156 

DESCRIPTION OF FEMALE.—Integument hardly 

indurate. Body proportions: C = 1< 2 = 3 = 

4 < 5 > 6 > 7. Cephalon with dorsolateral eyes; 

tiny triangular rostrum. Pleonites 1-5 fused; 6 

free, with middorsal notch in posterior margin. 
340 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 154.—Mesanthura punctillata new species, holotype, 9: a, complete specimen; b, antenna; 

c, antennule; d, mandible; e, maxilliped;/ pereopod 1; g, pereopod 7; h, maxilla; i, pleopod 1. 
NUMBER 12 341 

FIGURE 155.—Mesanthura punctillata new species, 8: a, pereopod 1; b, mandible; c, pleon in lateral view; d, pleopod 2. 

Telson dorsally flattened, distal margin evenly 

rounded. 

Basal antennular peduncle segment broader 

than, but equal in length to, 2 distal segments; 

flagellum of 3 articles. Antennal flagellum of 3 

articles. Mandibular palp, distal segment with 6 

finely serrate spines; incisor with 3 acute cusps; 

lamina dentata with 5 marginal serrations; molar 

thumb-like, blunt. Maxilla with 1 strong and 4 

slender distal spines. Maxilliped 5-segmented, 

lacking endite. Pereopod 1 unguis about onethird length of dactylus, propodus broad, palm 

with hyaline toothed lobe at midlength; carpus 

triangular, distal rounded part with about 6 serrations. Posterior pereopods with propodus bearing 3 strong posterodistal sensory spines; several 

spinules on posterior margin; carpus with short 

anterior margin, underriding propodus, with single sensory spine at anterodistal corner. Pleopod 

1 exopod operculiform, 3 times width and subequal in length to endopod, both rami with distal 

plumose setae; basis with 5 retinaculae. Uropodal 

exopod ovate, outer margin sinuous, extending 

slightly beyond distal end of basis, fringed with 

plumose setae. 

COLOR NOTES.—Red-brown chromatophores 

form reticulate pattern on dorsal surface of cephalon, pereon, and pleon. Scattered chromatophores on telson and uropods. 

MATERIAL EXAMINED.—Carrie Bow Cay, coarse 

sediment, 24 m. 

Holotype: 9 (TL 6.1 mm), Carrie Bow Cay, 

USNM 171161. 

REMARKS.—Mesanthura reticulata somewhat resembles M. punctillata in having scattered chromatophores with a concentration of pigment on 

the cephalon between the eyes, as well as in the 

shape of the maxilliped and first pereopod of the 

female. Mesanthura punctillata, however, does not 

have a notched uropodal exopod, the third mandibular palp segment has 7 more slender spines, 

rather than the 6 found in M. reticulata, and the 

body proportions of the 2 species also differ. 

Although only 1 specimen of this species was 

collected, the pigment pattern is distinctive 

enough to warrant the formation of a new species. 

ETYMOLOGY.—The specific name derives from 

the reticulate dorsal pigment pattern. 

Minyanthura, new genus 

DIAGNOSIS.—Antennular flagellum of 1 article; 

antennal flagellum of 4 articles. Mandible lacking 

palp and molar process. Maxilliped 7-segmented, 

bearing endite. Pleonites 1-5 fused; pleonite 6 

fused with telson. Telson with 2 basal statocysts. 

Pleopod 1, exopod and endopod together forming 

operculum over branchial chamber. Carpus of 

pereopods 4-7, rectangular, not underriding propodus. 

TYPE-SPECIES.—Minyanthura corallicola, new species. 
342 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 156.—Mesanthura reticulata new species, holotype, 9: a, complete specimen; b, antenna; c, 

antennule; d, mandible; e, maxilliped;/ pereopod 1; g, maxilla; h, pereopod 7; i, uropodal 

exopod; j , pleopod 1. 
NUMBER 12 343 

ETYMOLOGY.—The generic name is derived 

from the Greek minus (tiny), and anthura, the suffix 

used for many anthurid genera. 

REMARKS.—Three anthurid genera, Belizanthura, Ocsanthura, and Neohyssura, possess a 7-segmented maxilliped with an endite. All 3 genera, 

however, possess nonoperculiform first pleopods 

and pleonites 1-5 free, and Ocsanthura and Neohyssura have a short triangular carpus on the 

posterior 3 pairs of pereopods, which underrides 

the propodus. The present specimens (assigned 

herein to Minyanthura corallicola), with pleonites 1-

5 fused, and pleonite 6 fused with the telson, and 

operculiform pleopod 1, and a rectangular carpus 

on the posterior pereopods, obviously cannot be 

members of any of the above-mentioned genera. 

Table 25 summarizes these differences. 

Minyanthura corallicola, new species 

FIGURES 157, 158 

DESCRIPTION OF FEMALE.—Integument not indurate. Body proportions: C > 1 < 2 < 3 < 

4 = 5 > 6 > 7. Cephalon with broadly rounded 

rostrum extending beyond anterolateral corners; 

eyes dorsolateral. Pereonite 7 very short. Pleon 

only slightly longer than pereonite 7; pleonites 1-

5 fused, only indicated ventrolaterally; posterior 

margin of pleonite 5 with row of plumose setae; 

pleonite 6 fused with telson. Telson broad, distal 

margin crenulate, broadly rounded or truncate, 

with few setae; gently convex, longitudinal middorsal ridge present; broad hyaline margin; 2 

large basal statocysts. 

Antennular peduncle 3-segmented, basal segment as long as rest of appendage, segments 3 

and 4 equal in length, and half width of second 

segment; flagellum reduced to single very short 

setose article. Antennal peduncle 5-segmented, 

second segment curved ventrally, subequal in 

length to segments 3 and 4; flagellum of 4 articles, 

basal article longer than 3 distal articles together. 

Mandible with palp represented by single simple 

seta; incisor of 3 cusps, lamina dentata narrow, 

with 6 marginal serrations; molar absent. Maxilla 

with single stout spine and 5 shorter hooked 

spines. Maxilliped 7-segmented, 5 distal segments 

together shorter than second segment; thinwalled endite tipped with single seta, which 

reaches base of terminal palp segment. Pereopod 

1 unguis one-third length of dactylus; propodus 

with straight unarmed palm, with single stout 

serrate spine and irregular row of fine combs of 

setules on inner face. Pereopod 2 similar to pereopod 1. Posterior pereopods propodus with 2 

distal serrate spines; ventral margin with row of 

setule-combs; carpus rectangular, not underriding propodus. Pleopod 1 exopod and endopod 

lying side by side, subequal in length, together 

forming operculum over branchial chamber; exopod broadening distally, almost 3 times wider 

than endopod; both rami with distal plumose 

setae. Uropodal basis with row of plumose setae 

on outer margin: exopod widening distally, outer 

distal angle produced into acutely triangular 

lobe; inner distal angle rounded and dentate; 

endopod oval, distal margin serrate, endopod and 

exopod with broad hyaline border. 

DESCRIPTION OF MALE.—Body proportions as 

in female. Eyes larger than in female. Antennular 

peduncle 3-segmented, flagellum of 2 articles with 

single terminal aesthetasc. Pereopod 1 as in female. Pleopod 2 endopod with copulatory stylet 

formed by club-shaped extension of distal end of 

ramus. 

COLOR NOTES.—Female, cephalon with broad 

dark-red-brown band between and posterior to 

eyes, with lateral unpigmented spots, centrally 

continuous with pattern on pereonite 1; latter 

with anterodorsal branching patch; pereonite 2 

with dark medio-dorsal ramifying pattern; pereonite 3 with 2 slender lateral bars; pereonite 4 

with broad posterodorsal rectangle; pereonite 5 

with slender posterodorsal rectangle; tiny wedge 

between pereonites 6 and 7; pleon with solid 

middorsal patch with lateral extensions. Male, 

pigment pattern less defined than in female, with 

scattered ventral patches. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble, 6-24 m. 

Holotype: larvigerous 9 (TL 1.7 mm), Carrie 

Bow Cay, USNM 171169 (2 larvae in brood 

pouch, with pigment pattern developed). 

Allotype: 6 (TL 1.3 mm), Carrie Bow Cay, 

USNM 171170. 
344 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 157.—Minyanthura corallicola new species, holotype, 9: a, complete specimen; b, antenna; 

c, antennule; d, lateral margin of anterior pleon; e, mandible; / maxilla; g, maxilliped; h, 

pereopod 1; i, pereopod 6. 
NUMBER 12 345 

FIGURE 158.—Minyanthura corallicola new species, 8: a, pleopod 2; b, pleopod 1; c, antennule; d, 

telson and uropod. 

Paratypes: 49 (TL 1.5-1.8 mm), Carrie Bow 

Cay, USNM 171171. 

ETYMOLOGY.—The specific name corallicola 

which means coral-dwelling, is used because all 

the specimens were removed from coral rubble. 

Pendanthura, Menzies and Glynn 

Pendanthura tanaiformis Menzies and Glynn 

FIGURES 159, 160 

Pendanthura tanaiformis Menzies and Glynn, 1968:32, fig. 

12A-I. 

DESCRIPTION OF FEMALE.—Integument moderately indurate. Cephalon half length of pereonite 

1, with rostrum extending beyond anterolateral 

corners; eyes present in bases of anterolateral 

lobes. Body proportions: C<1> 2 = 3< 4 = 

5 > 6 > 7 > P. Brood pouch formed by 3 pairs 

oostegites on pereonites 3-5. Pleon very reduced, 

one-third length of pereonite 7; pleonites indicated only on ventrolateral margins by very short 

sutures. Telson dorsally flattened, distal margin 

broadly rounded, pair of statocysts situated at 

about midlength. 

Antennular peduncle 3-segmented, basal segment longest and broadest, distal segment with 

reduced flagellum of 2 short articles, plus setabearing papilla; peduncle armed with large pinnate setae, each articulating on distinct slightly 
346 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 159.—Pendanthura tanaiformis Menzies and Glynn, 9: a, complete specimen; b, antenna; 

c, antennule; d, maxilla; e, mandible;/ maxilliped; g, pleopod 1; h, pereopod 1; i, pereopod 7. 
NUMBER 12 347 

FIGURE 160.—Pendanthura tanaiformis Menzies and Glynn, 8: a, antennule; b, pereopod 1; c, 

pleopod 2 endopod. 

broader base. Antennal peduncle 5-segmented, 

segments with minute setules; flagellum reduced 

to single short article. Mandibular palp reduced 

to small papilla bearing single seta; incisor of 3 

rounded, slightly sclerotised cusps; molar reduced; lamina dentata plate with 7 marginal 

serrations. Maxilla slender, with 8 distal curved 

spines, terminal spine strongest. Maxilliped 3-segmented, terminal segment with several distal simple setae; thin-walled endite present on inner 

face, with 2 fine distal setae. Pereopod 1 unguis 

almost half length of dactylus, ventral margin of 

latter with fine spinules, strong supplementary 

spine at base of unguis; propodus with 6 serrate 

spines on inner face, rounded, very thin transparent lobe on palm with few simple setae. Pereopods 

2-7 similar, propodus with posterior margin with 

crenulations bearing clusters of short spinules and 

with strong serrate posterodistal spine; carpus 

short, triangular, underriding propodus; propodus, carpus, merus, and ischium bearing numerous short fine setules. Pleopod 1 exopod operculiform, with numerous laterodistal plumose setae; 

endopod half width of exopod, with 6 distal 

plumose setae; basis with 2 retinaculae. Uropodal 

exopod distally narrowly rounded, outer (dorsal) 

margin crenate, with numerous plumose setae, 

reaching slightly beyond basis; endopod ovoid, 

bearing plumose and elongate simple setae. 

DESCRIPTION OF MALE.—Antennular peduncle 

3-segmented, flagellum of 4 articles, each with 

cluster of aesthetascs. Pereopod 1 propodus with 

dense cluster of curved serrate spines on inner 

surface, palm with rounded transparent lobe at 

about midlength. Copulatory stylet of pleopod 2 

endopod attached at about proximal third, cylindrical, longer than ramus, apically narrowly 

rounded, with minute scattered spinules. 

COLOR NOTES.—Strong reticulate red-brown 

pigmentation dorsally on cephalon, pereonites, 

antennae and first pereopods, becoming diffuse 

on pleon, telson and uropods. The young are 

released from the brood pouch fully pigmented. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble: 37 ovigerous 9, 829, 456*, 4 juveniles. 

PREVIOUS RECORDS.—Puerto Rico. 
348 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

REMARKS.—This is the second record of this 

unusual genus and species, and the first record of 

the male. It was thought useful to supplement 

Menzies and Glynn's description and to figure 

most of the appendages. A few features require 

further comment. Menzies and Glynn (1968) described the mandibular palp as being reduced to 

2 setae. In fact, the palp may be regarded as 1-

segmented, this seta-bearing segment being reduced to a small papilla. Menzies and Glynn 

mentioned the possible existence of a second flagellar ramus of a single article, on the antennule. 

More likely, this apparent segment is the slightly 

enlarged articulated base of one of the large 

specialized pinnate setae. 

Family PARANTHURIDAE 

Accalathura, Barnard 

Accalathura crenulata (Richardson) 

Calathura crenulata Richardson, 1901:509, figs. 1-4; 1905:74, 

figs. 58-61. 

Accalathura crenulata.—Barnard, 1925:147, pl. 4: fig. 18.— 

Nierstrasz, 1941:242.—Menzies and Glynn, 1968:33, fig. 

13A-H.—Schultz, 1969:96, fig. 128. 

MATERIAL EXAMINED.—Carrie Bow Cay, intertidal to 12 m, sediments, 99, 38, 9 juveniles; Twin 

Cays, under mangroves, 19. 

PREVIOUS RECORDS.—Bahamas; Puerto Rico; 

Yucatan, 40 m; Brazil; Cape Verde Is. 

Paranthura, Bate and Westwood 

Paranthura caribbiensis, new species 

FIGURES 161, 162 

DESCRIPTION OF FEMALE.—Integument not indurate, with sparse scattered chromatophores. 

Cephalon with patch of chromatophores between 

dorsolateral eyes; tiny, triangular rostrum, not 

extending beyond anterolateral corners. Body 

proportions: C<1<2>3<4>5>6>7 . 

Pleonites 1-5 free, subequal; pleonite 6 with bilobed posterodorsal margin. Telson dorsally flat, 

basally constricted, distal margin evenly rounded, 

with few setae. 

Antennular peduncle 3-segmented, basal segment wider than and subequal in length to 2 

distal segments; flagellum of 4 articles bearing 

aesthetascs. Antennal flagellum of single flattened 

triangular article. Mandibular palp 3-segmented, 

second segment two and one-half times length of 

basal segment, third segment with 4 stout fringed 

spines. Maxilla slender, with 8 distal serrations. 

Maxilliped 3-segmented, with short endite at base 

of third segment. Pereopod 1 unguis about onethird length of dactylus; propodus proximally 

broad, with convex flange and row of 12-14 

spines on inner face. Pereopods 2 and 3 similar, 

subchelate, smaller than pereopod 1; propodal 

palm armed with 5 sensory spines. Pereopods 4-

7 ambulatory, propodus with 2 posterior sensory 

spines; carpus rectangular, with 2 posterior sensory spines. Pleopod 1 exopod operculiform, four 

times wider and slightly longer than endopod, 

both rami with distal plumose setae; basis with 4 

retinaculae. Uropodal endopod subcircular; exopod oval, outer margin slightly sinuous, extending 

beyond endopod base. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble, shallow sediments, and under mangroves. 

Holotype: Larvigerous 9 (TL 4.5 mm), Carrie 

Bow Cay, USNM 171164 (8 larvae in brood 

pouch). 

Paratype: Ovigerous 9 (TL 4.5 mm), Carrie Bow 

Cay, USNM 171165. 

Additional Material: 109, 4 juveniles. 

REMARKS.—This small species of Paranthura is 

easily distinguished from the larger P. infundibulata, which also occurs at Carrie Bow Cay, by the 

serrate and curved telson in the latter species. 

Paranthura barnardi Paul and Menzies, 1971, recorded from Venezuela, is more similar to P. 

caribbiensis in both size and structure. These 2 

species can be separated on the uropodal structure (broadly ovate and crenulate in P. barnardi, 

narrowly ovate/sinuous, and entire in P. caribbiensis), the telson, which is distally more rounded in 

P. caribbiensis, the presence of a small maxillipedal 

endite in this species (absent in P. barnardi), the 
NUMBER 12 349 

I /. 

FIGURE 161.—Paranthura caribbiensis new species, holotype, 9: a, complete specimen; b, antennule; 

c, antenna; d, mandible; <?, maxilla;/ maxilliped; g, telson; h, pleopod 1; i, uropodal exopod. 
350 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 162.—Paranthura caribbiensis new species, 9: a, pereopod 1; b, pereopod 2; c, pereopod 7. 

relative proportions of the antennular flagellar 

articles, and the number of spines on the terminal 

segment of the mandibular palp (5 in P. caribbiensis, 8 in P. barnardi). The patch of chromatophores 

between the eyes in the present species is also a 

distinctive feature. 

ETYMOLOGY.—The specific name is derived 

from the Caribbean Sea in which Carrie Bow 

Cay is situated. 

Paranthura infundibulata Richardson 

Paranthura infundibulata Richardson, 1902:284, figs. 15-20; 

1905:76, fig. 62.—Barnard, 1925:155.—Menzies and 

Glynn, 1968:1 l.—Schultz, 1969:95, fig. 126.—Camp, 

Whiting, and Martin, 1977:16. 

Paranthura verrillii Richardson, 1902:286, figs. 21, 22; 1905: 

77, fig. 63.—Barnard, 1925:155. Menzies and Glynn, 

1968:11. 

MATERIAL EXAMINED.—Carrie Bow Cay, coral 

rubble: 29, 16\ 12 juveniles. 

PREVIOUS RECORDS.—Bermuda; Florida. 

Discussion 

MICROHABITATS.—The five microhabitats that 

were sampled for anthuridean isopods in and 

around Carrie Bow Cay ranged in depth from 

the intertidal zone to about 24 meters. A general 

description of the location is presented in Riitzler 

and Macintyre (herein: 9). Although quantitative sampling was not done, about 5 liters of 

sediment, plant material, and/or rubble was collected at each station, and subjective measures of 

anthuridean abundance (Table 26) are given in 

addition to the comments on these microhabitats 

and their fauna that follow. 

1. Coral rubble and algae atop the reef, east 

side of island. Because much of the reef top is in 

very shallow water (intertidal to 30 cm) this area 

is subject to considerable wave action and turbulence under windy or stormy conditions, and 

to local warming under calm conditions. This 

environment—with the numerous crevices provided by the Corallinacea-encrusted pieces of 

coral—shelters a rich population of cryptic ani-
NUMBER 12 351 

TABLE 26.—Distribution of anthuridean species at Carrie Bow Cay and vicinity by substrate 

and depth (A = abundant, > 20 specimens; FC = fairly common, 5-19 specimens; P = present, 

< 5 specimens) 

Species 

Accalathura crenulata 

Apanthura geminsula 

A. signata 

Apanthuroides millae 

Belizanthura imswe 

Mesanthura fasciata 

M. paucidens 

M. pulchra 

M. punctillata 

M. reticulata 

Minyanthura corallicola 

Paranthura caribbiensis 

P. infundibulata 

Pendanthura tanaiformis 

Coral 

rubble 

0-1.5 m 

A 

P 

P 

A 

FC 

FC 

FC 

FC 

FC 

A 

Coarse 

sediments 

0-1.5 m 

FC 

A 

A 

P 

A 

A 

P 

P 

P 

Algal mat 

under 

0 

mangroves 

-1.5 

P 

A 

-

-

A 

-

P 

-

-

-

-

P 

-

-

m 

Coarse 

sediments 

6-12 m 

P 

P 

P 

-

P 

-

P 

-

-

-

Coarse 

sediments 

24 m 

_ 

-

P 

P 

P 

P 

P 

-

-

-

mals, including isopods, amphipods, small mollusks, polychaetes, sipunculans, and pycnogonids. 

Ten species of anthurideans were collected here, 

several of which also occurred in coarse sediments 

of weed beds and at the bases of patch corals. 

Pendanthura tanaiformis and Paranthura infundibulata 

were found only in the coral rubble, the former 

in great numbers. The dark wine-red pigmentation of both species, which live in the tiny holes 

in the Corallinacea-encrusted coral fragments, 

probably has a protective function. Probably also 

connected with this overriding red coloration of 

the coral rubble is the fact that 8 of the 10 species 

occurring here have some degree of integumental 

pigmentation. 

2. Coarse white sandy sediments in shallow 

water. These sediments were taken from Thalassia 

weed beds, Syringodium weed beds, or from the 

sandy patches between patch reefs at a depth of 

1.5 m or less. These sites are in protected areas 

with little wave action, either in the lagoonal area 

west of the island, or in the shallows between the 

main reef and the island. Eight species of anthurideans were common to these habitats and to the 

coral rubble area atop the reef, with Mesanthura 

fasciata, M. paucidens, and Apanthura geminsula being 

fairly common to abundant in both habitats. 

3. Shallow water sediments and algae at bases 

of mangroves. This habitat is characterized by 

calm water in the shade of the mangrove trees, 

with high accumulations of organic debris, especially mangrove leaves and rootlets, and Thalassia 

leaves. In the several large samples taken under 

the mangroves, three of the five species of anthurideans collected occurred only rarely, 

whereas Belizanthura imswe and Apanthura geminsula 

were abundant, especially in the dense carpet of 

Caulerpa verticillata J. Agardh growing in less than 

15 cm of water between the mangrove trunks and 

roots. Although Belizanthura was taken rarely from 

other living plants, it is abundant only in the 

Caulerpa verticillata. This algal mat, with its high 

organic debris content, and with a complex web 

of fine algal species (including Cladophora sp., 

Centroceras sp., Ceramium sp., blue-green algae and 

diatoms), which floats above the Caulerpa at hightide and sinks onto the Caulerpa at low-tide, harbors a rich fauna, including many asellote and 

gnathiid isopods, amphipods, a high diversity of 

pycnogonids (C. A. Child, pers. comm.), poly-
352 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

chaetes, nematodes, cumaceans, leptostracans, 

and small mollusks, especially the bivalved gastropod Berthelinia. 

4. Coarse sediments and rubble from the channel between Carrie Bow Cay and South Water 

Cay. This area is somewhat protected from wave 

action, but subject to the scouring action of water 

moving through the channel at a depth of 6 to 12 

m. Five species of anthurideans were collected 

here, none of which appeared to be abundant. 

5. Sand trough at base of coral and gorgoniancovered slope. Only two species, Apanthura signata 

and Mesanthura fasciata were collected from this 

area (~24 m depth) and from the intertidal zone. 

Both were abundant only in the very shallow 

sediments. Interestingly, Minyanthura corallicola resembles Pendanthura tanaiformis both in overall size 

and in having a very short anterior pleonal area. 

Both species seem to live in crevices in coral 

rubble, for which a reduction of the body length 

may be an advantage. 

GENERAL OBSERVATIONS.—Only male anthurids were taken in the horizontal plankton samples, 

in shallow water over the reefs suggesting that the 

males leave the substrate in search of females. 

The development of enlarged eyes, filiform aesthetascs of the antennular flagella, and pigmentation over the entire body (rather than confined to 

the dorsum in female Mesanthura spp., for example) are probably adaptations for this reproduction-centered activity. It is unlikely that feeding 

is a reason for this increased activity, considering 

the reduction of mouthparts seen in the males of 

Belizanthura imswe, Apanthura geminsula, and Mesanthura punctillata. 

The genus Mesanthura, represented here by five 

species (plus single specimens of two additional 

species not dealt with in this report) shows very 

successful radiation into several of the microhabitats mentioned above. Four of these five species 

co-occur in the coral rubble environments, as well 

as in the coarse sediments from shallow water. 

Closer investigation will probably reveal distinctive feeding preferences, and even behavioral differences (considering the distinctive and constant 

dorsal pigmentation of each species), which 

would account for this apparent overlap of several 

species of the same genus. 

Literature Cited 

Amar, R. 

1952. Isopodes marins du littoral corse. Bulletin de Societe 

Zoologique de France, 77:349-355. 

Barnard, K. H. 

1925. A Revision of the Family Anthuridae (Crustacea 

Isopoda), with Remarks on Certain Morphological Peculiarities. Journal of the Linnaean Society of 

London, Zoology, 36:109-160. 

Camp, D. K., N. H. Whiting, and R. E. Martin 

1977. Nearshore Marine Ecology at Hutchinson Island, 

Florida, 1971-1974, V: Arthropods. Florida Marine 

Research Publications, 25:1-63. 

Kensley, B. K. 

1978a. The South African Museum's Meinng Naude 

Cruises: The Isopoda Anthuridea from the 1975, 

1976, and 1977 Cruises. Annals of the South African 

Museum, 77(1): 1-25. 

1978b. A New Genus and Species of Anthurid Isopod 

from Deep Water off the East Coast of the United 

States. Proceedings of the Biological Society of Washington, 91:558-562. 

Menzies, R. J., and J. L. Barnard 

1959. Marine Isopoda of Coastal Shelf Bottoms of 

Southern California: Systematics and Ecology. 

Pacific Naturalist, 1 (11 -12): 1 -35. 

Menzies, R. J., and D. Frankenberg 

1966. Handbook of the Common Marine Isopod Crustacea of 

Georgia. 93 pages. Athens, Georgia: University of 

Georgia Press. 

Menzies, R. J., and P. W. Glynn 

1968. The Common Marine Isopod Crustacea of Puerto 

Rico. Studies on the Fauna of Curacao and Other Caribbean Islands, 27:1-133. 

Nierstrasz, H. F. 

1941. Gnathiidea, Anthuridea, Valvifera, Asellota, 

Phreatocoidea. In Die Isopoden der Siboga-Expedition, IV: Isopoda Genuina. Siboga-Expeditie, Vitkomslen op Zoologisch, Botanisch, Oceanographisch en 
NUMBER 12 353 

Geologisch Gebied . . . uitgegeven door . . . M. Weber, 

Monographic, 32(d): 1-308. 

Paul, A. Z., and R. J. Menzies 

1971. Sub-Tidal Isopods of the Fosa de Cariaco, Venezuela, with Descriptions of Two New Genera and 

Twelve New Species. Boletin de Instituto Universidade 

Oriente, 10:29-48. 

Richardson, H. 

1901. Key to the Isopods of the Atlantic Coast of North 

America with Descriptions of New and Little 

Known Species. Proceedings of the United Stales National Museum, 23:493-579. 

1902. The Marine and Terrestrial Isopods of the Bermudas, with Descriptions of New Genera and 

Species. Transactions of the Connecticut Academy of 

Sciences, 11:277-310. 

1905. A Monograph of the Isopods of North America. 

Bulletin of the United States National Museum, 54:1— 

727. 

Schultz, G A. 

1969. How to Know the Marine Isopod Crustaceans, i-vii + 

359 pages. Dubuque, Iowa: W. C. Brown. 
Pycnogonida from Carrie Bow Cay, Belize 

C. Allan Child 

ABSTRACT 

This is a systematic account of pycnogonids 

found in the area of Carrie Bow Cay, Belize. 

Thirty one identified species in 14 genera are 

discussed of which four are described as new 

species: Hedgpethius mamillatus, Callipallene belizae, 

Anoplodactylus imswe, and Rhynchothorax crenatus. 

Most species discussed were found within their 

previously known range of geographical distribution. Three species are reported for only the 

second time: Hedgpethius tridentatus, previously 

known from Florida, Parapallene bermudensis, from 

Bermuda, and Anoplodactylus bahamensis, from the 

Bahamas. At Carrie Bow Cay most pycnogonids 

occur within the littoral depth zone, some extend 

their range into the sublittoral to a depth of 33 

m. Habitats include mangrove roots, algae, and 

seagrasses, as well as coral and coral rubble. 

Introduction 

This is the first report of pycnogonids from the 

Belizean barrier reef. Only a small collection from 

the northern portion of this barrier reef in the 

Territorio de Quintana Roo, Mexico, has been 

treated in a previous publication (Child, 1979). 

Previous to that report, the nearest occurrence of 

pycnogonids reported in the literature were deep 

water captures of Anoplodactylus lentus Wilson and 

Ascorhynchus serratus Hedgpeth off Yucatan (Hedgpeth, 1948:226, 259). Anoplodactylus lentus is also 

listed from deep water in the Yucatan Channel 

(Stock, 1975:1055). Other nearby captures are 

recorded from Cuba to the northeast and from 

the Caribbean coast of Panama to the south. 

C. Allan Child, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. 

Most published work on pycnogonids centers 

on descriptive systematics. The small size, cryptic 

coloration, and very slow movement of these 

predators make it difficult to record their habits 

and habitats in the field. Therefore, our knowledge of the ecology of this group is rather poor. 

Although no attempt was made in this study to 

observe pycnogonids alive, ecologically relevant 

information on substrates and associations was 

obtained and is presented. 

All specimens are deposited in the pycnogonid 

collections of the National Museum of Natural 

History, Smithsonian Institution, and bear the 

catalog numbers of the United States National 

Museum collection (USNM). 

ACKNOWLEDGMENTS.—My appreciation is extended to the various collectors listed under "material examined" and to P. M. Kier for supporting 

my 1976 field work in Belize. I am grateful to T. 

E. Bowman for his critical reading of the manuscript. 

Methods 

Pygnogonids were found in a wide variety of 

microhabitats such as bryozoans, hydroids, and 

encrusting sponges, algae, or seagrass, and rock 

rubble covered with one or more of these. Most 

samples were collected in a depth of less than one 

meter, some to a maximum depth of 33 m. The 

gross samples were either examined live under a 

low-power microscope or preserved before study. 

Plants and broken-up rock and rubble were agitated in a bucket with dilute formalin in seawater 

to separate and preserve the microfauna. Floating 

organisms were then skimmed off and the remainder of the liquid poured through a net to 

355 
356 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

concentrate the sample. Other suitable substrates 

such as dock pilings, coral heads, mangrove roots, 

and rock faces were scraped or chipped into 

plastic bags for preservation and later examination. 

All specimens illustrated in this report were 

drawn by the author with the aid of a compound 

microscope and a camera lucida. 

Species List 

Family AMMOTHEIDAE 

Achelia Hodge 

Achelia sawayai Marcus 

Ammothella Verrill 

Ammothella appendiculata (Dohrn) 

A. exornata Stock 

A. marcusi Hedgpeth 

A. rugulosa (Verrill) 

Ascorhynchus Sars 

Ascorhynchus latipes (Cole) 

Ascorhynchus sp. cf. serratus Hedgpeth 

Eurycyde Schiodte 

Eurycyde raphiaster Loman 

Hedgpethius Child 

Hedgpethius mamillatus, new species 

H. Tridentatus Child 

Nymphopsis Haswell 

Nymphopsis duodorsospinosa Hilton 

Tanyslylum Miers 

Tanystylum birkelandi Child 

7A. tubirostrum Stock 

Family CALLIPALLENIDAE 

Callipaltene Flynn 

Callipallene belizae, new species 

C. emaciata (Dohrn) 

Parapallene Carpenter 

Parapallene bermudensis Lebour 

Pigrogromilus Caiman 

Pigrogromitus timsanus Caiman 

Family PHOXICHILIDIIDAE 

Anoplodactylus Wilson 

Anoplodactylus bahamensis Child 

A. balangensis (Heifer) 

A. evelinae Marcus 

A. imswe, new species 

A. jonesi Child 

A. mantimus Hodgson 

A. monolrema Stock 

A. mulliclavus Child 

A. pectinus Hedgpeth 

A. partus Caiman 

Anoplodactylus sp. 

Family ENDEIDAE 

Endeis Philippi 

Endeis spinosa (Montagu) 

Family NYMPHONIDAE 

Nymphon Fabricius 

Nymphon floridanum Hedgpeth 

Family RHYNCHOTHORACIDAE 

Rhynchothorax Costa 

Rhynchothorax architectus Child 

R. crenatus, new species 

Famil y AMMOTHEIDAE 

Achelia Hodge, 1864 

Achelia sawayai Marcus, 1940 

Achelia sawayai.—Krapp and Kraeuter, 1976:342-343 [literature].—Child, 1979:7-8. 

MATERIAL EXAMINED.—Carrie Bow Cay: On 

corals, 9.1 to 30.5 m, by hand with SCUBA, 7 

Apr 1973; 16 with eggs. Tidal flats in 0.5 m, coll. 

K. Riitzler, 2 May 1974; 19. Tidal flats in 0.5 m, 

coll. K. Riitzler, 4 May 1974; 16. Tidal flats in 

0-1 m, coll. K. Riitzler, 7 May 1974; 16. Windward side, reef crest and flat in 0.5 m, 12 Jan 

1976; 1 juvenile. Broken rock and rubble at tide 

line, coll. J. Clark, 17 Jan 1976; 66, 19, 2 juveniles. 

Tidal flats, coll. M. Jones, 8 Apr 1976; 16. Coral 

rubble on inner side of reef crest in 0.2-0.4 m, 

coll. A. Cohen, 15 Jan 1978; 19. Rubble at tide 

line, coll. C. A. Child, 27 Jan 1978; 3 specimens. 

Rubble at tide line, ocean side, coll. C. A. Child, 

28 Jan 1978; 7 specimens. Rocks with algae on 

reef crest, coll. C. A. Child, 29 Jan 1978; 8 

specimens. Algae of the genera Halimeda and Caulerpa from outer reef crest, coll. C. A. Child, 30 

Jan 1978; 9 specimens. From the seagrass Syringodium on lagoon flats in 1.3 m, coll. B. Kensley, 

I Feb 1978; 19. Ocean side flats, plankton net on 

bottom in 0.4 m, coll. R. Larson, 30 Jan 1978; 1 

juvenile. Syringodium bed at S end of lagoon in 1.3 

m, coll. B. Kensley, 3 Feb 1978; 66, 1 juvenile. 

Rubble from tide line in 0.3 m, coll. C. A. Child, 

4 Feb 1978; 3 specimens. Rubble in front of reef 

crest in 0.6 m, coll. B. Kensley, 5 Feb 1978; 26, 

19. Carpet of coralline and fine red algae at shore, 

coll. B. Kensley, 5 Feb 1978; 39. From Halimeda 
NUMBER 12 357 

in same place; 7 specimens. Halimeda and sparse 

rubble from top of reef ridge, in 18 m, coll. C. A. 

Child, 7 Feb 1978; 19. From other Halimeda and 

rubble nearby, coll. B. Kensley, 7 Feb 1978; 2 

specimens. 

Tobacco Reef: About 500 m N of South Water 

Cay, debris at low tide, coll. M. Carpenter and 

R. Larson, 23 Mar 1977; 26. 

Twin Cays: N end of dividing channel, on 

Halimeda in 1 m, coll. C. A. Child, 31 Jan 1978; 

19. NW coast, mat oi Caulerpa verticillata]. Agardh 

and mangrove rootlets beneath mangroves, intertidal, coll. C. A. Child, 31 Jan 1978; 3 specimens. 

NW coast, another Caulerpa mat under mangroves, coll. C. A. Child, 2 Feb 1978; 4 specimens. 

NW coast, mat of Halimeda under mangroves, 

intertidal, coll. C. A. Child, 2 Feb 1978; 4 specimens. 

REMARKS.—This species is by far the most common pycnogonid in the Carrie Bow area and 

occurs from the tide line to a depth of 18 meters. 

It is also common elsewhere in the tropical North 

and South Atlantic to depths of 65 meters. It has 

also been found in Madagascar. 

ECOLOGY.—Several general collections of algae 

made in the shallow waters of various cays indicate that Achelia sawayai lives on associated rubble 

and other substrates rather than on the algae 

itself. This species and others were generally not 

found on clean algae at Carrie Bow Cay, but 

were common where the algae and rubble were 

associated with adherent detritus and sessile 

fauna. Therefore, it is concluded that algae do 

not provide food for A. sawayai, but only a substrate for food attachment. 

Ammothella Verrill, 1900 

Ammothella appendiculata (Dohrn, 1881) 

Ammothella appendiculata.—Stock, 1955:250-252, fig. 18 [literature]; 1975:973-975.—Child, 1974:497; 1979:9. 

MATERIAL EXAMINED.—Carrie Bow Cay: Sand 

trough behind outer reef ridge in 27 m, in coral 

sand and rubble, coll. C. A. Child, 7 Feb 1978; 2 

juveniles. Rubble with Halimeda from top of outer 

reef ridge in 18 m, coll. B. Kensley, 7 Feb 1978; 

16 juvenile. 

Blue Ground Range (Cays): At S end of northernmost cay, on mangrove roots in 0-1 m, coll. 

C. A. Child, 30 Apr 1976; 1 juvenile. 

Twin Cays: NW coast, from Rhizophora roots 

with algae, sponges, ascidians, and hydroids in 

0.5 m, coll. C. A. Child, 31 Jan 1978; 36 with 

eggs, 19, 3 juveniles. From Halimeda mats along 

N channel wall in 1 m, coll. C. A. Child, 2 Feb 

1978; 36 with eggs, 16, 79, 15 juveniles. From red 

sponge along wall of N channel in 1.0-1.5 m, coll. 

M. Carpenter, 2 Feb 1978; 26 with eggs. 

REMARKS.—There is some controversy over the 

status of this species and Ammothella rugulosa (page 

358). The wide morphological variation displayed 

by A. appendiculata almost bridges the gap between 

the two species. Stock (1955:250-252, fig. 18) 

described two forms of A. appendiculata: one with 

the relatively short appendages of European specimens, and another form with very long appendages from the Caribbean. Stock considered that 

his two forms reflected differences between the 

ultimate and penultimate molt in adults, the 

ultimate molt producing the "long form." The 

above material is in agreement with the "long 

form" but includes several chelate juveniles, thus 

invalidating the use of adult morphology as an 

explanation for the range of variation. Raising 

even more questions than these littoral specimens 

are the juveniles from depths of 18 and 27 meters 

with both abdomen and ocular tubercle more 

than twice as long as the "short form." 

As Stock pointed out, apparently no single 

visible character can be used to separate Ammothella appendiculata from A. rugulosa taxonomically, if 

indeed they are separate species. It is somewhat 

easier to separate them on the basis of chelifore 

scape segment lengths (subequal for A. rugulosa) 

and ocular tubercle and abdomen length (measurably shorter for A. rugulosa). Stock (1975:973) 

noted that these differences are probably insufficient to separate the two species. I have no 

stronger criteria for keeping the species separate, 

but with the inadequate material currently available, I will regard them as separate until a larger 
358 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

collection can be measured and diagnosed with 

more certainty. 

ECOLOGY.—This species has been found in a 

wide variety of littoral habitats throughout the 

Caribbean and on the Pacific side of Panama 

besides Europe and the Middle East. It has been 

collected in association with Rhizophora, several 

species of algae, Thalassia, sand, rubble, corals, 

sponges, and ascidians. This distribution indicates 

a wide food preference, probably including hydroids and other coelenterates, sponges, and possibly ascidians. 

Ammothella exornata Stock, 1975 

Ammothella exornata Stock, 1975:975-978, figs. 7c-d, 8.— 

Child, 1979:9, fig. 3a-c. 

MATERIAL EXAMINED.—Twin Cays: NW coast, 

mat of Caulerpa verticillata and mangrove rootlets 

under Rhizophora in intertidal, coll. C. A. Child, 

2 Feb 1978; 39, 2 juveniles. 

REMARKS.—The male characters were described and figured by Child (1979:9, fig. 3a-c). 

This is the third record of this species. Ammothella 

exornata is so distinctive that it is easily recognized, 

even as a juvenile. It is the only known species 

from the Caribbean littoral having median trunk 

tubercles. 

ECOLOGY.—In Belize, this species is known only 

in association with Rhizophora mangle Linnaeus 

and Caulerpa sp. I collected several juveniles associated with algae (not Caulerpa) on the prop 

roots of R. mangle in the U.S. Virgin Islands. The 

type-specimens, from St. Martin, were also collected on and near this mangrove. The only other 

capture record is Stock's (1975) two specimens 

collected on algae at Bonaire. The species, however, is not common in this mangrove habitat. At 

least six algae samples were taken from mangrove 

roots at about the time of the above capture but 

no other specimens of Ammothella exornata were 

found. The preferred sessile fauna used for food 

probably was not with the algae on and around 

the roots where the six samples were taken. No 

bryozoans, sponges, or hydroids were found in the 

Caulerpa sample. Therefore, the primary food of 

this species, as with most pycnogonid species, 

remains unknown. 

Ammothella marcusi Hedgpeth, 1948 

Ammothella marcusi Hedgpeth, 1948:247-249, fig. 39b-g; 

1954:427.—Stock, 1975:975, fig. 7a-b.—Child, 1979:9, 11. 

MATERIAL EXAMINED.—Carrie Bow Cay: tide 

flats in 0.5 m, coll. K. Riitzler, 4 May 1974; 16. 

Twin Cays: NW Coast, mat of Caulerpa verticillata and mangrove rootlets under Rhizophora in 

intertidal, coll. C. A. Child, 2 Feb 1978; 16. 

REMARKS.—These specimens agree well with 

Hedgpeth's figures of the type, also a male, and 

Stock's clarifying figures of the palp and terminal 

leg segments. Superficially, Ammothella marcusi is 

much more setose than the other species of Ammothella in this report. There are many more 

clubbed, plain, and feathered setae on these specimens than have been figured before. 

The species is much less common than its 

nearest Caribbean relation, Ammothella appendiculata, which is known to have an extremely wide 

range of habitats in its amphi-Atlantic distribution. The above records extend the known range 

of A. marcusi to Belize from Florida, Panama, the 

eastern Caribbean islands, and the Mexican Pacific coast. 

ECOLOGY.—This species has been found in shallow water on debris, a sandy reef, Lithothamnion 

flats, rubble, algae, and on tide flats. This distribution shows no pattern except that the principal 

food of the species is not restricted to particular 

substrates. 

Ammothella rugulosa (Verrill, 1900) 

Ammothella rugulosa.—Stock, 1975:972 [literature].—Child, 

1979:11. 

MATERIAL EXAMINED.—Carrie Bow Cay: Rubble along tide line on ocean side, coll. J. Clark, 17 

Jan 1976; 16. Large Halimeda clump from SE 

shore, coll. B. Kensley, 5 Feb 1978; 96, 29. 

South Water Cay: SW end, scrapings from 

dock pilings in 0.5 m, coll. C. A. Child, 30 Jan 

1978; 1 juvenile. 
NUMBER 12 359 

Twin Cays: N channel, sponge with ectoprocts 

in 0.5 m, coll. R. Larson, 31 Jan 1978; 2 juveniles. 

NW coast, mat of Caulerpa verticillata and mangrove rootlets under Rhizophora in intertidal, coll. 

C. A. Child, 31 Jan 1978; 76 with eggs, 19, 3 

juveniles. Another mat from nearby, coll. C. A. 

Child, 2 Feb 1978; 56 with eggs, 26, 39, 2 juveniles. Halimeda mat from nearby, coll. B. Kensley, 

2 Feb 1978; 19, 1 juvenile. N channel, red sponge 

from wall of channel in 1.0-1.5 m, coll. M. Carpenter, 2 Feb 1978; 16 with eggs. 

REMARKS.—All specimens of Ammothella with 

short ocular tubercle, short abdomen, and first 

chelifore segment approximately equal to the second are tentatively placed under A. rugulosa. 

Those without these qualifications are placed 

with A. appendiculata (see remarks under that species). 

The female from the Halimeda mat of 2 Feb 

1978, has an extra eye situated below the normal 

left anterior eye. It is round, half the size of the 

one above, and is darkly pigmented, matching 

the four eyes above. 

Scattered records indicate that Ammothella rugulosa is distributed from Bermuda to Florida and 

through the Caribbean to Brazil. 

ECOLOGY.—This species lives among fouling 

organisms on ships and piers and has been taken 

in association with Sargassum, in addition to the 

habitats listed above. 

Ascorhynchus Sars, 1877 

Ascorhynchus latipes (Cole, 1906) 

Ascorhynchus latipes.—Hedgpeth, 1948:256 [literature]; 1954: 

427.—Fage, 1952:530.—Stock, 1953:304 [key]; 1954:116; 

1975:969.—Child, 1979:15-16. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tide 

line on ocean side among rocks and rubble, coll. 

J. Clark, 17 Jan 1976; 19, 1 juvenile. Tidal flats 

on ocean side, from plankton net resting on bottom in 0.5 m at night, coll. R. Larson, 30 Jan 

1978; 19. 

REMARKS.—This is another species for which 

the range is extended to the western Caribbean. 

Its occurrence from Florida and the Bahamas to 

Bonaire, and also at Dakar, Senegal, gives it an 

amphi-Atlantic distribution. 

ECOLOGY.—This littoral species appears to 

have some preference for sand and rock habitats, 

but the collecting records are too scarce to be 

certain. 

Ascorhynchus sp. cf. serratum Hedgpeth, 1948 

Ascorhynchus serratum Hedgpeth, 1948:259-260, fig. 44a-f. 

Ascorhynchus serralus.—Stock, 1975:969. 

MATERIAL EXAMINED.—Carrie Bow Cay: In 

coral and algae on outer vertical reef face in 27-

30 m, coll. C. A. Child, 29 Apr 1976; 1 juvenile. 

REMARKS.—This specimen probably represents 

a new species, but is sufficiently immature that a 

description should be postponed until an adult 

can be collected. It shows several similarities to 

Ascorhynchus serratum, in having 3-segmented chelifores, tall ocular and median trunk tubercles, 

and a long curved oviger claw. The differences 

exhibited by this juvenile are that it (1) is without 

lateral process tubercles; (2) has Ammothella-\ike 

terminal leg segments, but without auxiliary 

claws; (3) has very short legs and oviger segments 

in comparison with A. serratum; (4) has very short 

first tibiae; and (5) has very long dorsodistal leg 

setae. Some or most of these characters may be 

the result of growth stage, but tubercles are usually well developed by this stage. The absence of 

lateral process tubercles possibly places this specimen in a separate species. 

The depths at which Ascorhynchus serratum has 

been captured, roughly from 400 to 700 meters 

off Florida and in the Yucatan Channel, would 

remove this specimen from consideration as the 

same species except that we know nothing of 

vertical migration during growth in pycnogonids. 

We do know that some pycnogonids have a narrow range of temperature tolerances, and perhaps 

vertical migration is limited by this factor. Certainly, the temperatures from 30 meters to 700 

meters cover a wide thermal spectrum. 
360 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Eurycyde Schiodte, 1857 

Eurycyde raphiaster Loman, 1912 

Eurycyde raphiaster.—Stock, 1975:979 [literature].—Child, 

1979:21, fig. 5i-j. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats in 0.5 m, coll. K. Riitzler, 4 May 1974; 16. 

Rock and algae from both sides of reef crest at 

low tide, coll. M. Carpenter, 20 Mar 1977; 19. 

Rocks and algae from just in front of reef crest in 

0.4 m, coll. B. Kensley, 29 Jan 1978; 19. From 

sandy rubble with Halimeda behind outer reef 

ridge in 27 m, coll. C. A. Child, 7 Feb 1978; 1 

juvenile. From Halimeda clump behind outer reef 

ridge in 27 m, coll. B. Kensley, 7 Feb 1978; 1 

juvenile. 

REMARKS.—Females of this species usually have 

lateral process and first coxa tubercles that are 

much smaller than thos£ of the male, and the 

lateral processes are sometimes placed closer together, imparting a dimorphic appearance to the 

species. Otherwise, Eurycyde raphiaster is easily distinguished from others of the genus in the western 

Atlantic. 

The juveniles collected from 27 meters appear 

to mark the deepest limit at which this species 

has been taken in the western Atlantic. Loman 

(1912:13) lists the capture depth as 91 meters for 

his type from the Cape Verde Islands. 

ECOLOGY.—Three of the above records show 

this species to be associated with Halimeda. It has 

many other habitats throughout its amphi-Atlantic range. It was one of the few pycnogonids 

found in the high energy wave action area of the 

reef crest, but it was also taken in the calm water 

of the sand trough behind the outer reef ridge. 

Hedgpethius Child, 1974 

EMENDED DIAGNOSIS.—Ammotheidae. Ascorhynchus-Mke with minutely papillose body surface, 

without trunk or lateral process tubercles. Anterior trunk segment longer than combined length 

of posterior 3 segments; first lateral processes at 

extreme posterior of first segment, imparting 

"long necked" appearance. Proboscis with 3 anterior-pointing tubercles arranged laterally and 

ventrally around its largest circumference. Scape 

2-segmented, very short, chela vestigial. Palp 7-

or 8-segmented, third segment with swelling. Female oviger rudimentary, 3-segmented. Male oviger 9-segmented, Ammothella-\ike, without denticulate spines. Propodus with much reduced or 

missing main claw, auxiliaries large, very curved. 

Hedgpethius mamillatus, new species 

FIGURE 163a-/ 

MATERIAL EXAMINED.—Carrie Bow Cay: Broken rock and rubble at tide line, coll. J. Clark, 17 

Jan 1976; 1 ovigerous female, holotype (USNM 

170997). 

DESCRIPTION.—First body segment 0.5 longer 

than combined length of posterior 3 segments. 

Lateral processes short, less than half trunk diameter, separated by greater than their own diameter, without setae or tubercles. Ocular tubercle a rounded cone as tall as neck diameter, 

situated at midlength of first trunk segment. Eyes 

lightly pigmented. Abdomen cylindrical, carried 

half erect" not extending beyond posterior lateral 

processes, armed with 2 distal setae. 

Proboscis large, ovoid, without marked constrictions, with 2 dorsolateral and 1 ventral anterior-pointing tubercles at widest diameter of 

proboscis and at one-third its length. Anterior to 

each tubercle is a slight bulge in same longitudinal axis. Mouth with 3 lateral and ventral slits 

with flat distinct lips. 

Scape 2-segmented, short, carried in cowling 

around anterior of first trunk segment. First scape 

segment longest, with single dorsodistal seta. Second segment minute. Chela tiny, with anterior 

crease and no fingers. 

Palp 7-segmented, thin. First 2 segments small, 

no longer than wide. Third segment about 5 times 

as long as wide, with a distinct posterior bulge. 

Fourth and sixth segments tiny, only slightly 

longer than wide. Fifth segment longest, extremely slender, with 3-4 endal setae. Terminal 

segment thin, 4 times as long as wide, armed with 
NUMBER 12 361 

FIGURE 163.—Hedgpethius mamillatus, new species (holotype): a, trunk; b, trunk, lateral; c, oviger; 

d, third leg; e, terminal segments of third leg; / palp. Hedgpethius tridentatus Child, male: g, 

anterior trunk segments, lateral; h, anterior of trunk, dorsal; i, femur with enlargement of 

femoral cement gland;/ oviger; k, oviger terminal segments. 
362 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

endal and distal setae longer than segment diameter. 

Oviger (female) rudimentary, tiny, of 3 segments, arising ventrally just anterior to first lateral processes. 

Legs thin, femorae with eggs. Second tibia 

slightly longer than first, femur slightly shorter 

than both. Major segments with single long dorsodistal seta, longer than segment diameter, and 

several short setae. Tarsus and propodus short, 

without spines but having a row of sole setae as 

long as segment diameter. Main claw lacking, 

auxiliaries large, strongly curved. 

MEASUREMENTS (mm).—Ocular segment 

length, 0.44; total trunk length (anterior tip to tip 

of fourth lateral processes), 0.74; trunk width 

(across second lateral processes), 0.23; proboscis 

length, 0.37; abdomen length, 0.1; third leg, coxa 

1, 0.09, coxa 2, 0.15, coxa 3, 0.09, femur, 0.34, 

tibia 1, 0.37, tibia 2, 0.38, tarsus, 0.04, propodus, 

0.13, auxiliary claw, 0.05. 

DISTRIBUTION.—Known only from the type-locality, Carrie Bow Cay, Belize, littoral. 

ETYMOLOGY.—Named for the two breast-like 

lateral tubercles on the proboscis. 

REMARKS.—The new species differs from Hedgpethius tridentatus in having one less palp segment, 

a longer abdomen, the ocular tubercle placed on 

the middle of the neck, and by having large 

auxiliary claws without a main claw, however 

small, between them. The male oviger of this new 

species remains unknown, but will probably resemble that of H. tridentatus, with the same extreme sexual dimorphism displayed for that species. 

Hedgpethius tridentatus Child, 1974 

FIGURE \63g-k. 

Hedgpethius tridentatus Child, 1974:494-497, fig. 1. 

MATERIAL EXAMINED.—Twin Cays: NW coast, 

mat of Caulerpa verticillata and mangrove rootlets 

beneath Rhizophora in intertidal, coll. C. A. Child, 

31 Jan 1978; 16, 19. Another mat of the same 

substrate from just S of first collection, coll. C. A. 

Child, 2 Feb 1978; 13 specimens. 

DESCRIPTION OF MALE.—Proboscis tubercles 

slightly larger than those of female. Femoral cement gland a tall tube placed at midlength of 

dorsal surface, perpendicular to femur. Oviger 

with 9 segments: second the longest; sixth ovoid; 

terminal 3 segments (strigilis) placed anaxially on 

middle of sixth segment; strigilis segments short, 

curved, with simple spines; all but first oviger 

segment armed with several setae; without terminal claw. 

EMENDED DESCRIPTION OF FEMALE.—Palp of 8 

segments: first two very short; third and fifth 

longest; sixth and seventh very short; terminal 

segment thin, with ventral setae. Chelifores of 3 

segments, partly hidden by anterior cowling of 

first trunk segment. Scape with 2 segments, first 

a short cylinder with dorsodistal seta, second a 

wrinkled bud, no longer than wide. Chela vestigial, with distal crease but no fingers. Juvenile 

chela with strongly curved fingers overlapping at 

tips. 

REMARKS.—The smallest palp and chelifore 

segments are extremely difficult to see and were 

missed in making the first description. The males 

of this species have the same number of segments 

in palps and chelifores as listed above in the 

emended female description. 

The extreme sexual dimorphism found in the 

ovigers of this species is very rare in pycnogonids. 

In most pycnogonid genera, either ovigers are 

lacking in the female or the ovigers are smaller 

slightly modified versions of the male ovigers. 

Enough females of this genus have now been 

collected to show that the very reduced ovigers 

reported in the original description are not misinformation based on damaged specimens, but 

are the natural state for females of Hedgpethius, in 

which they appear almost embryonic. 

This genus was named and published before I 

received the paper by Turpaeva (1973), in which 

she assigns a number of species of Rhopalorhynchus 

(Colossendeidae) to a new genus, Hedgpethia. According to the International Code of Zoological 
NUMBER 12 363 

Nomenclature, Article 56a, both names must be 

retained. The two genera are sufficiently different 

in most characters that they will never become 

synonymous. 

The known distribution for Hedgpethius tridentatus is extended from Florida to the Belizean 

barrier reef islands; all records are littoral. 

Nymphopsis Has well, 1885 

Nymphopsis duodorsospinosum Hilton, 1942a 

Nymphopsis duodorsospinosum Hilton, 1942a:303-305, pl. 45.— 

Hedgepeth, 1948:250-252, fig. 40; 1954:427.—Child and 

Hedgpeth, 1971:609 [list].—Kraeuter, 1973:496.—Stock, 

1975:978.—Krapp and Kraeuter, 1976:342.—Child, 

1979:21. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats in 0.5 m, coll. K. Riitzler, 7 May 1974; 19. 

REMARKS.—This species is easily recognized by 

its large size in comparison with tiny littoral 

species, its legs and trunk, which are crowded 

with dorsal tubercles, and particularly the two 

tall trunk tubercles, which separate it from all 

other pycnogonids known from the Belizean 

coast. It has been found from the tide line to 

about 60 meters of depth on various substrates, 

but its habits and feeding preferences remain 

unknown. 

Nymphopsis duodorsospinosa has a wide tropical 

and temperate distribution, from Georgia and 

Florida through the Caribbean to Panama and 

from the Gulf of California to the Pacific coast of 

Panama. It is also found in the Galapagos Islands. 

Tanystylum Miers, 1879 

Tanystylum birkelandi Child, 1979 

Tanystylum birkelandi Child, 1979:23, fig. 7. 

MATERIAL EXAMINED.—Carrie Bow Cay: Flats 

on ocean side, rubble and calcareous algae from 

tide line, coll. C. A. Child, 27 Jan 1978; 29. 

REMARKS.—These two females agree exactly 

with the female paratype from Galeta Island, on 

the Caribbean side of Panama. Since this is only 

the second capture record for the species, little 

can be said concerning its habitats, except that it 

has been taken with coralline and calcareous 

algae in the littoral. Its distribution is here extended north from Panama to the Belizean coast. 

Tanystylum tubirostrum Stock, 1954 

Tanystylum tubuostre Stock, 1954:117-120, figs. 24-25.—Bourdillon, 1955:600, pl. 3: figs. 2-4.—Child, 1979:34-35. 

Tanystylum tubirostrum.—Stock, 1975:984. 

MATERIAL EXAMINED.—Twin Cays: NW coast, 

on roots of Rhizophora mangle with adherent red 

and green sponges, algae, ascidians and hydroids 

in 0.5 m, coll. C. A. Child, 31 Jan 1978; 26, 19. 

REMARKS.—These specimens agree in all respects with Stock's (1954) description and figures. 

The present capture extends the known distribution of this species from Bermuda, Puerto Rico, 

Curasao and Bonaire, the western Caribbean, 

and the Pacific shores of Mexico and Panama. 

ECOLOGY.—There are few capture records for 

this species, but these are fairly well documented. 

It lives in littoral habitats and has been found 

associated with Sargassum, hydroids, and, in this 

case, with a wealth of potential food. Pycnogonids 

are known to feed on hydroids and ascidians and 

because they have been taken in association with 

sponges, it is assumed that they also feed on the 

soft parts of sponges. 

Famil y CALLIPALLENIDAE 

Callipallene Flynn, 1929 

Callipallene belizae, new species 

FIGURE 164. 

MATERIAL EXAMINED.—Carrie Bow Cay: Halimeda and rubble from outer reef ridge in 18 m, 

coll. B. Kensley, 7 Feb 1978; 16 with eggs, holotype (USNM 171035), 2 juvenile paratypes 

(USNM 171036). Large clump oi Halimeda from 

same area, coll. C. A. Child, 7 Feb 1978; 19 
364 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 164.—Callipallene belizae, new species (holotype): a, trunk; b, trunk, lateral; c, proboscis, 

ventral; d, chela with enlargement of tip of movable finger; e, third leg;/ terminal segments of 

third leg; g, oviger with single egg; h, terminal segments of oviger with enlargement of two 

spines. 

paratype (USNM 171037). 

DESCRIPTION.—First 2 trunk segment lines present, third lacking. Neck short, unadorned. Lateral processes short, only as long as their diameters, separated by less than their diameters, glabrous. Ocular tubercle a broad truncated cone 

capped with a thin tubercle as tall as ocular cone. 

Eyes large, lightly pigmented. Abdomen an in-
NUMBER 12 365 

flated cylinder slightly longer than twice its maximum diameter, armed with 2 distal setae. 

Proboscis short, inflated distally and with 

rounded ventrodistal bulges. 

Chelifore scape equal in length to proboscis, 

armed with lateral and dorsodistal setae. Chela 

with slightly inflated palm, armed with 7-8 dorsal 

and 1 ventral setae. Fingers overlap at tips, without setae. Movable finger with single tiny distal 

tooth, broader than long. 

First 3 segments of oviger moderately short, 

fourth longer, fifth more than twice length of 

fourth, armed with several setae longer than fifth 

segment diameter. Distal apophysis of male fifth 

segment armed with 2 setae. Terminal 4 segments 

(strigilis) armed with denticulate spines in the 

formula 6:5:5:5. Denticulate spines dimorphic: 

proximal spines oval with fine denticulations; 

distal spine larger, with fan-like projection of 

denticulations pointing distally on segment. 

Legs long, slender, very setose distally. First 

and third coxae short, little longer than their 

diameters. Second coxa long, curved, over 5 times 

longer than its maximum diameter. Femur 

slightly longer than tibia 1, tibia 2 slightly longer 

than femur. Cement glands not found. Tarsus 

short, armed with 2 smooth ventral spines, 1 

ventral and 1 dorsal setae. Propodus short, only 

slightly curved, armed distally and laterally with 

many long setae. Sole with 4 smooth heel spines 

and 7 distal spines. Main claw half as long as 

propodus, auxiliaries about 0.9 times length of 

main claw, without endal setae or teeth. 

MEASUREMENTS (mm).—Trunk length (chelifore insertion to tip of fourth lateral processes), 

0.89; trunk width (across second lateral processes), 0.37; abdomen length, 0.12; proboscis 

length, 0.28; third leg, coxa 1, 0.12, coxa 2, 0.65, 

coxa 3, 0.19, femur, 0.96, tibia 1, 0.77, tibia 2, 

1.1, tarsus, 0.06, propodus, 0.28, claw, 0.14. 

DISTRIBUTION.—Known only from the type-locality, Carrie Bow Cay, Belize, in depths of 18 

meters. 

ETYMOLOGY.—Named for the country where it 

was discovered, Belize. 

REMARKS.—This new species resembles American specimens of Callipallene brevirostris (Johnston), but has a shorter neck; it is also similar to 

Caribbean specimens of C. emaciata. It differs from 

both of these species in the following respects: the 

new species has a tall thin tubercle on top of its 

truncated ocular cone; it has smooth tarsus and 

propodus spines whereas the other two have variously crenulated spines; its chela fingers are without teeth except for a single tiny distal tooth on 

the movable finger; the auxiliary claws are longer 

than for most American specimens, although 

those of C. brevirostris are sometimes as long; and 

the posterior trunk segmentation line is lacking, 

although this is not a reliable taxonomic character with this genus. 

Taxonomic distinction among Callipallene species is often very difficult to make, not only 

because several species are very similar, but also 

because apparently they have more than one 

adult molt stage with resulting changes in setae, 

teeth, denticulate spine number and shape, and 

mensural characters and their ratios. These molt 

changes add vastly to the difficulty in deciding 

where variation ends and species begin, a dividing 

line that will undoubtedly remain uncertain for 

most Callipallene species until large numbers can 

be compared interspecifically and intraspecifically. 

The propodus of Callipallene belizae is shorter, 

less curved, and has many more setae than either 

C brevirostris or C. emaciata. Neither of these species 

has the very marked dimorphism shown in the 

denticulate oviger spines of C. belizae. The terminal spine on each of the four distal segments is 

splayed out and canted forward so as to resemble 

a fan. 

ECOLOGY.—Since this species has been found 

only in rubble and Halimeda at depths of 18 

meters, and the bottom samples from which the 

species was taken were not preserved for possible 

food preference organisms, its habits are unknown. 

Callipallene emaciata (Dohrn, 1881) 

Callipallene emaciata emaciata.—Stock, 1952a:8 [literature]. 

Callipallene emaciata.—Stock, 1975:1011.—Child, 1979:41-42. 
366 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

MATERIAL EXAMINED.—Carrie Bow Cay: From 

Thalassia bed in 0.5 m, coll. R. Larson, 31 Mar 

1977; 16 with eggs, 16, 29, 2 juveniles. From 

Dictyota on reef flat in 0.5 m, coll. R. Larson, 16 

Mar 1977; 1 juvenile. Rubble and calcareous 

algae at wall on ocean side at tide line, coll. C. A. 

Child, 28 Jan 1978; 19. Syringodium and sediment 

from lagoon in 1.5 m, coll. B. Kensley, 1 Feb 

1978; 19. Plankton sampler on bottom on ocean 

side flats in 0.5 m, coll. R. Larson, 30 Jan 1978; 

1 larva (probably this species). 

South Water Cay: Piling scrapings from dock 

at S end on lagoon side, coll. C. A. Child, 30 Jan 

1978; 1 juvenile. 

REMARKS.—These are the first records of Callipallene emaciata from the western Caribbean. This 

species has been found in the Mediterranean, 

Portugal, the Azores, and from Florida and the 

Caribbean archipelago to the Guianas, primarily 

in littoral and shallow depths. 

ECOLOGY.—This species and two others from 

the Caribbean coasts, Callipallene brevirostris and 

C. phantoma (Dohrn) are generally found associated with fouling organisms on bridge and dock 

pilings. 

Parapallene Carpenter, 1892 

Parapallene bermudensis Lebour, 1949 

FIGURE 165. 

Parapallene bermudensis Lebour, 1949:930-932, figs. 2-3. 

MATERIAL EXAMINED.—Carrie Bow Cay: 

Coral, sand and rubble from sand trough behind 

outer reef ridge at 27 m, coll. C. A. Child, 6 Feb 

1978; 19 subadult. Sand and rubble from slightly 

S of above sample in 27 m, coll. C. A. Child, 7 

Feb 1978; 16 juvenile. 

REMARKS.—Although neither of these specimens is fully mature, they agree with Lebour's 

(1949) description and represent only the second 

record of this species; the type-locality is Bermuda. An unreported female from about the 

same depth in the Bahama Islands is in the 

collections of the National Museum of Natural 

History. 

Since Lebour's (1949) figures are somewhat 

stylized and several details are omitted from her 

drawings and description, I have prepared a set 

of figures of the male for clarification. Lebour 

stated that the chela fingers lack denticulations, 

but both the Bahama specimen and the above 

two have very small denticulations or teeth on 

one or both fingers of the chela. 

The species has raised lips and an oral fringe of 

tiny setae. The propodus has two major heel 

spines unlike the figure and description of Lebour's (1949:931-932, figs. 3-7) specimen. The 

ventrodistal second tibia spine, major tarsal spine, 

and the sole spines all show a slight serration on 

their inner surfaces. The auxiliary claws are 

slightly shorter than those figured by Lebour. 

The ovigers of the Carrie Bow female are 

shorter than those of the type-specimen, but the 

female was about to molt and the next (adult) 

stage oviger can be seen clearly within the outer 

chitinous layer. The oviger of the Carrie Bow 

male is still the unsegmented curled appendage 

of a juvenile. 

A distinctive character of this species is its 

spination. There are short broad spines on the 

first coxae and a series of these spines around the 

insertion of the chelifores and proboscis. The 

spines have an annulated hollow interior without 

the annulations being carried through to the 

outer surface (Figure 165/). Elsewhere on the 

animal, where other typical spines and setae are 

present, these annulated spines are absent. 

The Carrie Bow specimens represent a range 

extension to the western Caribbean for this species. 

ECOLOGY.—The four available records for this 

species indicate that it has a very restricted depth 

preference. All four specimens reportedly have 

been taken in depths of 27 to 33 meters (90 to 

100 ft). Lebour's (1949) type was taken "with 

fragments of hydroids and weeds." The Carrie 

Bow specimens were taken from rubble with some 

algae and probably a number of hydroids, although none of the latter were saved. No habitat 

data are available on the Bahamas specimen. 

The Carrie Bow specimens were light brownish 

green. Pigmentation and feeding preference have 
NUMBER 12 367 

FIGURE 165.—Parapallene bermudensis Lebour, juvenile male: a, trunk; b, anterior trunk segments, 

lateral; c, third leg; d, terminal segments of third leg with enlargement of spine; e, chela with 

enlargement of movable finger; / annulated spine. 

not been correlated, but most literature references 

concerning live pycnogonids state that generally 

they are colored the same as the substrate making 

them difficult to see. Their color probably represents some of the algal substrate on which many 

pycnogonids find their food source. 

Pigrogromitus Caiman, 1927 

Pigrogromitus timsanus Caiman, 1927 

Pigrogromitus timsanus Caiman, 1927:408-410, fig. 104a-f.— 

Hedgpeth, 1947:7 [text]; 1948:214-216, fig. 23.—Stock, 

1968a:46; 1975:1015-1016.—Lipkin and Safriel, 1971: 

9.—Arnaud, 1972:159-160.—Child, 1979:46,47. 

Clotenopsa prima Hilton, 1942b:52-53, fig. 8. 

MATERIAL EXAMINED.—Carrie Bow Cay: 

Among rocks and rubble on ocean side at tide 

line, coll. J. Clark, 17 Jan 1976; 2 juveniles. 

Twin Cays: Among Rhizophora roots along N 

edge of dividing channel at S end of Cays in 0.6 

m, coll. C. A. Child, 25 Apr 1976; 16 with eggs. 

REMARKS.—This species has a pantropical distribution. This is the second record of capture in 

the western and southwestern Caribbean, the first 

being from the Caribbean coast of Panama. 
368 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ECOLOGY.—The many and varied habitats of 

this species include corals, algae, zoanthids, mangroves, rubble, and molluscan cavities. It was also 

found among unspecified fouling organisms on 

pilings, almost always at intertidal and subtidal 

depths. 

Family PHOXICHILIDIIDAE 

Anoplodactylus Wilson, 1878 

Anoplodactylus bahamensis Child, 1977 

Anoplodactylus bahamensis Child, 1977:587-589, fig. 2. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats in 0.5 m, coll. K. Riitzler, 2 May 1974; 16. 

REMARKS.—This species was originally described (Child, 1977:587) as having teeth only on 

the movable fingers of the chelae. The male 

collected here has 3 tiny low teeth on the immovable finger also, but it is otherwise indistinguishable from the type, also a male. 

This Carrie Bow male extends the known distribution to the western Caribbean. 

The habitats or associations of this species remain unknown. It has been found at depths of 

0.5 and 12 meters. 

Anoplodactylus batangensis (Heifer, 1938) 

Pycnosoma batangense Heifer, 1938:174-176, fig. 6a-c. 

Anoplodactylus batangensis.—Stock, 1968a:54 [literature]; 1975: 

1082-1083, fig. 43c-d.—Arnaud, 1973:957, figs. 3-4.— 

Child, 1979:50. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats at tide line, coll. K. Riitzler, 23 Apr 1974; 

16 with eggs, 19. Tidal flats at 0.5 m, coll. K. 

Riitzler, 2 May 1974; 16. From Dictyota on reef 

flat in 0.5 m, coll. R. Larson, 16 Mar 1977; 16 

juvenile. From Thalassia bed in 0.5 m, coll. R. 

Larson, 31 Mar 1977; 1 juvenile. Rubble and 

Halimeda from outer reef crest in 0.5 m, coll. B. 

Kensley, 30 Jan 1978; 16 with eggs. Mixed carpet 

of red algae and compact corallines at SE end 

shore, coll. B. Kensley, 5 Apr 1978; 16. 

South Water Cay: Thalassia and red sponge 

beyond dock at S end in 1 m, coll. B. Kensley, 30 

Jan 1978; 39. 

Twin Cays: NW coast, mat of Caulerpa verticillata and mangrove rootlets from under Rhizophora, 

intertidal, coll. C. A. Child, 31 Jan 1978; 16, 

several juveniles. Another C. verticillata and rootlet 

mat from nearby, coll. C. A. Child, 2 Feb 1978; 

36. A mat of Halimeda from nearby, coll. B. 

Kensley, 2 Feb 1978; 19. 

REMARKS.—This species is easily recognized by 

its anteriorly curved and tapered proboscis, 

unique among the many species of this genus. 

Several live and even freshly killed specimens, 

particularly the 36 found on Caulerpa, 2 Feb 1978, 

had a broad chalk-white stripe running from the 

posterior of the ocular tubercle to the base of the 

abdomen. The remainder of these animals ranged 

from cream to slightly straw colored, except for a 

chalk-white band around each distal leg segment 

suture. This white color was not associated with 

the intestinal diverticula, which could be seen 

below and separate from the color line. 

ECOLOGY.—Stock (1975:1083) mentioned that 

males seem to be rare, but both in this collection 

and in another from Panama, males appear in 

equal or greater numbers than females. Of the 14 

adults in this collection, there are 9 males and 5 

females; 2 of the males bear eggs. This imbalance 

may be related to collecting methods. Pycnogonids are generally found incidentally by sorters 

who are looking for other organisms and therefore 

many are missed. 

The habitats of this species are quite varied, 

and include algae, Thalassia, sponges, rubble, and 

Rhizophora. All specimens were taken in littoral 

depths and most records in the literature are also 

from shallow water. 

Anoplodactylus evelinae Marcus, 1940 

Anoplodactylus evelinae Marcus, 1940:55-58, pl. 4.—Hedgpeth, 

1948:232, fig. 31.—Child, 1979: 53. 

Anoplodactylus (Labidodactylus) evelinae.—Stock, 1954:128; 

1975:1083. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats at tide line, coll. K. Riitzler, 23 Apr 1974; 
NUMBER 12 369 

16. Thalassia bed in 0.5 m, coll. R. Larson, 31 

Mar 1977; 16 with eggs. Rubble and calcareous 

algae on ocean side in 0.3 m, coll. C. A. Child, 27 

Jan 1978; 19. Halimeda, Caulerpa, and rubble from 

outer reef crest in 0.5 m, coll. B. Kensley, 30 Jan 

1978; 19. Clump of Halimeda at shore, SE end, 

coll. B. Kensley, 5 Feb 1978; 19. 

Twin Cays: NW coast, Rhizophora roots with 

algae, hydroids, ascidians, and bryozoans in 0.5 

m, coll. C. A. Child, 31 Jan 1978; 2 juveniles. 

NW coast, mat of Caulerpa verticillata and mangrove rootlets under Rhizophora, intertidal, coll. C. 

A. Child, 2 Feb 1978; 46, 29. Mat of Halimeda 

from same area, coll. B. Kensley, 2 Feb 1978; 19. 

REMARKS.—This is another easily recognized 

species. It is very "stumpy" in appearance, with 

a short and broad ocular tubercle, abdomen, and 

proboscis. The legs are robust and the very 

pointed heel with its short spine makes a reliable 

recognition character. 

It is common in the western Caribbean. The 

above collections produced 14 specimens from 

the littoral. 

ECOLOGY.—The habits of Anoplodactylus evelinae 

are unknown, but it is found together with other 

species of pycnogonids in algae, Thalassia, Rhizophora, rubble, and among sessile animals. The 

Belizean collections do not support Stock's (1975: 

1083) suggestion that this species is a sand burrower. 

Anoplodactylus imswe, new species 

FIGURE 166 

MATERIAL EXAMINED.—Tobacco Reef: Reef 

top about 500 m N of South Water Cay, intertidal, coll. M. Carpenter and R. Larson, 23 Mar 

1977; 16 holotype (USNM 171122). 

Carrie Bow Cay: Tidal flats in 0.5 m, coll. K. 

Riitzler, 4 May 1974; 19 paratype (USNM 

171123). Tidal flats in 0.5 m, coll. K. Riitzler, 7 

May 1974; 19 paratype (USNM 171124). 

DESCRIPTION.—Trunk with first two intersegmental lines well marked, third incomplete, 

marked only by slight depression dorsally. Lateral 

processes separated by slightly less than their 

diameters, each as long as trunk diameter, without tubercles, armed with 1-3 dorsodistal setae. 

Ocular tubercle a moderately tall cylinder twice 

as tall as its diameter, capped by triangular cone. 

Eyes distally on cylinder, darkly pigmented. Neck 

armed with single seta lateral to, and in front of, 

ocular tubercle. Abdomen almost 3 times its diameter, bent erect, armed with several distal setae. 

Chelifores thin, scape armed with several distal 

setae. Chela ovoid, fingers shorter than palm, well 

curved at tips, without teeth. Palm with several 

distal setae; movable finger with 3 ectal setae. 

Oviger with robust curved basal segment. Second segment only slightly longer, cylindrical, 

armed with ectal row of 6-7 short setae. Third 

segment longest, slightly less than half again as 

long as second segment, armed with several ectal 

and endal setae. Terminal 3 segments each 

shorter than last, moderately setose distally with 

distal setae longer than segment diameter. Terminal segment thin, pointed at tip. 

Leg with first coxae armed with 2-4 dorsodistal 

setae, without tubercles. Second coxae of third 

and fourth legs with ventrodistal genital spur 

almost as long as segment diameter, carrying 

genital orifice at tip. Coxae of first and second 

legs without spurs or orifices. Third coxae armed 

with several distal setae. Femur the longest leg 

segment with first tibia longer than second, each 

armed with several short setae and a single long 

dorsodistal seta, as long or longer than segment 

diameter. Seta of femur mounted on short tubercle. Femoral cement gland a long, thin, flaskshaped tube, canted distally, situated at less than 

one-third length of femur. Tarsus roughly triangular, with several ventral setae. Propodus stout; 

heel perpendicular, armed with single large spine 

and 5 smaller spines. Sole armed with 6-7 curved 

spines and several lateral setae; distal lamina onefifth sole length. Claw robust, strongly curved. 

Auxiliary claws lacking. 

MEASUREMENTS (mm).—Trunk length (chelifore insertion to tip of fourth lateral processes), 

0.93; trunk width (across first lateral processes), 

0.63; proboscis length, 0.52; abdomen length, 
370 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 166—Anoplodactylus imswe, new species (holotype): a, trunk; b, trunk, lateral; c, proboscis, 

ventral; d, chela; e, third leg with enlargement of femoral cement gland;/ terminal segments 

of third leg; g, oviger terminal segments. 
NUMBER 12 371 

0.24; third leg, coxa 1, 0.21, coxa 2, 0.42, coxa 3, 

0.2, femur, 0.55, tibia 1, 0.52, tibia 2, 0.47, tarsus, 

0.09, propodus, 0.34, claw, 0.23. 

DISTRIBUTION.—Known only from the type-locality, the Belizean barrier reef at Tobacco Reef 

and Carrie Bow Cay at a depth of 0.5 meters. 

ETYMOLOGY.—This species is named for the 

Smithsonian Institution Investigations of Marine 

Shallow-Water Ecosystems (IMSWE) Project. 

REMARKS.—The femoral cement glands of this 

species are not unique among species of Anoplodactylus. They are almost exactly like the glands 

of A. batangensis, but they are placed slightly 

farther back toward the proximal end of the 

segment in the new species. Other differences in 

A. imswe remove it from consideration as A. batangensis. Most notably, the proboscis in the new 

species is cylindrical whereas in A. batangensis it is 

constricted distally to a thin tube. The new species also has longer first oviger segments, longer 

and less setose leg segments, longer ocular tubercle and abdomen, and lacks low lateral process 

tubercles. The propodus, however, with its heel 

and heel spine, sole spines, and lack of auxiliaries, 

is strikingly similar in both species. 

This species is similar to Anoplodactylus erectus 

Cole in the shape of the trunk, ocular tubercle, 

oviger, and abdomen, but the propodus characters and the cement glands are very different. It 

is also similar to A. maritimus. In comparison with 

Giltay's male type specimen of A. parvus {= A. 

maritimus, fide Stock, 1975:1069-1074), the ovigers, chelifores, trunk, and propodus shape of A. 

imswe are very close to those of the type of A. 

parvus. The differences are (1) the propodus of A. 

parvus has two heel spines and auxiliary claws, (2) 

the abdomen and ocular tubercle are shorter, (3) 

there are no discernible trunk segmentation lines, 

and (4) the cement glands are tiny short tubes 

where they emerge from the femorae. 

ECOLOGY.—The three specimens from Belize, 

although collected in separate locations, all share 

a distinct color pattern. When freshly killed, the 

Tobacco Reef specimen had a bright purple gut. 

After storage in alcohol for a year or more, all 

three specimens have deep blue intestinal tracts. 

This blue pigment appears to be a reliable recognition character for this species, at least on the 

Belizean barrier reef. The purple or blue gut color 

is shared by Anoplodactylus lentus, a much larger 

pycnogonid which has not yet been found in 

Belize. The coloration of A. imswe indicates that 

it might feed on organisms containing blue-green 

algae, but since collecting data for this species do 

not contain information on habitat associates, its 

feeding preference remains unknown. 

Anoplodactylus jonesi Child, 1974 

Anoplodactylus jonesi Child, 1974:497-500, fig 2; 1979:56, fig. 

19a-b. 

Anoplodactylus (?) antillianus Stock, 1975:1081-1082, fig. 57. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats in 0.5 m, coll. K. Riitzler, 2 May 1974; 26. 

REMARKS.—Both of these specimens are normal 

males with full ovigers and a tall tubular cement 

gland on each femur. Several questionable females have been found in the past that bear male 

ovigers, ova in the legs, and that lack cement 

glands. This seemingly abnormal situation has 

led to speculation on the status of this species in 

a genus in which the females are entirely without 

ovigers. Further discussion and figures of this 

species appear in Child (1979:56, fig. 19a-b). 

ECOLOGY.—Habits and habitats are lacking for 

the Belizean specimen, but in Panama the species 

has been found most often among algae. 

Anoplodactylus maritimus Hodgson, 1914 

Anoplodactylus maritimus Hodgson, 1914:164; 1915:148; 1927: 

357.—Marcus, 1940:60.—Stock, 1975:1069-1074, fig. 54. 

Not Anoplodactylus maritimus.—Hedgpeth, 1948:230, fig. 

29d-e [= A. mleus Stock, 1975:1069]. 

Anoplodactylus parvus Giltay, 1934:1-3, figs. 1-5.—Hedgpeth, 

1948:223-224, fig. 27e-f.—Stock, 1951:13, figs. 14-16; 

1954:127; 1955:235; 1957:85; 1975:1069-1074, fig. 54.— 

Bourdillon, 1955:590-591, fig. 1, pl. 1: fig. 2.—Fage and 

Stock, 1966:326.—Kraeuter, 1973:494-495. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats, coll. M. L. Jones, 6 Apr 1976; 19. 

South Water Cay: Rubble from edge of chan-
372 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

nel in 6 m, coll. C. A. Child, 1 Feb 1978; 26 with 

eggs, 2 juveniles. 

Twin Cays: NW coast, Halimeda mat from 

under mangroves in 0.3 m, coll. B. Kensley, 2 Feb 

1978; 19. 

REMARKS.—In agreement with Stock (1975: 

1072), I have combined the two species under 

Hodgson's (1914) earlier designation. There is 

ample cause for confusion here as Hodgson's type 

was never figured. Giltay's (1934) Anoplodactylus 

parvus has been figured by several authors, including Giltay, but was never compared with Hodgson's type (which appears to have been lost). As 

presently designated, the species can also be easily 

confused with A. petiolatus (Kroyer), which itself 

is sufficiently variable to suggest the possibility of 

two species (Stock, 1975:1075). Stock (1975, figs. 

53, 54) attempted to sort out and figure the 

differences between the present species and A. 

petiolatus. 

Association with floating substrates has given 

this species an extremely wide distribution in the 

Atlantic. It has been found from Chesapeake Bay 

in Virginia, south to Bermuda and through the 

West Indies to Brazil. Hodgson (1914) originally 

described it from south of the Azores and it is 

known also from the Cape Verde Islands. 

ECOLOGY.—This species is common over vast 

areas of the mid-Atlantic as one of the many 

inhabitants of the seaweed Sargassum (Bourdillon, 

1955, fig 1). The type oi Anoplodactylus maritimus 

was collected in Sargassum, and many other records confirm this association. I have also seen 

specimens feeding on hydroids attached to Sargassum dipped from Florida waters. The present 

collection, however, shows that this species occupies other habitats as well. 

Anoplodactylus monotrema Stock, 1979 

Anoplodactylus robustus.—Child and Hedgpeth, 1971:612-613 

[literature].—Stock, 1975:1080. 

Anoplodactylus monotrema Stock, 1979:15-18, figs. 4-5.—Child, 

1979:56, 58, fig. 19c. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats in 0-1 m, coll. 7 Apr 1973; 16 with eggs. 

REMARKS.—This species has a "fat" appearance with a short blunt proboscis, abdomen, and 

ocular tubercle. 

All specimens previously assigned to Anoplodactylus robustus, whether from European collections 

or from western hemisphere collections, were 

thought to agree. It recently became evident that 

the European specimens have more than one 

(usually 3) cement gland per femur whereas the 

American specimens have only one gland per 

femur. Other slight differences, when taken in 

combination, caused American specimens to be 

recognized as a separate species, A. monotrema. 

Therefore, the known distribution of the American species is confined to the east and west coasts 

of southern North America and northern South 

America, including the Galapagos Islands. 

ECOLOGY.—The color of this littoral and sublittoral species is usually straw or lighter. Its 

feeding habits are not yet identified. It has been 

taken on a wide variety of substrates. 

Anoplodactylus multiclavus Child, 1977 

Anoplodactylus multiclavus Child, 1977:593-596, fig. 4; 1979: 

58, fig. 19d. 

MATERIAL EXAMINED.—South Water Cay: SE 

side, in shallow grass bed with sand patches near 

Rhizophora stumps in 0.5 m, coll. J. Clark, 15 Jan 

1976; 16 with eggs. 

REMARKS.—This specimen agrees with the 

type, also a male, in all respects including the 

multiple cement glands, except that it appears 

slightly more robust. 

ECOLOGY.—Both the type and this specimen 

have been found in association with or near mangroves. 

Anoplodactylus pectinus Hedgpeth, 1948 

Anoplodactyluspectinus Hedgpeth, 1948:234-236, fig. 34; 1954: 

427.—Stock, 1955:235; 1974:17.—Arnaud, 1973:955-

957.—Child, 1974:500; 1979;58. 

Anoplodactylus pectims [sic].—Stock, 1975:1050-1052, fig 41a. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats at tide line, coll. K Riitzler, 23 Apr 1974; 

16. From Dictyota on reef flat in 0.5 m, coll. R. 

Larson, 16 Mar 1977; 16 with eggs, 16. Rubble 
NUMBER 12 373 

from sand trough behind outer reef ridge in 27 

m, coll. C. A. Child, 6 Feb 1978; 36, 29. Rubble 

from just S of 6 Feb collection, in 27 m, coll. C. 

A. Child, 7 Feb 1978; 36. 

Twin Cays: NW coast, mat of Caulerpa verticillata and mangrove rootlets beneath Rhizophora, 

intertidal, coll. C. A. Child, 2 Feb 1978; 19. Mat 

oi Halimeda from nearby, intertidal, coll. B. Kensley, 2 Feb 1978; 19. 

REMARKS.—This species, including its females, 

is easily recognized by the pectinate major heel 

spine that can be seen under high magnification. 

This marks the second time Anoplodactylus pectinus has been collected in the western Caribbean. 

Previously, it has been collected on the Caribbean 

coast of Panama, in Florida, the Caribbean Leeward Islands, and in Madagascar. 

ECOLOGY.—Freshly killed specimens had light 

green intestinal diverticula. This color may be a 

character useable to distinguish fresh specimens 

of this species, at least on the Belizean barrier 

reef. Anoplodactylus portus shares this green color, 

but retains coloration even after long storage, 

unlike A. pectinus. The color is undoubtedly a 

function of diet, although the particular green 

food matter is unknown. 

Anoplodactylus portus Caiman, 1927 

Anoplodactylus portus.—Stock, 1975:1052-1053, fig. 41b-e [literature].—Child, 1978:133-144, figs. 1-4; 1979:58-59. 

MATERIAL EXAMINED.—Carrie Bow Cay: Rubble and Halimeda from reef crest in 0.5 m, coll. C. 

A. Child, 29 Jan 1978; 16 with eggs. Lagoon flats, 

with Syringodium and sediment in 1.2 m, coll. B. 

Kensley, 1 Feb 1978; 16 juvenile, 1 larva. Mixed 

algae and compact corallines at shore, coll. B. 

Kensley, 5 Feb 1978; 19 juvenile. Halimeda and 

rubble from outer reef ridge in 18 m, coll. B. 

Kensley, 7 Feb 1978; 19 juvenile. 

South Water Cay: Rubble from edge of channel in 6 m, coll. C. A. Child, 1 Feb 1978; 16, 29. 

Twin Cays: NW coast, mat of Caulerpa verticillata and mangrove rootlets under Rhizophora, 

intertidal, coll. C. A. Child, 2 Feb 1978; 19. 

REMARKS.—This is a pantropical species with 

a robust appearance which aids in distinguishing 

it from the many other tropical species of this 

genus. As with all species of Anoplodactylus, the 

configuration of the male femoral cement 

gland(s) is a key character. In conjunction with 

oviger, chela, and propodus characters, it facilitates recognition among these otherwise difficult 

species. This observation does not hold for females, which remain difficult if not impossible to 

identify if they are not taken in the same sample 

or area with recognizable males. 

ECOLOGY.—This species is often collected with 

gut diverticula in the legs showing green chloroplasta or at least chlorophyllous coloration ingested from the algal habitat in which it is often 

found. Whether or not it picks this up directly as 

food or as a biproduct of eating algal-grazing 

fauna has never been demonstrated. 

Anoplodactylus sp. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats, coll. K. Riitzler, 4 May 1974; 19 juvenile. 

Lagoon surface among Thalassia and Sargassum, 

coll. R. Larson with plankton net, 25 Jan 1978; 

1 larva. Tidal flat, plankton net resting on bottom 

in 0.5 m, coll. R. Larson, 30 Jan 1978; 1 larva. 

With Syringodium in 1.0-1.5 m, coll. B. Kensley, 3 

Feb 1978; 29 juveniles. 

REMARKS.—None of these specimens is mature 

enough to identify. 

Family ENDEIDAE 

Endeis Philippi, 1843 

Endeis spinosa (Montagu, 1808) 

Endeis spinosa.—Hedgpeth, 1948:238-240 [early literature].—Stock, 1952b: 185-186; 1954:128; 1957:85; 1962: 

218; 1968a:59 [key]; 1968b:32, fig. 25.—Soyer, 1966:3.— 

deHaro, 1966:9; 1967:109, 112-113, fig. 5.—Krapp, 1973: 

72.—Child, 1979:66. 

MATERIAL EXAMINED.—Carrie Bow Cay: Thalassia beds in lagoon, 1.5 m, coll. J. D. Ferraris, 11 

May 1975; 16 with eggs. 

Stann Creek: Surface plankton tow off Pelican 
374 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Beach, 0.5 mi out, coll. R. Larson, 13 Apr 1978; 

19. 

REMARKS.—This species has been found from 

Norway to Argentina, including the Mediterranean Sea, at mainly littoral and sublittoral 

depths. 

ECOLOGY.—Endeis spinosa is a frequent inhabitant of floating Sargassum in the Caribbean and 

elsewhere. It is also common in Thalassia grass 

that is supporting colonies of hydroids, bryozoans, 

and ascidians. 

Family NYMPHONIDAE 

Nymphon Fabricius, 1794 

Nymphon floridanum Hedgpeth, 1948 

Nymphon flondanum Hedgpeth, 1948:196-199, fig. 17 [longnecked form only].—Stock, 1955:215, fig. la [long-necked 

form only]; 1975:994-998, figs. 14-15.—Kraeuter, 1973: 

494.—Krapp and Kraeuter, 1976:336-337.—Child, 1979: 

37. 

MATERIAL EXAMINED.—Carrie Bow Cay: Tidal 

flats at low water, coll. K. Riitzler, 23 Apr 1974; 

16. Tidal flats, coll. M. L. Jones, 5 Apr 1976; 19. 

Halimeda and rubble from outer reef ridge in 18 

m, coll. B. Kensley, 7 Feb 1978; 1 juvenile. 

REMARKS.—These specimens are all the longnecked form of N. floridanum (sensu strictu) as 

defined by Stock (1975:994-998). In this extensive genus, this species (along with its look-alike, 

N. aemulum) is one of the few tropical members 

that is both littoral and sublittoral. It is known 

from Georgia through the Caribbean to as far 

south as French Guiana. It has been found in 

association with coral, algae, and sandy habitats. 

Family RHYNCHOTHORACIDAE 

Rhynchothorax Costa, 1861 

Rhynchothorax architectus Child, 1979 

Rhynchothorax architectus Child, 1979:68-72, figs. 23, 24a-g, 

25a-e. 

MATERIAL EXAMINED.—Carrie Bow Cay: Broken rock and rubble at tide line on ocean side, 

coll. J. Clark, 17 Jan 1976; 16. 

REMARKS.—This is the third capture locality 

for this variable species, the first two being the 

Caribbean and Pacific sides of the Isthmus of 

Panama. This specimen agrees well with male 

paratype specimens from Panama. It has low 

middorsal tubercles instead of the taller ones of 

the holotype. The species is known only from the 

intertidal. 

ECOLOGY.—This species probably came from 

the coral sand around or under rock rubble. The 

most common habitats of this genus, as reported 

in the literature, are interstitial, with the animals 

living between sand grains around or under rocks. 

The above capture probably substantiates this 

mode of living and suggests that coralline and 

coral sand are the primary substrates. 

Rhynchothorax crenatus, new species 

FIGURE 167 

MATERIAL EXAMINED.—Carrie Bow Cay: Coral 

sand and rubble in sand trough behind outer reef 

ridge in 27 m, coll. C. A. Child, 6 Feb 1978; 1 

subadult specimen, holotype (USNM 170996). 

DESCRIPTION.—Trunk compact, first 2 segment 

lines complete, third lacking. First 3 trunk segments with tall conical median dorsal tubercles 

almost as tall as ocular tubercle. Surface of entire 

animal except proboscis with minute scattered 

papillae. Dorsal and ventral trunk surfaces with 

pattern of lightly pigmented molt sutures similar 

to reticulations. Ocular tubercle tall, pointing 

about 30 degrees anteriorly, with 2 tiny lateral 

tubercles flanking a conical cap, a posterior triangular median tubercle and 2 thin short tubercles lateral and posterior to triangular tubercle. 

Conical cap with single posterior seta. Eyes large 

with medium dark pigment. Abdomen extending 

to tip of second coxae of fourth legs, cylindrical, 

and with proximal and distal constrictions, armed 

with 3 dorsodistal setae. 

Proboscis cylindrical-conical, with 2 dorsolateral bulges flanking a tall spike-like tubercle in 

median line halfway along proboscis. Mouth with 

2 laterally flattened antimeres confining it to a 

vertical slit. 

Palps 4-segmented, arising from rather long 
NUMBER 12 375 

FIGURE 167—Rhynchothorax crenatus, new species (holotype): a, trunk; b, trunk, lateral; c, palp; d, 

terminal segments of palp, dorsally, with enlargement of endal spine; e, third leg. 

trunk tubercle bases lateral to and closely set 

against proboscis. First segment very short, half 

as long as its diameter. Second segment longest, 

with small triangular dorsodistal tubercle armed 

with 2-3 setae and tiny ventrodistal tubercle 

armed with single seta. Third segment slightly 

less than half as long as second, with narrow 

median dorsal tubercle armed with seta, and 

small ventrodistal bulge armed with 2-3 setae. 

Terminal segment round in lateral aspect, armed 

with many setae dorsally, distally, and ventrally. 

Third palp segment armed on endal surface with 

single large spine bearing denticulations on posterior surface only. Spine slightly longer than 

diameter of segment. 

Chelifores entirely lacking. 

Oviger incomplete (?) or vestigial (?) in holotype, only small unsegmented bud present. 

Legs moderately thin, armed with a few short 

setae and a very long single seta on first and 

second tibiae. Femorae lacking long setae. Femur 

the longest segment; first and second tibiae each 

shorter than preceeding segment. Major leg segments of anterior 4 legs slightly longer than posterior four. Tarsus with single ventral spine and 

seta. Propodus without heel, moderately curved, 

armed with 5-6 sole spines. Terminal segments 

of anterior 4 legs armed with 4-6 tarsus setae and 

up to 12 sole spines. Claw robust, less than half 

propodus length. Auxiliary claws lacking. 

DISTRIBUTION.—Known only from the type-locality, Carrie Bow Cay, Belize, in a depth of 27 

meters. 

ETYMOLOGY.—From Latin, meaning notched, 
376 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

in reference to the appearance created by the 

median trunk, ocular, and proboscis tubercles in 

lateral view. 

MEASUREMENTS (mm).—Trunk length (tip of 

ocular tubercle to tip of fourth lateral processes), 

0.65; trunk width (across first lateral processes), 

0.42; abdomen length, 0.14; proboscis length, 

0.35; third leg, coxa 1, 0.1, coxa 2, 0.08, coxa 3, 

0.07, femur, 0.23, tibia 1, 0.2, tibia 2, 0.17, tarsus, 

0.05, propodus, 0.21, claw, 0.09. 

REMARKS.—This subadult specimen (sex pores 

not evident) has a unique set of characters although it superficially resembles Rhynchothorax unicornis Fage and Stock from the Cape Verde Islands. The trunk of R. crenatus is thinner, with 

more widely spaced lateral processes. None of the 

appendage tubercles are nearly as long or thin, 

the ocular tubercle is very different with its cone 

and posterior tubercles, and the median trunk 

tubercles are simple cones instead of complex 

tuberculate and papillose tubercles. 

On the other hand, there are several striking 

similarities between the two species. Both share a 

single mid-dorsal proboscis tubercle, both have 

similar propodus configuration except for the 

strong distal sole spine of Rhynchothorax unicornis, 

and both have similar palps although the tall 

tubercle of the second segment and the tiny terminal segment are not present on R. crenatus. Both 

species lack auxiliary claws and have lateral process and coxa 1 tubercles in the same places 

although their sizes are very different. Finally, 

both species have small lateral tubercles placed 

behind the ocular tubercle and rudimentary ovigers (signifying that both are females?) of a button-like appearance. 

The two species are similar enough perhaps to 

form a geminate pair, separated by the Atlantic 

Ocean and Caribbean Sea. Probably, each is a 

valid species, but the discovery of a specimen 

intermediate between the two, if such exists, 

would invalidate any suggestion that these examples represent speciation by geographic isolation due to plate tectonics. Although the male 

ovigers in Rhynchothorax species are much alike, it 

might be worthwhile to study the character of 

male ovigers in these two species when more 

specimens are collected. 

Thompson (1909:535) first proposed a separate 

family for this genus and I concur that its combination of characters fit none of the currently 

accepted genera with which it has been placed. 

Conclusions 

The pycnogonid fauna of Carrie Bow Cay and 

its vicinity on the Belizean barrier reef is extremely rich. Extensive collecting during this 

study has produced approximately 300 specimens 

from mainly littoral habitats. Thirty-one identified species in 14 genera are represented, including four new species. The habitat diversity on the 

reefs makes it probable that collecting in nearby 

unsampled areas will further increase this number. For instance, two samples taken from a single 

mangrove habitat on different days contained no 

less than 10 species of pycnogonids, and five small 

samples of rubble and algae from depths of 18 

and 27 meters on the outer reef yielded nine 

species. 

Algae appear to be a preferred habitat but 

seldom is an alga associated with just one species 

of pycnogonid. Sargassum is regularly colonized by 

Anoplodactylus maritimus, Endeis spinosa, Tanystylum 

tubirostrum, Numphopsis duodorsospinosum, and occasionally other species. All these species, however, 

are also found in other habitats, such as rubble, 

other algae, sea grass, coral, and mangrove roots. 

Without an analysis of gut content there is no 

proof that pycnogonids ingest algae. More likely, 

they eat the soft parts of animals living on the 

algae or on rubble having associated algal growth. 

Specimens of Achelia sawayai, for example, were 

captured on rubble with algae supporting sessile 

organisms and were not found on "clean" algae 

collected at the same time. Pycnogonids with 

green intestinal diverticula {Anoplodactylus pectinus 

and A. portus) possibly form the second step in the 

food chain by ingesting the tissues of animals that 

feed primarily on green algae. Pycnogonids move 

very slowly so that it is almost impossible for 

them to feed on mobile fauna. A stand of foliose 
NUMBER 12 377 

TABLE 27.—Geographical distribution of Belizean pycnogonid species 

Species 

Achelia sawayai 

Ammothella appendiculata 

A. exornata 

A. marcusi 

A. rugulosa 

Ascorhynchus latipes 

A. serratus 

Eurycyde raphiaster 

Hedgpethius mamillatus 

H. tridentatus 

Nymphopsis duodorsospinosa 

Tanystylum birkelandi 

T. tubirostrum 

Callipallene belizae 

C emaciata 

Parapallene bermudensis 

Pigrogromitus timsanus 

Anoplodactylus bahamensis 

A. batangensis 

A. evelinae 

A. imswe 

A. jonesi 

A. maritimus 

A. monotrema 

A. multiclavus 

A. pectinus 

A. portus 

Endeis spinosa 

Nymphon floridanum 

Rhynchothorax architectus 

R. crenatus 

Pantropical 

X 

X 

X 

u is 

1 CU 

rri Middle Ameria 

X 

X 

X 

X 

X 

X 

-o Galapagos Islan 

X 

X 

c 

ed 

<u 

_D Panama (Caribl 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

Belize 

X 

X 

X 

X 

X 

X 

X? 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

c Other Caribbea 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

o 

G Western N At la 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

— ^ 03 Guianas and Br 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

Q. 

o ( H 

3 

W Africa and E 

X 

X 

X 

X 

X 

X 

X 

X 

Indian Ocean 

X 

X 

X 

green algae can support an enormous number of 

attached organisms, many of which are known to 

be food for pycnogonids. Among these organisms 

are hydroids—long a classic example of pycnogonid food—sponges, tube worms, ascidians, and 

anemones. Since Pycnogonids have been captured on living corals, they probably also feed on 

these cnidarians. 

Most of the Belizean species have a littoral 

depth distribution with the exception of Parapallene bermudensis which has only been captured 

within the narrow limits of 27 to 33 meters. This 

rarely collected species may have escaped owing 

to inefficient collecting techniques. More sampling on reefs at this depth range may provide 

many other species previously thought to be rare. 
378 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

The geographical distribution of pycnogonids 

from Belize is summarized in Table 27. Most of 

the species also occur in the eastern Caribbean 

and some are present along the Caribbean shores 

of Panama. Three species were found in the 

Caribbean for the first time: Hedgpethius tridentatus, previously known from Florida; Anoplodactylus 

bahamensis, Bahamas; and Parapallene bermudensis, 

Bermuda. The four new species presented here 

are known only from Belize. Nineteen species are 

known from the western North Atlantic beyond 

the Caribbean, and at least 10 range to the 

Guianas and Brazil. There are eight known amphi-Atlantic species, two of which are also known 

from the Indian Ocean. The Pacific distribution 

of Belize species is much more restricted, as is to 

be expected. The isthmian barrier has permitted 

colonization of only six species on the Pacific 

shores of Central America. Two species are also 

known from the Galapagos Islands and one of 

these has been captured in other areas of the 

Eastern Pacific. Three additional species are pantropical in distribution and are known from most 

of the above localities. 

Literature Cited 

Arnaud, F. 

1972. Pycnogonides des recifs coralliens de Madagascar, 

3: Famille des Callipallenidae. Telhys, supplement, 

3:157-164, 6 figures. 

1973. Pycnogonides des recifs corraliens de Madagascar, 

4: Colossendeidae, Phoxichilidiidae et Endeidae. 

Telhys, 4(4):953-960, 8 figures. 

Bourdillon, A. 

1955. Les Pycnogonides de la croisiere 1951 du "President Theodore Tissier." Revue des Travaus de 

L'Inslitute des Peches Man times, 19 (4): 581-609, 2 

figures, 3 plates. 

Caiman, W. T. 

1927. Zoological Results of the Cambridge Expedition 

to the Suez Canal: Report on the Pycnogonida. 

Transactions of the Zoological Society of London, 22(3): 

403-410, figures 102-104. 

Carpenter, G. H. 

1892. Pycnogonida: Reports on the Zoological Collections Made in Torres Straits by Prof. A. C. Haddon, 1888-89. Scientific Proceedings of the Royal Dublin 

Society, 7(5): 552-558, 1 plate. 

Child, C. A. 

1974. Hedgpethius tridentatus, a New Genus and New Species, and Other Pycnogonida from Key West, 

Florida, U.S.A. Proceedings of the Biological Society of 

Washington, 87(43):493-500, 2 figures. 

1977. Four new species of Anoplodactylus (Pycnogonida) 

from the Western North Atlantic. Proceedings of the 

Biological Society of Washington, 90(3):584-596, 4 

figures. 

1978. Gynandromorphs of the Pycnogonid Anoplodactylus 

portus. Zoological Journal of the Linnean Society, 63(1-

2): 133-144, 4 figures. 

1979. The Shallow-Water Pycnogonida of the Isthmus 

of Panama and the Coasts of Middle America. 

Smithsonian Contributions to Zoology, 293:1-86, 25 figures. 

Child, C. A., and J. W. Hedgpeth 

1971. Pycnogonida of the Galapagos Islands. Journal of 

Natural History, 5:609-634, 8 figures. 

Cole, L. J. 

1906. A New Pycnogonid from the Bahamas. The American Naturalist, 40(471):217-227, 2 plates. 

Costa, O. G. 

1861. Microdoride mediterranea, o descrizione de'poco ben conosciuti od affatto ignoti viventi minuti e microscopici del 

Mediterraneo, etc. Volume 1, xviii + 80 pages, 12 

plates. 

De Haro, A. 

1966. Picnogonidos del alga parda Halopteris scoparia (L.) 

de las islas Medas (Gerona). Boletin Real Sociedad 

Espanola de Historia Natural (Biologica), 64(1):5—14, 

5 figures. 

1967. Picnogonidos de las posidonias de Blanes (Gerona). Publicaciones de la Instituto de Biologica Aplicada, 

Barcelona, 43:103-124, 10 figures. 

Dohrn, A. 

1881. Die Pantopoden des Golfes von Neapel und der 

angrenzenden Meeresabschnitte. Monographic der 

Fauna und Flora des Golfes von Neapel, 3:1-252, plates 

1-18. 

Fabricius, J. C. 

1794. Enlomologia Systematica Emendata el aucta, 4:416-417. 

Hafniae. 

Fage, L. 

1952. Sur quelques Pycnogonides de Dakar. Bulletin du 
NUMBER 12 379 

Museum d'Histoire Naturelle (Paris), series 2, 24(6): 

530-533, 1 figure. 

Fage, L., and J. H. Stock 

1966. Pycnogonides: Resultats scientifiques du campagne de la Calypso aux iles de Cap Vert (1959). 

Annates de ITnstitut Oce'anographique de Monaco, 44(7): 

315-327, figures 1-4. 

Flynn, T. T. 

1929. Pycnogonida from the Queensland Coast. Memoirs 

of the Queensland Museum, 9(3): 252-260, 9 figures. 

Giltay, L. 

1934. A New Pycnogonid from Bermuda. Bulletin du 

Muse'e Royal d'Histoire Naturelle de Belgique, 10(42): 

1-3, 5 figures. 

Haswell, W. A. 

1885. On the Pycnogonida of the Australian Coast, with 

Descriptions of New Species. Proceedings of the Linnean Society of New South Wales, 9(4): 1021-1033, 

plates 54-57. 

Hedgpeth, J. W. 

1947. On the Evolutionary Significance of the Pycnogonida. Smithsonian Miscellaneous Collections, 

106(18): 1-54, 16 figures, 1 plate. 

1948. The Pycnogonida of the Western North Atlantic 

and the Caribbean. Proceedings of the United States 

National Museum, 97:157-342, figures 4-53, 3 

charts. 

1954. Pycnogonida. In Gulf of Mexico: Its Origin, Waters, and Marine Life. United Stales Fish and Wildlife 

Fishery Bulletin, 89:425-427, figure 69, table 1. 

Heifer, H. 

1938. Einige neue Pantopoden aus der Sammlung des 

Zoologischen Museums in Berlin. Sitzungsbenchte 

der Gesellschaft Naturforschender Freunde zu Berlin, 

1937(2): 162-185, 11 figures. 

Hilton, W. A. 

1942a. Pycnogonids from Allan Hancock Expeditions. 

Allan Hancock Pacific Expeditions, 5(9):277-338, 

plates 35-48. 

1942b. Pycnogonids from Hawaii. Occasional Papers, Bernice 

P. Bishop Museum, 17(3):43-55, 10 figures. 

Hodge, G. 

1864. List of the British Pycnogonidea, with Descriptions 

of Several New Species. Annals and Magazine of 

Natural History, series 3, 13:113-117, plates 12-13. 

Hodgson, T. V. 

1914. Preliminary Report on the Pycnogonida of the 

German Southpolar Expedition 1901-3. Zoologischer Anzeiger, 45:158-165. 

1915. Preliminary Report on the Pycnogonida of the 

German Southpolar Expedition 1901-3. Annals and 

Magazine of Natural History, series 8, 15:141-9. 

1927. Die Pycnogoniden der Deutschen Siidpolar-Expedition 1901-1903. In Deutsche Sixdpolar-Expedition 

1901-1903, 19 (Zoologie ll):305-358, 17 figures. 

Kraeuter, J. N. 

1973. Pycnogonida from Georgia, U.S.A. Journal of Natural History, 7:493-498. 

Krapp, F. 

1973. Pycnogonida from Pantelleria and Catania, Sicily. 

Beaufortia, 21(277):55-74, 7 figures. 

Krapp, F., and J. N. Kraeuter 

1976. Additions to the Pycnogonida of Georgia. Bonner 

Zoologische Beilrages, 27(3/4): 336-346, 3 figures. 

Lebour, M. V. 

1949. Two New Pycnogonids from Bermuda. Proceedings 

of the Zoological Society of London, 118(4):929-932, 3 

figures. 

Lipkin, Y., and U. Safriel 

1971. Intertidal Zonation on Rocky Shores at Mikhmoret (Mediterranean, Israel). Journal of Ecology, 

59(1): 1-30, 7 figures, 3 photos, 1 table. 

Loman, J.C.C. 

1912. Note preliminaire sur les "Podosomata" (Pycnogonides) du Musee Oceanographique de Monaco. 

Bulletin de L'Institut Oceanographique de Monaco, 238: 

1-14, 9 figures. 

Marcus, E. 

1940. Os Pantopoda brasileiros e os demais sul-americanos. Boletim Faculdade de Filosofia, Ciencias e Letras 

da Universidade de Sao Paulo, 19 (Zoologia) 4:3-179, 

17 plates. 

Miers, E. J. 

1879. Crustacea (and Pycnogonida) of the Transit of 

Venus Expedition, Kerguelen Island. Philosophical 

Transactions of the Royal Society of London, 168:200-

214, plate 11. 

Montagu, G. 

1808. Description of Several Marine Animals Found on 

the South Coast of Devonshire. Transactions of the 

Linnean Society of London, 9:81-113, 5 plates. 

Philippi, A. 

1843. Uber die neapolitanischen Pycnogoniden. Archiv 

fiir Naturgeschichte, series 1, 9:175-182, pl. 9: figs. 

1-3. 

Sars, G. O. 

1877. Podromus descriptionis Crustaceorum et Pycnogonidarum, quae in expeditione norvegica anno 

1876 observavit. Archiv for Mathematik og Naturvidenskab, Krisliania, 2:237-271. 

Schiodte, J. C. 

1857. Udsigt over Gr0nlands Land Ferskvands og 

Strandbreds-Arthropoder. In H. Rink, Gr0nland 

Geografisk og Statistisk Beskrevet, 2(3):50-74. Kj0-

benhavn. 

Soyer, J. 

1966. Sur quelques Pycnogonides du Golfe de Genes. 

Doriana, 4(174): 1-5. 

Stock, J. H. 

1951. Pantopoda: Resultats Scientifiques des Croisieres 
380 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

du Navire-Ecole Beige "Mercator," 5. Memoires 

Institut Royal des Sciences Naturelles de Belgique, series 

2, 43(5): 1-23, 24 figures. 

1952a. Revision of the European Representatives of the 

Genus Callipallene Flynn, 1929. Beaufortia, 13:1-14, 

27 figures. 

1952b. The pycnogonids of the Lagoon of Venice. Bollettino Della Societa Veneziana di Storia Naturale, 6(2): 

179-186, 3 figures. 

1953. Contribution to the Knowledge of the Pycnogonid 

Fauna of the East Indian Archipelago. Temminckia, 

9:276-313, 18 figures. 

1954. Four New Tanystylum Species, and Other Pycnogonida from the West Indies. Studies on the Fauna of 

Curacao and other Caribbean Islands, 5(24): 115-129, 

figures 24-29. 

1955. Pycnogonida from the West Indies, Central America, and the Pacific Coast of North America. Videnskabelige Meddeleser fra Dansk Naturhistorisk Forening, Kobenhavn, 117:209-266, 26 figures. 

1957. Pantopoden aus dem Zoologischen Museum 

Hamburg. Mittheilungen aus dem Hamburg Zoologischen Museum Institut, 55:81-106, 20 figures. 

1962. Some Turkish Pycnogonid Records. Entomologische 

Berichten, series 22, 1(11):218-219. 

1968a. Pycnogonida Collected by the Galathea and Anton 

Bruun in the Indian and Pacific Oceans. Videnskabelige Meddeleser fra Dansk Naturhistorisk Forening, 

Kobenhavn, 131:7-65, 22 figures. 

1968b. Faune marine des Pyrenees Orientales. Vie et Mi1974. 

1975. 

1979. 

lieu, series A (Biologie Marine), 19(1-A), supplement 6:5-36, 30 figures. 

Medio- and Infralittoral Pycnogonida Collected 

during the I.I.O.E. near the Landbase on NossiBe, Madagascar. Bulletin of the Zoological Museum, 

University of Amsterdam, 4(3): 11-22, 4 figures. 

Pycnogonida from the Continental Shelf, Slope, 

and Deep Sea of the Tropical Atlantic and East 

Pacific: Biological Results of the University of 

Miami Deep-Sea Expeditions, 108. Bulletin of Marine Science, 24(4) :957-1092, 59 figures. 

Pycnogonida from the Mediolittoral and Infralittoral Zones in the Tropical Western Atlantic. Studies on the Fauna of Curacao and other Caribbean Islands, 

59(184): 1-19, 9 figures. 

Thompson, D'A. W. 

1909. Pycnogonida. In S. F. Harmer and B. E. Shipley, 

editors, The Cambridge Natural History, 4:501-542, 

figures 262-287. London: Macmillan. 

Turpaeva, E. P. 

1973. [Pantopoda from the Northeast Pacific Ocean.] 

Trudy Instituta Okeanologii P. P. Shirshov, Academia 

Nauk SSSR, 91:178-191, 2 figures. [In Russian.] 

Verrill, A. E. 

1900. Additions to the Crustacea and Pycnogonida of 

the Bermudas. Transactions of the Connecticut Academy 

of Arts and Sciences, 10(15):580-582, figures 2-4, 

plate 70: figures 9, 10. 

Wilson, E. B. 

1878. Descriptions of Two New Genera of Pycnogonida. 

American Journal of Science and Arts, 15:200-203. 
Pontomyia Edwards (Diptera: Chironomidae), 

a Member of the Coral Reef Community 

at Carrie Bow Cay, Belize 

Gemot Bretschko 

ABSTRACT 

The occurrence of the genus Pontomyia (Chironomidae, Tanytarsini) in the Caribbean Sea is 

reported for the first time. Larvae were found 

around Carrie Bow Cay, Belize, in the lagoon, on 

the reef flat, on the fore reef and along the forereef slope to a depth of 25-30 m (maximum 

sampling depth). Distribution of larvae indicates 

that Pontomyia is a true member of the off-shore 

benthic community. Pupal skins and females 

closely resembling those of P. natans Edwards, 

1926, were common in plankton samples. Species 

identification is not possible, however, because no 

males were found. 

Introduction 

Previous studies have reported species of the 

genus Pontomyia from only Pacific locations. It was 

first found in Samoa by Buxton (1926), whose 

collection was described by Edwards (1926). 

Characteristic features of the genus are larvae 

that morphologically resemble normal Tanytarsini larvae, a highly specialized adult male, and 

a completely reduced female that lacks all appendices on head and thorax except inoperative 

stumps of the second and third pairs of legs. 

Buxton (1926) reported that the species he studied, P. natans, spends its entire life-cycle under 

water, that males swim with their legs, and that 

females do not leave their pupal housings. 

Gemot Bretschko, Biologische Station Lunz, A-3293 Lunz am See, 

Austria. 

Another species, Pontomyia pacifica, was described from Japan by Tokunaga (1932), whose 

detailed field observations and laboratory experiments revealed that males of P. pacifica are unable to swim. They are true surface gliders propelled by their wing activity. Like the males, 

females emerge on the water surface, where they 

await copulation with a male passing by. 

Recently, Hashimoto (1959) showed that the 

biology of Pontomyia natans is very similar to that 

of P. pacifica in that adults are confined to the 

surface and are unable to copulate while submerged. Flying males of P. natans have been reported from the Red Sea and from northeast 

Australia (Thienemann, 1974). 

Larvae of Pontomyia have been found in association with various algae and in sandy mud below 

plant beds (Tokunaga, 1932; Hashimoto, 1959). 

All marine chironomid species are reported to be 

restricted to the intertidal zone; only P. natans has 

been found below the extreme low water level 

(Hashimoto, 1976), but always near the shore 

(Hashimoto, 1962). This author concluded that 

"in site of the excellent adaptation to the open 

sea, the territories of Clunio and Pontomyia are 

confined to the coast" (1962:241). 

ACKNOWLEDGMENTS.—I am grateful to F. Reiss 

(Munich) for stimulating discussions, to J. D. 

Ferraris for scanning all plankton samples and 

for counting the chironomids, to A. Cohen for 

providing additional material, and to W. R. 

Brown for operating the scanning electron microscope. 

381 
382 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Study Area and Methods 

Sampling for this study took place in May 1977 

around Carrie Bow Cay (16°48'N, 88°05'W) a 

small island on the Caribbean barrier reef off 

Belize (Riitzler and Macintyre, herein: 9) Benthic samples (Figure 168) were taken in the 

Thalassia stands between patch reefs in the lagoon, and from sand substrates on the fore reef, 

including sand trough and fore-reef slope, to a 

depth of about 25-30 meters. All samples were 

qualitative; two to three liters of sediments were 

brought to the laboratory in a plastic bag and 

subsequently were well mixed with sea water in 

a bucket. After a few hours, the bottom layers in 

the bucket became anoxic, and chironomid larvae 

swam to the surface and were collected. Chironomid larvae were also present in four semiquantitative samples taken on the reef flat by K. 

Riitzler in April-May 1974. 

Seventy-one quantitative plankton tows (250 

jum mesh size, water depth of 0.-0.5 and 2.5-3.0 

m) were taken by J. D. Ferraris (herein: 143) 

parallel to the reef, in the lagoon and outside the 

fore-reef slope, and during different lunar and 

tidal phases, in April-May 1976 (Figure 168). 

Nets were towed horizontal distances that varied 

from 200 to 552 m, and the volume of water 

filtered per tow ranged from 13.2 to 36.4 m . 

Results and Discussion 

Out of 13 large benthic samples, 11 contained 

chironomid larvae and a few females (Figure 

169). Chironomids were absent only in one sample of coarse sand from inside of a patch reef and 

in one sample of detached Thalassia seagrass 

leaves. The positive samples included muddy 

sands—with and without Thalassia—from the lagoon (2-5 m deep), coral rubble from the reef flat 

(0.5 m), and fine sands from the fore reef (3-23 

m) and along the reef drop-off (12-30 m) Figure 

168). Although the abundance of larvae is rather 

low everywhere, it is slightly higher in the lagoon. 

In the semiquantitative reef-flat samples densities 

range from about 10 to 150 larvae/m and are 

comparable to plankton samples having maximum counts of 2.2 pupal skins/m3

 in the lagoon 

meters 

Sponges 

FIGURE 168.—Transect across Belizean barrier reef near Carrie Bow Cay indicating sample 

locations (circle = plankton tow, arrow = benthic sample). 
NUMBER 12 383 

WBHBum meMm 

lif e 

FIGURE 169.—Pontomyia sp., scanning electron micrographs: a, larva, fourth instar; b, female, 

vermiform body somewhat shrunk; c, front end of female, with eyes and first two pairs of legs 

largely reduced, mouthparts, wings, and last pair of legs absent (scales = 200 /xm). 

and 0.4/m3 along the drop-off. The wide distribution of Pontomyia sp. around Carrie Bow Cay 

and its occurrence in relatively deep waters 

clearly indicates—contrary to earlier reports— 

that this chironomid species is a member of the 

marine offshore benthic community. 

The occurrence in the samples of all developmental stages at the same time suggests continu-
384 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ous reproduction. Analysis of 273 specimens indicated the absence of males, which in such a 

large sample points to a parthenogenetic population—a characteristic previously unknown for 

members of the genus Pontomyia. 

Thirty-four plankton samples contained skins 

of chironomid pupae and a few partly macerated 

females. Considerably more pupal skins were present in planktonic samples from the lagoon than 

in samples from the outer fore-reef {t = 3.882, d.f 

= 34 for 0.0-0.5 m depth; and t = 4.772, d.f. = 33 

for 2.5-3.0 m depth, respectively). These results 

support the findings drawn from the qualitative 

benthic samples. A correlation with moon phases 

and tide could not be established. A distinct 

relationship, however, exists between the abundance of exuviae and the diurnal cycle: pupalskin density is significantly higher in samples 

taken between 1800 and 2400 h compared to 

samples taken at all other 6-hour intervals {t = 

3.803-4.786, d.f. = 29-37, for all samples taken 

together). Surface tows (0-0.5 m) yielded more 

chironomids at all times of day than did tows 

from depths of 2.5 to 3.0 m (lagoon: t = 1.883, 

d.f. = 37; off fore-reef slope: t = 4.999, d.f. = 30). 

The specific gravity of the pupal skins may well 

be the reason for these differences. 

Of the four species described for the genus 

Pontomyia, P. cottoni Womersley (1937) was 

thought to be a junior synonym of P. natans 

Edwards (Hashimoto, 1959). In a recently published paper (Cheng and Hashimoto, 1978), the 

species P. cottoni is listed, yet without further 

taxonomic comments. Morphological differences 

between P. cottoni and P. natans are still considered 

of doubtful value (Hashimoto, 1973) but from an 

ecological point of view P. cottoni could be separated as a dweller of rather shallow water and 

tide pools (H. Hashimoto, pers. comm., 1979). 

The remaining three species are P. natans Edwards 

(1926), P. pacifica Tokunaga (1932), and P. oceana 

Tokunaga (1964). Immature stages and females, 

in addition to the males, are described for P. 

pacifica and P. natans. The larvae described for P. 

natans may not belong to Pontomyia at all, but to 

some other species of the Tanytarsini group (Tokunaga, 1932). 

This is the first observation of a Pontomyia species outside the Indo-Pacific region. Confident 

specific identification of the present material (larvae, pupal skins, a few females) is not possible 

because chironomid taxonomy is based on the 

morphology of the adult male which has not yet 

been found in the Caribbean population. The 

presence of four joints on the female leg excludes 

Pontomyia oceana and P. pacifica, although the larval 

atennae resembles the latter species more than 

the remaining P. natans. If we are dealing with a 

new species it is probably most closely related to 

P. natans. The taxonomic status of P. cottoni might 

require further collection and reevaluation. 

Literature Cited 

Buxton, P. A. 

1926. The Colonisation of the Sea by Insects: With an 

Account of the Habits of Pontomyia, the Only 

Known Submarine Insect. Proceedings of the Zoological Society of London, 3:807-814. 

Cheng, L., and H. Hashimoto 

1978. The Marine Midge Pontomyia (Chironomidae) with 

a Description of Females of P. oceana Tokunaga. 

Systematic Entomology, 3:189-196. 

Edwards, F. W. 

1926. On Marine Chironomidae (Diptera): With Description of a New Genus and Four New Species 

from Samoa. Proceedings of the Zoological Society of 

London, 3:779-806. 

Hashimoto, H. 

1959. Notes on Pontomyia natans from Sado (Diptera, 

Chironomidae). Science Reports of the Tokyo Kyoiku 

Daigaku, section B, 9:57-64. 

1962. Ecological Significance of the Sexual Dimorphism 

in Marine Chironomids. Science Reports of the Tokyo 

Kyoiku Daigaku, section B, 10:221-252. 
NUMBER 12 385 

1973. Marine Chironomids from Australia, with Description of a New Species of the Genus Clunio 

(Diptera: Chironomidae). Bulletin of the Faculty of 

Education, Shizuoka University, Natural Science Series, 

24:1-17. 

1976. Non-biting Midges of Marine Habitats (Diptera: 

Chironomidae). In L. Cheng, editor, Marine Insects, 

pages 377-414. New York: American Elsevier Publishing Company. 

Thienemann, A. 

1974. Chironomus: Leben, Verbreitung und wirtschaftliche Bedeutung der Chironomiden. 834 pages. Stuttgart: E. 

Schweizerbart. 

Tokunaga, M. 

1932. Morphological and Biological Studies on a New 

Marine Chironomid Fly, Pontomyia pacifica, from 

Japan, Part I: Morphology and Taxonomy. Memoirs of the College of Agriculture, Kyoto Imperial University, 19:1-56. 

1964. Chironomidae. In Insects of Micronesia, 12(5):485-

628. Honolulu, Hawaii: Bernice P. Bishop Museum. 

Womersley, H. 

1937. A New Marine Chironomid from South Australia. 

Transactions of the Royal Society, South Australia, 61: 

102-103. 
Ophiuroidea (Echinodermata) 

from Carrie Bow Cay, Belize 

Frederick H. C. Hotchkiss 

ABSTRACT 

Thirty-six species of ophiuroids representing 16 

genera in eleven families occur at Carrie Bow 

Cay, Belize. Th e rank order of abundanc e of 

species in various parts of the reef complex is 

estimated on the basis of numerical representation in the collection. Th e four most common 

species in the lagoon amon g discarded conch 

shells are Ophiothrix oerstedi, Ophiactis savignyi, 

Ophiothrix angulata, an d Ophiolepis paucispina; on 

the reef flat are Ophioderma appressum, Ophiocoma 

echinata, Ophiolepis impressa, and Ophiocoma wendti; 

on the reef crest are Ophiocoma wendti, Ophiocoma 

paucigranulata, Ophiocoma pumila and Ophiocoma 

echinata; on the fore reef including the accessible 

part of the fore-reef slope are Ophiocoma pumila, 

Ophiothrix angulata, Ophioderma rubicundum, and 

Ophiurochaeta littoralis. Fifteen species are reported 

for the first time from Belize, including a new 

species of Ophioderma. All specimens of Ophiostigma sp. are six arme d and fissiparous; they 

almost certainly represent a new species different 

from 0. isacanthum. Th e occurrence of young in 

the bursae of Ophiurochaeta littoralis and Ophiolepis 

paucispina indicates but does not prove viviparity. 

A small amphiuri d that clings mout h to mout h 

to an Ophiopsila riisei has three arms aligned and 

in contact with those of its host. Th e blue-purple 

color of man y preserved specimens of Ophiothrix 

angulata and 0. oerstedi is an artifact, the color in 

life usually being shades of brown. 

Introduction 

This report on the ophiuroids of Carrie Bow 

Cay is based on field work conducted 21 April 

Frederick H. C. Hotchkiss, Department of Biology, Moravian College, 

Bethlehem, Pa. Department of Geology and Paleontology, Academy of 

Natural Sciences of Philadelphia, Pa. Mailing address: Sherry Road, 

R.D. 3 Box 77, Harvard, Mass. 01451 

through 9 Ma y 1974. Carrie Bow Cay is a small 

island situated 22 km southeast of Dangriga 

(Stann Creek) (approximately 16°48'N, 88° -

05'W), on the barrier reef that extends along the 

coast of Belize (formerly British Honduras). An 

extensive collection of ophiuroids was mad e from 

various habitats and depths for the purpose of 

inventorying the species composition and distribution. Th e principal collecting localities categorized by major physiographic features of the 

reef complex are listed below. Detailed descriptions of these habitats are given by Riitzler and 

Macintyre (herein: 9). 

Thirty-six species in 16 genera and 11 families 

of ophiuroids are recognized in this report, including two not fully identified and one tha t is a 

new species of Ophioderma. All the species collected 

by Devaney (1974) at Turneffe Islands, Lighthouse Reef, and Glover's Reef are represented at 

Carrie Bow Cay except, perhaps, his Ophioderma 

sp., which I could not recognize from his description. Fifteen of the species are reported from 

Belize for the first time, although they have been 

known from other West Indian localities. Almost 

all species were photographed in color while alive, 

in order to document colors that are lost or 

changed with the fixatives used. 

ACKNOWLEDGMENTS.—My participation in this 

project was mad e possible by a Smithsonian Postdoctoral Fellowship (1973-1974). Th e field party 

consisted of K. Riitzler, E. Kirsteuer, K. Sandved, 

B. Spracklin, and myself. K. Sandved took the 

color photographs in the field and collected man y 

ophiuroids. Th e completion of this manuscript 

was facilitated by the assistance of J. Jonigian, 

M. Roberson, T. Burrows, J. Arguelles, E. Ball, 

D. Dejianne , A. Pruzan, and by a Short-Term 

387 
388 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Visit Program award from the Office of Academic 

Studies, Smithsonian Institution. 

Methods 

Ophiuroids were collected at 13 stations by 

hand using snorkel or SCUBA, according to 

depth. In most cases specimens were found hidden under coral, coral rock, or dead shells, or in 

nooks and crannies of living coral or sponges. 

Bulk samples of substrate for sorting ashore were 

collected from stations 1, 10, 12, and 13. Live 

specimens were photographed with 35 mm color 

slide film and electronic flash illumination in 

temporary aquaria or in trays. Specimens were 

relaxed in magnesium chloride solution judged 

isotonic with sea water by optical refractometer 

readings. Specimens were fixed in 10% formalin 

in sea water (4% formaldehyde solution) neutralized and buffered with 20 g/1 hexamethylenetetramine and transferred to 70% ethanol on return 

to the National Museum of Natural History. 

Most of the material has been kept in alcohol for 

future dissection and histological analysis, but 

some specimens were dried for identification purposes. Photographs and voucher specimens have 

been deposited in the National Museum of Natural History, Smithsonian Institution, under the 

catalog numbers of the United States National 

Museum (USNM), uncataloged material designated CBC. 

Station List 

(Station numbers keyed to Figure 170) 

Lagoon 

1. Northwest side of cay, foot of pier; among discarded 

conch shells, 0.5 m. 

2. Southwest end of cay; rubble area between sand and 

Thalassia, 0.5 m. 

Back Reef 

3. Main reef transect; patch-reef zone, 1 m. 

Reef Flat 

4. Northeast end of cay; Penicillus zone, 0.5 m. 

5. Southeast end of cay; Penicillus zone, 0.5 m. 

6. Coral-head/coral-pavement zone, 1 m. 

7. Inner reef-crest zone, 0.5 m. 

Reef Crest 

8. Coral-rubble berm (storm ridge), intertidal to 0.5 m 

(Plate 5: top left). 

9. Outer reef-crest zone, 0.5 m. 

Inner Fore Reef 

10. High-relief spur and groove zone, along the reef transect; bulk sample, 10-12 m. 

Outer Fore Reef 

11. Sand trough near transect; bottom of shoreward slope 

of outer ridge, 24 m. 

12. Outer ridge near transect; bulk sample, 12 m. 

13. Fore-reef slope near transect; bulk sample, 26 m; 

incidental 1973 samples, 15-28 m. 

Species List 

(* = new records for Belize) 

Superfamily OPHIUROIDEA 

Family GORGONOCEPHALIDAE 

* Astrophyton muricatum (Lamarck) 

Family OPHIOMYXIDAE 

Ophiomyxa flaccida (Say) 

Family OPHIACANTHIDAE 

*Ophiomelrella glabra (H. L. Clark) 

Family HEMIEURYALIDAE 

*Sigsbeia murrhina Lyman 

Family AMPHIURIDAE 

* Amphiura fibulala Koehler 

*A. stimpsonii Liitken 

* Axiognalhus squamatus (Delle Chiaje) 

*Ophiostigma sp. 

Family OPHIACTIDAE 

* Ophiactis algicola H. L. Clark 

Family OPHIOTRICHIDAE 

0. savignyi (Muller and Troschel) 

Ophiothrix (Acanlhophiothnx) suensom Liitken 

O. (Ophiothrix) angulata (Say) 

0. (Ophiothrix) lineala Lyman 

O. (Ophiothrix) oerstedi Liitken 

Family OPHIONEREIDAE 

Ophionereis reticulata (Say) 

*0. squamulosa Koehler 

Family OPHIOCOMIDAE 

Ophiocoma echinata (Lamark) 

0. paucigranulata Devaney 

0. pumila Liitken 

0. wendti Muller and Troschel 

* Ophiocomella ophiactoides (H. L. Clark) 

*Ophwpsila riisei Liitken 

Family OPHIODERMATIDAE 

*Ophioderma anilae, new species 

O. appressum (Say) 

O. brevicaudum Liitken 
NUMBER 12 389 

Sand/Rubble/ 

Pavement 

B Thalassia 

FIGURE 170.—Sketch map of collecting stations described in text (1, 2 = lagoon, 3 = back reef, 

4-7 = reef flat, 8, 9 = reef crest; 10 = inner fore reef, 11-13 = outer fore reef; A-A' = study 

transect). 

*0. brevispinum (Say) 

0. cinereum Muller and Troschel 

*0. guttatum Liitken 

O. phoenium H. L. Clark 

0. rubicundum Liitken 

*0. squamosissimum Liitken 

Ophioderma sp. 

Ophioderma sp. juv. 

Ophiurochaeta littoralis (Koehler) 

Family OPHIURIDAE 

Ophiolepis impressa Liitken 

0. paucispina (Say) 

Superfamily OPHIUROIDEA 

Family GORGONOCEPHALIDAE 

Astrophyton muricatum (Lamarck) 

Euryale muricatum Lamarck, 1816:538. 

Astrophyton muricatum.—Agassiz, 1835:172.—Muller and 

Troschel, 1842:122.—Liitken, 1859:258; 1869:68,106.— 

Lyman, 1865:192.—Ljungman, 1866:336.—Verrill, 1867: 

341; 1899a:84; 1899b: 381.—Drjderlein, 1911:7, 18, 53, 

108, pl. 5: fig 1; 1927:24, 25.—H. L. Clark, 1915:188; 

1919:56, 71; 1933:34, 42.—A. H. Clark, 1922:210.— 

Boone, 1933:103.—Engel, 1939:4.—Fontaine, 1953: 

200.—John and Clark, 1954:140.—Cherbonnier, 1959: 

261.—Fell, 1960:8.—Parslow and Clark, 1963:43.—Davis, 

1966:435.—Roa, 1967:283.—Fricke, 1968:1.—Tommasi, 

1970:11, pl. 2: fig. 3.—Godcharles and Jaap, 1973:56.— 

Zeiller, 1974:106 [color fig.]. 

Astrophyton costosum.—Lyman, 1865:192; 1882:256, 257, 266, 

271, 290, 313, 326, pl. 35: figs. 17-25.—Nutting, 1895: 

172.—Koehler, 1907:349. 

Astrophylum muricatum.—A. H. Clark, 1921:52; 1954:376. 

MATERIAL DEPOSITED.—USNM El 7735 (1 

spec); USNM E19478 (1 spec). 

DISTRIBUTION.—One specimen was secured 

from a coral sample taken in 10-12 m in the spur 

and groove zone of the inner fore reef along the 

main study transect (station 10); the other specimen was collected by B. Spracklin on a lagoon 

patch reef, 1 km WSW of Carrie Bow Cay, 4 m 

(24 May 1975). 

REMARKS.—The biology and living haits have 

been described by Davis (1966) and Fricke (1968). 

Family OPHIOMYXIDAE 

Ophiomyxa flaccida (Say) 

Ophiura flaccida Say, 1825:151. 

Ophiomyxa flaccida.—Liitken, 1859:79, 138.—Lyman, 1865: 
390 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

178.—Ljungman, 1866:327.—Verrill, 1867:329; 1899a: 

65; 1899b:366.—Rathbun, 1879:155.—Ives, 1889:178.— 

H. L. Clark, 1901:251; 1915:169; 1919:56, 71, pl. 1: figs. 

1,2; 1933:41.—Koehler, 1907:341; 1913:379; 1914:3, 132, 

162.—A. H. Clark, 1921:51; 1922:210; 1939a:446; 1939b: 

2; 1954:376.—Engel, 1939:4.—Fontaine, 1953:200.—Roa, 

1967:282, fig. 10.—Tommasi, 1970:13, pl. 2: figs. 4, 5.— 

Devaney, 1974:129.—Zeiller, 1974:106 [color fig.]. 

MATERIAL DEPOSITED.—USNM El 7759 (1 

spec); USNM E17779 (1 spec); USNM E17780 

(4 spec); USNM El7782 (1 spec); USNM 

El7857 (4 spec). 

DISTRIBUTION.—Found on the reef flat in the 

north Penicillus zone (station 4), in the coral-head/ 

coral-pavement zone (station 6), just inside the 

reef crest (station 7), on the reef crest in the rubble 

of the coral berm (station 8), and just seaward in 

0.5 m of water (station 9). Collected by bulk 

sample from the fore-reef slope (station 13). 

REMARKS.—Taken by Devaney (1974:129) at 

Lighthouse Reef and Glover's Reef. As noted by 

H. L. Clark (1933), this species occurs along shore 

in very shallow water to depths below 183 m (100 

fathoms). 

FIGURE \7\.—Ophiomilrella glabra, USNM El7679, dry: a, aboral; b, oral. (Scale = 10 mm. 
NUMBER 12 391 

Family OPHIACANTHIDAE 

Ophiomitrella glabra (H. L. Clark) 

FIGURE 171 

Op/nalcaea glabra H. L. Clark, 1901:249, pl. 15: figs. 1-4. 

Ophiomitrella glabra.—H. L. Clark, 1915:210, 1919:56, 71; 

1933:34, 43.—Parslow and Clark, 1963:26. 

MATERIAL DEPOSITED.—USNM "El7679 (1 

spec); USNM El7681 (1 spec). 

DISTRIBUTION.—One specimen was found on 

the reef crest in the rubble of the coral berm 

(station 8) by Kjell Sandved and one in a general 

sample taken from the fore-reef slope (station 13). 

REMARKS.—New to Belize, the species was previously known only from the unique holotype 

found at Playa de Ponce, Puerto Rico. 

Family HEMIEURYALIDAE 

Sigsbeia murrhina Lyman 

FIGURE 172 

Sigsbeia murrhina Lyman, 1878:234, pl. 3: figs. 55, 58; 1882: 

250, pl. 43: figs. 4-6; 1883:277; 1888:114, fig. 399.— 

Nutting, 1895:79.—Verrill, 1899a:72, pl. 2: figs. 1, la; 

1899b:365, 381, pl. 42: fig. 7.—Koehler, 1914:133, 135, 

163.—H. L. Clark, 1915:192; 1941:78.—A. H. Clark, 

1921:52; 1954:376. 

MATERIAL DEPOSITED.—UNSM El 7682 (3 

spec). 

DISTRIBUTION.—Six specimens were found in 

general samples from the fore-reef slope at depths 

from 16.0 to 27.5 m (station 13). 

Family AMPHIURIDAE 

Amphiura fibulata Koehler 

Amphiurafibulata Koehler, 1913:359; 1914:2, 56, 64, 65, pl. 7: 

figs. 3-5.—H. L. Clark, 1915:234; 1933:35, 45.—A. H. 

Clark, 1954:377.—Thomas, 1962a:638, fig. 4.—Tommasi, 

1965:3. 

Amphiura (Monamphiura) fibulata.—Tommasi, 1970:45. 

MATERIAL DEPOSITED.—USNM El 7762 (1 

spec); USNM El7793 (1 spec); USNM El7803 

(1 spec). 

FIGURE 172.—Sigsbeia murrhina, USNM El7682, live. (Scale 

= 5 mm.) 

DISTRIBUTION.—Three specimens were found 

among the discarded conch shells at the foot of 

the pier on the lagoon side of Carrie Bow Cay 

(station 1). 

Amphiura stimpsonii Liitken 

Amphiura stimpsonii Liitken, 1859:116.—Lyman, 1865:12; 

1875:4, pl. 5: fig. 66; 1882:124-143; 1883:252.—Ljungman, 1866:320; 1871:631.—Rathbun, 1879:154.—Marktanner, 1887:299.—Verrill, 1899b:376.—H. L. Clark, 

1901:247; 1915:232; 1919:56, 71; 1933:35, 45.—Koehler, 

1914:2, 59, 64, pl. 7: figs. 1, 2.—A. H. Clark, 1922:210; 

1954:377.—Mortensen, 1936:207.—Engel, 1939:4, 7.— 

Fontaine, 1953:200.—Thomas, 1962a:636, fig. 3.—Parslow and Clark, 1963:43. 

Amphiura vivipara H. L. Clark, 1918:268, pl. 1: figs. 1, 2; 1919: 

56, 71. 

Monamphiura stimpsonii.—Fell, 1962:11. 

Amphiura (Monamphiura) stimpsonii.—Tommasi, 1970:50, pl. 

24: figs. 51-52. 

MATERIAL DEPOSITED.—UNSM El 7763 (2 

spec); USNM El7765 (1 spec); USNM El7775 

(1 spec); USNM El7794 (4 spec); USNM 

E17796 (1 spec); USNM E17798 (1 spec); 

USNM El7801 (I spec); USNM El7802 (1 

spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); and in a 

general sample from the fore-reef slope (station 

13). 

REMARKS.—H. L. Clark (1918:268) showed 

that this species is viviparous. 

Axiognathus squamatus (Delia Chiaje) 

Ophiura elegans.—Leach, 1815:59 

Asterias squamata Delle Chiaje, 1829:74. 
392 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Ophiocoma neglecta.—Forbes, 1841:30 [text-fig.]. 

Ophiolepsis squamata.—Muller and Troschel, 1842:92. 

Amphiura squamata.—-Lyman, 1865:122; 1882:125, 136, 291, 

309, 310, 314, 321, 322, 324.—Koehler, 1907:305; 1913: 

356. 

Amphipholis elegans.—Ljungman, 1866:312. 

Amphipholis squamata. — Ljungman, 1871:633.—Verrill, 

1899b:312.—H. L. Clark, 1904:560, pl. 6: figs. 33, 34, pl. 

7: figs. 43, 44; 1914:105, pl. 2: figs. 1-6; 1915:242. 1933: 

36, 50.—Coe, 1912:81, pl. 17.—Koehler, 1914:2, 52, 66, 

156.—A. H. Clark, 1922:210; 1954:377.—Bernasconi, 

1926:146, pl. 3.—Engel, 1939:4, 8.—Fontaine, 1953: 

201.—Thomas, 1962a:662, fig. 13.—Parslow and Clark, 

1963:44.—Roa, 1967:286. 

Amphiura lenera Rathbun, 1879:154. 

Axiognathus squamatus.—Thomas, 1966:831.—Martin, 1968: 

65.—Tommasi, 1970:37, pl. 16: fig. 36.—Boffi, 1972: 

319.—Hendler, 1975:703. 

MATERIAL DEPOSITED.—USNM El7764 (12 

spec); USN M El7792 (2 spec). 

DISTRIBUTION.—Collected amon g the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1). 

REMARKS.—Th e development of this viviparous species has been studied by numerous authors. H. L. Clark's (1933:50) statement that very 

young specimens may have six arms and show 

evidence of autotomy is probably based on Koehler's (1913:356) report of a six-armed specimen 

from Tortuga s in which three arms are very short 

and in the process of regeneration; most likely 

either the specimen was misidentified or it regenerated a supernumerary arm following injury. 

Martin (1968) and Boffi (1972) have described 

aspects of the ecology of this species. 

Ophiostigma sp. 

FIGURE 173 

MATERIAL DEPOSITED.—USN M E l 7688 (1 

spec); USN M El768 9 (1 spec). 

DISTRIBUTION.—Collected amon g the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); in the spur 

and groove zone of the inner fore reef from a 

coral bulk sample taken in 10 to 12 m of water 

(station 10); and from the fore-reef slope in 26 m 

of water (station 13). 

REMARKS.—All specimens have six rays and 

most show evidence of recent fissiparity. Although 

Koehler (1913) and H. L. Clark (1933; 1942) 

reported six-rayed fissiparous specimens of Ophiostigma isacanthum, the present material strongly 

suggests that the fissiparous six-rayed specimens 

are specifically distinct from the nonfissiparous 

five-rayed specimens of 0. isacanthum and belong 

to an undescribed species. A. M. Clark (1967a) 

did not include Ophiostigma in her list of fissiparous ophiuroids, having previously suggested (in 

Parslow and Clark, 1963:37) that H. L. Clark's 

FIGURE 173.—Ophiostigma sp., USNM E17688, dry: a, aboral; b, oral. (Scale = 2 mm.) 
NUMBER 12 393 

(1942:377) Bermuda n fissiparous specimen of 0. 

isacanthum might instead be a specimen of Ophiocomella ophiactoides; that judgemen t now seems 

prematur e an d calls for reexamination of the 

material. Ophiostigma isacanthum has most recently 

been described in detail by Thoma s (1962a), but 

his material did not include six-rayed fissiparous 

specimens. 

Famil y OPHIACTIDAE 

Ophiactis algicola H. L. Clark 

FIGURE 174 

Ophiactis loncata.—H. L. Clark, 1901:246 [not Lyman, 1869: 

331]. 

Ophiactis plana.—H. L. Clark, 1915:264 Bermuda [not Lyman, 1869:330]. 

Ophiactis lymani.—H. L. Clark, 1918:303, pl. 4: figs. 5, 6; 

1919:57, 72.—Tommasi, 1965:5, 6; 1970:22, pl. 7: fig. 14. 

[Not Ljungman, 1871:629]. 

Ophiactis algicola H. L. Clark, 1933:38, 56; 1942:377. 

MATERIAL DEPOSITED.—USN M El 7647 (1 

spec). 

DISTRIBUTION.—One specimen collected from 

a general bulk sample from the fore-reef slope 

(station 13). 

REMARKS.—Th e species is fissiparous (H. L. 

Clark, 1933; but not listed by A. M. Clark, 1967a) 

and this specimen has three old and three regenerating arms. 

Ophiactis savignyi (Muller and Troschel) 

Ophiolepis savignyi Muller and Troschel, 1842:95. 

Ophiactis krebsu Liitken, 1856a: 12.—Lyman, 1865:111.— 

Verrill, 1868:366; 1899b:376.—Ljungman, 1871:627.— 

Rathbun, 1879:153.—H. L. Clark, 1901:246. 

Ophiactis virescens Liitken, 1856a:24. 

Ophiolepis sexradia Grube, 1857:343. 

Ophiactis reinhardlii Liitken, 1859:161, pl. 3: fig. 7a,b. 

Ophiactis savignyi.—Ljungman, 1866:323.—Koehler, 1907: 

311; 1913:351; 1914:2, 41, 154.—H. L. Clark, 1914:96, pl. 

1: figs. 1-19, pl. 3: figs. 12-14; 1915:265; 1918:305; 1919: 

57, 72; 1933:38, 59; 1942:377.—A. H. Clark, 1921:53; 

1922:211; 1939a:447, 1939b:3; 1954:377.—Engel, 1939:4, 

8.—Fontaine, 1953:201.—John and Clark, 1954:140.— 

Roa, 1967:288.—Tommasi, 1970:24, pl. 8: figs. 16, 17.— 

Boffi, 1972:319.—Devaney, 1974:129.—Pawson, 1978:11. 

Ophiactis incisa von Martens, 1870:248. 

Ophiactis brocki de Loriol, 1893:401. 

MATERIAL DEPOSITED.—USNM El 7732 (2 

spec); USN M El7797 (1 spec); USN M E17804 

(1 spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat in the south Penicillus zone (station 5); in the 

coral-head/coral-pavement zone (station 6); in 

the spur and groove zone of the inner fore reef 

from a coral bulk sample taken in 10 to 12 m of 

FIGURE 174.—Ophiactis algicola, USNM El 7647, dry: a, aboral; b, oral. (Scale = 1 mm.) 
394 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

water (station 10); from the sponge Siphonodictyon 

coralliphagum Riitzler at 24 m in the sand trough 

of the outer fore reef (station 11), and from the 

fore-reef slope in 26 m of water (station 13). 

REMARKS.—Two specimens from the fore-reef 

slope are five armed with the primary rosette 

perfectly intact, indicating that they never underwent autotomous fission. The relation of old 

to new arms in most specimens is 6 plus 0 or 3 

plus 3, but one from the lagoon has the relation 

2 plus 3. Previously reported from Glover's Reef 

(Devaney, 1974:129). 

Family OPHIOTRICHIDAE 

Ophiothrix (Acanthophiothrix) suensoni 

Liitken 

Ophiothrix suensonii Liitken, 1856a: 16; 1859:250, pl. 4: fig. 

2a-e.—Lyman, 1865:157; 1882:215, 218, 222, 223, 229, 

312, 315, 326.—Rathbun, 1879:153.—Ives, 1889:178.— 

Verrill, 1899a:21; 1899b:375— H. L. Clark, 1901:244; 

1915:279; 1918:313; 1919:57, 72; 1933:38, 62.—Koehler, 

1907:388; 1913:376; 1914:3, 121, 161.—A. H. Clark, 1921: 

54; 1922:211; 1939a:447; 1954:377.—Boone, 1933:111, pl. 

64.—Engel, 1939:4. 

Ophiothrix suensoni.—Ljungman, 1866:331; 1871:627.—Parslow and Clark, 1963:45.—Roa, 1967:289.—Tommasi, 

1971:7, figs. 21c, 22c, 23c. 

Ophiothrix (Acanthophiothrix) suensoni.—A. M. Clark, 1967b: 

638 643, 648, fig. 1 j.—Tommasi, 1970:62.—Devaney, 

1974:130. 

MATERIAL DEPOSITED.—USNM El 7734 (2 

spec); USNM E17787 (1 spec); USNM E17808 

(1 spec); USNM El7850 (2 spec). 

DISTRIBUTION.—Collected in the spur and 

groove zone of the inner fore reef from a coral 

bulk sample in 10 to 12 m of water (station 10), 

and in 24 m in the outer fore reef near the bottom 

of a trough among, and in crevices of, coral 

clumps (station 11). 

REMARKS.—Previously recorded from Belize 

(Koehler, 1914:122), Turneffe Islands, Lighthouse Reef, and Glover's Reef (Devaney, 1974: 

130). 

Ophiothrix ( Ophiothrix) angulata (Say) 

Ophiura angulata Say, 1825:145. 

Ophiothrix angulata.— Ayres, 1852:249.—Liitken, 1859:219.— 

Lyman, 1865:3, pl. 2: figs. 1-3; 1882:214, 216, 218, 219 

286, 287, 290, 312, 325.—Ljungman, 1866:331.—Ludwig 

1882:18.—Ives, 1889:178; 1890:332.—Verrill, 1899a: 18 

1899b:375.—H. L. Clark, 1901:244; 1914:111, pl. 2: figs 

7-12, pl. 3: figs. 1-10, 15-19; 1915:269; 1918:312; 1919 

57, 72; 1933:38, 60; 1941:86.—Koehler, 1907:330; 1913 

375; 1914:3, 118, 160.—A. H. Clark, 1921:53; 1922:211 

1939a:447; 1954:377.—Boone, 1933:110.—Engel, 1939:4 

8.—Fontaine, 1953:202.—Tabb and Manning, 1961 

566.—Parslow and Clark, 1963:45.—Roa, 1967:289.— 

Tommasi, 1971:7, figs. 21b, 22b, 23b.—Boffi, 1972:319.— 

Godcharles and Jaap, 1973:57. 

Ophiothrix violacea Muller and Troschel, 1842:115.—Liitken, 

1859, pl. 4: fig. la-g.—Verrill, 1868:366.—Rathbun, 

1879:152.—Koehler, 1907:339. 

Ophiothrix (Ophiothrix) angulata.—A. M. Clark, 1967b:646.— 

Tommasi, 1970:54.—Devaney, 1974:130. 

MATERIAL DEPOSITED.—USNM El 7805 (3 

spec); USNM El7809 (1 spec); USNM El7846 

(5 spec). 

DISTRIBUTION.—Found among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat in the south Penicillus zone among crevices of 

rocks and shells (station 5); in the spur and groove 

zone of the inner fore reef from a coral bulk 

sample in 10 to 12 m of water (station 10); from 

the outer ridge of the outer fore reef (station 12); 

and from the fore-reef slope to approximately 28 

m (station 13). 

REMARKS.—Previously recorded from Belize 

(Koehler, 1914:122) and from the Turneffe Islands, Lighthouse Reef, and Glover's Reef (Devaney, 1974:130). Shades of green and brown in 

life have turned blue in many of the preserved 

specimens. 

Ophiothrix ( Ophiothrix) lineata Lyman 

Ophiothrix lineata Lyman, 1860:201.—Ives, 1889:178.—Verrill, 1899b:375.—Koehler, 1907:334, pl. 13: figs. 46, 47 

1913:376; 1914:3, 120, 161.—H. L. Clark, 1915:273; 1918 

315; 1919:57, 72; 1933:38, 62.—A. H. Clark, 1922:211 

1954:377.—Engel, 1939:4.—Tommasi, 1970:54. 

Ophiothrix (Ophiothrix) tineata.—A. M. Clark, 1967b:647.— 

Devaney, 1974:131. 

Not Ophiothrix lineata.—A. H. Clark, 1921:54, reidentified in 

A. H. Clark, 1939a:449, as Ophiothrixplatyactis H. L. Clark, 

1939. 

MATERIAL DEPOSITED.—USNM El 7761 (1 

spec). 
NUMBER 12 395 

REMARKS.—One specimen was found in the 

temporary aquaria on Carrie Bow Cay. Previously reported from Turneffe Islands (Devaney, 

1974:131). 

Ophiothrix ( Ophiothrix) oerstedi Liitken 

Ophiothrix orstedu Liitken, 1856a: 15; 1859:149, pl. 4: fig. 

3a-e.—Ljungman, 1866:331; 1871:626.—Ives, 1889:178; 

1891:340.—Stearns, 1891:4.—Verrill, 1899a:20; 1899b: 

375.—H. L. Clark, 1901:244. 1915:276; 1918:313, 314; 

1919:57, 72, pl. 2; 1933:38, 63.—Koehler, 1907:336; 1913: 

376; 1914:3, 121, 161.—A. H. Clark, 1921:54; 1922:211; 

I939a:447; 1939b:3; 1954:377.—Engel, 1939:4, 8.—Roa, 

1967:290.—Tommasi, 1970:53. [Includes the spellings oersledii, oerstedi, and orstedi.] 

Ophiothrix (Ophiothrix) oerstedi.—A. M. Clark, 1967b:647.— 

Devaney, 1974:131. 

MATERIAL DEPOSITED.—USNM El 7739 (1 

spec); USNM El7843 (1 spec); UNSM El7844 

(1 spec); USNM El7849 (1 spec). 

DISTRIBUTION.—This species was widely taken. 

Collected among the discarded conch shells at the 

foot of the pier on the lagoon side of Carrie Bow 

Cay (station 1); under rocks in a rubble area 

between sand and Thalassia on the lagoon side of 

the south end of Carrie Bow Cay (station 2); on 

the reef flat in the coral-head/coral-pavement 

zone (station 6); just inside the reef crest (station 

7); on the reef crest in the rubble of the coral 

berm (station 8); in the spur and groove zone of 

the inner fore reef from a coral bulk sample taken 

in 10 to 12 m of water (station 10), and in 15 to 

28 m on the fore-reef slope (station 13). 

REMARKS.—Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

131). Shades of brown in life have turned blue in 

the preserved specimens. 

Family OPHIONEREIDAE 

Ophionereis reticulata (Say) 

Ophiura reticulata Say, 1825:148. 

Ophiolepis nereis Liitken, 1856a: 11; 1856b:24. 

Ophionereis reticulata.—Liitken, 1859:212, pl. 3: fig. 6a-c.— 

Lyman, 1865:141; 1875:4; 1878:224; 1882:162, pl. 40: figs. 

13-15; 1883:253.—Ljungman, 1866:310; 1871:620.—Verrill, 1867:342; 1868:366; 1899a:30; 1899b:377.—Rathbun, 1879:152.—Ives, 1889:177.—Liitken and Mortensen, 

1899:162, pl. 13: fig. 10.—H. L. Clark, 1901:249; 1915: 

289; 1919:57, 58, 72; 1933:39, 64; 1942:378.—Koehler, 

1907:315; 1913:360; 1914:2, 44, 59, 155.—A. H. Clark, 

1921:42, 55; 1922:211; 1939a:450; 1954:377.—Engel, 

1939:5, 8.—Fontaine, 1953:202.—A. M. Clark, 1953:67, 

68, 73, fig. 3a, pl. 1: figs. 1, 2.—Millott, 1953:96-99.— 

Brito, 1960:2, fig. 2.—Tabb and Manning, 1961:566.— 

Parslow and Clark, 1963:27, 47.—Tommasi, 1970:86, 88, 

pl. 40: figs. 92, 93.—Boffi, 1972:319.—Godcharles and 

Japp, 1973:57.—Thomas, 1973:586, figs. 1,5A.—Devaney, 

1974:132. 

Amphiura stearnsi Ives, 1891:339, 340, pl. 16: figs. 1-3. 

Ophionereis olivacea.—Roa, 1967:284, fig. 12 [not 0. olivacea H. 

L. Clark, 1901]. 

MATERIAL DEPOSITED.—USNM El 7738 (1 

spec); USNM El7851 (2 spec). 

DISTRIBUTION.—Collected in the lagoon at the 

south end of South Water Cay and from the pile 

of discarded conch shells at the foot of the pier of 

Carrie Bow Cay (station 1); on the reef flat in 

both the north and south Penicillus zone (stations 

4 and 5); and from just inside the reef crest 

(station 7). Two juvenile specimens were collected 

by bulk sample from the fore-reef slope (station 

13). 

REMARKS.—Previously reported from Lighthouse Reef (Devaney, 1974:132). 

Ophionereis squamulosa Koehler 

FIGURE 175 

Ophionereis dubia.—H. L. Clark, 1901:248 [not 0. dubia Muller 

and Troschel, 1842]. 

Ophionereis squamata.—Koehler, 1913:360, pl. 21: figs. 4-6 

[not 0. squamata Ljungman, 1866:310]. 

FIGURE 175.—Ophionereis squamulosa, USNM El7847, live. 

(Scale = 5 mm). 
396 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Ophionereis squamulosa Koehler, 1914:2, 44.—H. L. Clark, 

1915:290; 1918:326; 1919:57, 58, 59, 72; 1933:39, 65.— 

Mortensen, 1921:171-177, pl. 31: figs. 1-6.—Pearson, 

1936:70-72 [possibly 0. reticulata}.—A. H. Clark, 1939a: 

450.—A. M. Clark, 1953:67, 71, fig. 2, pl. 3: figs. 3, 4.— 

Parslow and Clark, 1963:27, 47.—Tommasi, 1970:87, 88, 

pl. 41: fig. 94.—Thomas, 1973:588, figs. 2, 5b. 

Ophionereis reticulata.—Roa, 1967:284 [probably 0. squamulosa] . 

MATERIAL DEPOSITED.—USNM El 7847 (1 

spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat in the coral head/coral pavement zone (station 6); and from the spur and groove zone of the 

inner fore reef in 10 to 12 m of water (station 10). 

Family OPHIOCOMIDAE 

Ophiocoma echinata (Lamarck) 

Ophiura echinata Lamarck, 1816:543. 

Ophiocoma echinata.—Agassiz, 1835:192.—Liitken, 1859:244, 

pl. 4: fig. 7a-d.—Lyman, 1865:81.—Ljungman, 1871: 

621.—Rathbun, 1879:152.—Ives, 1889:177.—Grave, 

1898:6, figs. 1-6.—Verrill, 1899a:22; 1899b:375.—H. L. 

Clark, 1901:245; 1915:291; 1919:57, 72; 1921:125; 1933: 

39, 65; 1941:89.—Koehler, 1907:325; 1913:374; 1914:3, 

117, 159.—A. H.Clark, 1921:55; 1922:212; 1939a:450, pl. 

54: fig. 4; 1954:377.—Engel, 1939:5, 8.—Fontaine, 1953: 

199, 203.—John and Clark, 1954:140.—Parslow and 

Clark, 1963:46.—Roa, 1967:292.—Tommasi, 1970:81.— 

Devaney, 1974:132.—Zeiller, 1974:107 [color fig.]. 

Ophiocoma crassispina Say, 1825:147.—Ljungman, 1866: 

329.—Ives, 1889:177. 

MATERIAL DEPOSITED.—USNM El 7845 (1 

spec); USNM El7854 (3 spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); under rocks 

in a rubble area between sand and Thalassia on 

the lagoon side of the south end of Carrie Bow 

Cay (station 2); in the patch reef zone of the back 

reef along the transect (station 3); on the reef flat 

in both the north and south Penicillus zone (stations 4 and 5); among coral rubble in the coralhead/coral-pavement zone (station 6); just inside 

the reef crest (station 7); on the reef crest in the 

rubble of the coral berm (station 8); and in the 

spur and groove zone of the inner fore reef from 

a coral bulk sample taken in 10 to 12 m of water 

(station 10). 

REMARKS.—Very common. Previously reported 

from Lighthouse Reef and Glover's Reef (Devaney, 1974:132). 

Ophiocoma paucigranulata Devaney 

Ophiocoma paucigranulata Devaney, 1974:132, figs. 5-8. 

MATERIAL DEPOSITED.—USNM El7783 (19 

spec); USNM E17784 (1 spec); USNM E17785 

(1 spec); USNM El7786 (3 spec). 

DISTRIBUTION.—Twenty-five specimens collected on the reef crest in the rubble of the coral 

berm (station 8), and one specimen taken by bulk 

sample from the outer ridge of the outer fore reef 

(station 12). 

REMARKS.—Devaney (1974:132) described this 

species on the basis of two specimens from Long 

Cay, Lighthouse Reef. Our specimens have the 

disc thoroughly covered with granules but otherwise they agree with Devaney's description. 

Ophiocoma pumila Liitken 

Ophiocoma pumila Liitken, 1859:141, 146, pl. 4: fig. 5a-c [not 

1859:248, pl 4: fig. 5d, = Ophiocomella ophiactoides (H. L. 

Clark)].—Lyman, 1865:71.—Ljungman, 1866:328; 1871: 

621.—Ives, 1889:177.—Verrill, 1899a:23; 1899b:375.—H. 

L. Clark, 1901:245; 1915:293; 1919:57, 72; 1921:131; 

1933:39, 67.—Koehler, 1907:326; 1913:375; 1914:3, 117, 

160.—A. H. Clark, 1921:55; 1922:212; 1939a:451 [not pl. 

54: fig. 3 = Ophiocomella ophiactoides}; 1954:377.—Engel, 

1939:5, 9.—Fontaine, 1953:199, 203.—Roa, 1967:292, fig. 

22.—Tommasi, 1970:82.—Devaney, 1970:10, 28, 29, 30-

32, fig. 16; 1974:139. 

MATERIAL DEPOSITED.—USNM El 7799 (2 

spec); USNM El7807 (3 spec); USNM El7853 

(1 spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat under coral rubble in the coral-head/coralpavement zone (station 6); just inside the reef 

crest (station 7); on the reef crest in the rubble of 

the coral berm (station 8); in the spur and groove 

zone of the inner fore reef from a coral bulk 

sample taken in 10 to 12 m of water (station 10), 
NUMBER 12 397 

and in about 26 m of water on the fore-reef slope 

(station 13). 

REMARKS.—Common. One specimen from station 1 has six arms and shows no evidence of 

fissiparity; it is not Ophiocomella ophiactoides. Previously reported from Belize (Koehler, 1914:160) 

and from Turneffe Islands, Lighthouse Reef, and 

Glover's Reef (Devaney, 1974:139). 

Ophiocoma wendti Muller and Troschel 

Ophiocoma wendti Miiller and Troschel, 1842:99.—Ljungman, 

1871:621.—Devaney, 1970:34,35,37; 1974:140. 

Ophiocoma riisei Liitken, 1859:243, 245, pl. 4; fig. 6a-d.— 

Lyman, 1865:76.—Ljungman, 1866:328.—Rathbun, 

1879:152.—Ives, 1889:177.—Stearns, 1891:4.—Verrill, 

1899a:22; 1899b:375.—H. L. Clark, 1901:245; 1915:293; 

1919:57, 72; 1921:128; 1933:39, 66.—Koehler, 1907:326; 

1913:375; 1914:3, 118, 159.—A. H. Clark, 1921:56; 1922: 

212; 1939a:450,pl. 54: fig. 5; 1954:377.—Engel, 1939:5.— 

Fontaine, 1953:199, 203.—Parslow and Clark, 1963:47.— 

Roa, 1967:292.—Tommasi, 1970:82. 

Not Ophiocoma wendti.—Koehler, 1907:327, pl. 13: fig. 38.— 

H. L.Clark, 1921:129. 

MATERIAL DEPOSITED.—USNM El 7733 (1 

spec); USNM El7737 (1 spec); USNM El7855 

(2 spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat in the north and south Penicillus zones (stations 4 and 5); under coral rubble in the coralhead/coral-pavement zone (station 6); on the reef 

crest in the rubble of the coral berm (station 8); 

in the spur and groove zone of the inner fore reef 

from a coral bulk sample taken in 10 to 12 m of 

water (station 10); and by bulk sample from the 

outer ridge of the outer fore reef (station 12). 

REMARKS.—Common. Previously reported 

from Belize (Koehler, 1914:159, as 0. nisei Liitken) and from Lighthouse Reef and Glover's Reef 

(Devaney, 1974:140). 

Ophiocomella ophiactoides (H. L. Clark) 

Ophiocoma pumila [part] Liitken, 1859:248, pl. 4: fig. 5d [not 

1859:141, 146, pl. 4: fig. 5a-c].—A. H. Clark, 1922:212; 

1939a:451, pl. 54: fig. 3.—H. L. Clark, 1942:378 [questionable]. 

Ophiacanlha ophiactoides H. L. Clark, 1901:249, pl. 15: figs. 5-

8; 1915:207; 1933:34,43. 

Ophiacanlha oligacantha H. L. Clark, 1918:265-267, pl. 7: fig 

5; 1919:56, 71. 

Ophiocomella canbbaea A. H. Clark, 1939b:7-8. 

Ophiocomella ophiactoides.—A. M. Clark in Parslow and Clark, 

1963:37, fig. 11; 1967:151.—Roa, 1967:291, fig. 21. 

MATERIAL DEPOSITED.—USNM El 7736 (1 

spec). 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1); on the reef 

flat in the south Penicillus zone (station 5); and in 

the coral-head/coral-pavement zone (station 6). 

REMARKS.—Five specimens have recently autotomized and have three arms and a broken disc. 

Of these, one is 2+1+0 and one is 1+2+0 (old to 

regenerating to new arms). Of the other specimens, one has five arms and shows no evidence of 

fissiparity; fourteen have six arms, showing three 

old and three regenerating arms when the distinction can be made, and two of these can be 

further categorized as 2+1+3; one has six arms 

with two old and four regenerating arms; and 

one has seven arms with four old and three 

regenerating arms. 

Ophiopsila riisei Liitken 

FIGURE 176 

Ophiopsila nisei Liitken 1859:136, pl. 5: fig. 2a-c.—Lyman, 

1865:150, figs. 16-17.—Ljungman, 1866:330; 1871:621.— 

Rathbun, 1879:152.—Verrill, 1899a:23.—H. L. Clark, 

1901:245; 1915:299; 1918:330; 1919:57,72; 1933:39,67.— 

Koehler, 1907:315; 1914:2, 45, 155.—A. H. Clark, 1939a: 

. 452; 1954:377.—Roa, 1967:293.—Tommasi, 1970:85. 

MATERIAL DEPOSITED.—USNM El 7648 (2 

spec); USNM El7758 (7 spec); USNM El7790 

(1 spec). 

DISTRIBUTION.—Twelve specimens from coral 

substrate from the fore-reef slope (station 13). 

REMARKS.—One specimen has an amphiurid 

clinging mouth to mouth. Three arms of the 

amphiurid are extended in the gutter formed by 

the lowermost Ophiopsila arm spines (Figure 176). 

Without separating the organisms, identification 

of the amphiurid as cf. Amphiura stimpsom is only 

tentative. 
398 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 176.—Ophiopsila nisei, with small amphurid cf. Amphiura stimpsonii clinging mouth to mouth; USNM El7648, 

dry. (Scale = 2 mm.) 

Family OPHIODERMATIDAE 

Ophioderma anitae new species 

FIGURES 177, 178a; PLATE 5: top right 

MATERIAL DEPOSITED.—Nine specimens collected on the reef crest in the rubble of the coral 

berm (station 8) on 3, 5, and 7 May 1974. 

Holotype: USNM El7641, 5 May 1974 (photographed) . 

Paratypes: USNM El7642 (1 spec, photographed), USNM El7643 (1 spec); USNM 

El7644 (2 spec), USNM El7645 (3 spec), all 3 

May 1974; USNM El7646 (1 spec), 7 May 1974. 

DIAGNOSIS.—Upper arm plates composed of a 

single piece, not fragmented; proximal upper arm 

plates nearly twice as broad as long, broadly in 

contact, with slight concave arc to distal edge. 

Disc granules fine, closely spaced, not squamose. 

Oral and adoral shields and small radial shields 

not covered by granules. Arms long, up to five 

and two-thirds times disc diameter, uniformly 

tapering, circular in section proximally, flat in 

section near the tip, exceedingly smooth where 

arm spines are pressed in. Arm spines 7 to 9, 

rarely 10. Disc grayish brown with yellow 

splotches, white underside; arms brown above, 

buff below, yellow-brown arm spines. 

DESCRIPTION.—Holotype: Disc diameter 18.4 

mm. Arm length from edge of disk 104.6 mm. 

The disc is about circular with slight interradial 

swellings. It is covered on both surfaces with very 

fine granules that are closely spaced but not 

touching, and that on the upper surface leave 

only the radial shields exposed. The radial shields 

are very small and only measure about 1.2 mm 

in length; they are irregularly oval, about twothirds as broad as long. The two shields of each 

pair are separated by a space about as wide as 

the width of the arm base. The granules of the 

upper surface of the disc do not pass out onto the 

upper surface of the arms or onto the few disc 

scales that form short radial notches, which encompass two minute upper arm plates, a halfexposed upper arm plate and an almost fully 

exposed upper arm plate. 

The under surface of the disc is covered with 

granules identical with those of the upper surface, 

but the oral and adoral shields and the scales 

between he outer genital slits and the arm bases 

are free of granules. The inner genital slits extend 

from near the distal edge of the first side arm 

plate to one or less granules short of the oral 

shields. The outer genital slits are located approximately at the level of the fifth side arm plate and 

are bordered on their interradial edge by 14 to 16 

granules. 

The oral shields are large, almost five-sided, 

not quite twice as wide as high; the distal border 

is almost straight or slightly concave except in the 

madreporic interradius where it is slightly convex; 

the proximal corner is broadly rounded and confluent with the adjoining sides, and at its tips 

there is a small to minute supplementary plate 

without granules. The adoral shields form broad 

lobes at the radial corners of the oral shields and 

adjoin the outermost oral papillae, the lower edge 

of the buccal tentacle scale and the lateral corners 

of the first (zeroeth in terms of visible arm segments) ventral arm plate; they are separated by 

granules from the first side arm plates. The oral 

plates are concealed by granules. Not counting 

the buccal tentacle, there are nine oral papillae 

on the side of most jaw angles, but sometimes 

eight or 10. The outermost are broadest, about as 
NUMBER 12 399 

FIGURE 177.—Ophioderma anitae, USNM El7641, holotype, live: a, b, aboral; c, oral. (Scale 

10 mm for a; 5 mm for b, c.) 

broad as long; the others are somewhat prism 

shaped, about twice as long as broad, becoming 

more pointed toward the apex of the jaw, overlapping in the direction of the jaw apex; the two 

at the apex of the jaw are again quite large and 

overlie the vertical row of teeth. 

The arms are circular in section proximally, 

long, and very slightly and uniformly tapering in 

both width and thickness, gradually becoming 

very flat in section near the arm tip; exceedingly 

smooth where arm spines are pressed against the 

side of the arms. 

Upper arm plates composed of a single piece, 

not fragmented. The upper arm plates beyond 

the radial notch are very wide, nearly rectangular 

but having a subtle yet distinct concave arc to 

the distal edge, changing to a similarly subtle 

convex arc which strengthens near the uppermost 

arm spine. Viewed from above, the proximal 

upper arm plates are slightly more than twice as 

wide as long. Near the arm tips the plates become 

trapezoidal, about as wide as long, as the side 

arm plates encroach on the upper surface of the 

arm. The upper arm plates are broadly in contact 

except for the last 10 or so segments at the arm 

tip. 

The first lower arm plate, not associated with 

a visible arm segment, is about twice as broad as 

long; the distal edge is shallowly rounded or very 

slightly angular. Between this and the lower arm 

plate of the first arm segment there is a pair of 

narrow slit-like, almost concealed, pores. There 

are no such pores between succeeding lower arm 

plates. The succeeding lower arm plates overlap 

slightly and appear nearly circular proximally, 

rounded-octagonal more distally, owing in part 
400 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

to strong overlap by the inner tentacle scales; 

close to the tip of the arm they ultimately become 

fan shaped as the side arm plates encroach further 

onto the under surface of the arm. 

The side arm plates abut in smooth contour 

the upper and lower arm plates, giving the arm 

its very smooth nearly circular section. The arm 

spines also match this contour when pressed 

against the side of the arm. The arm spines are 

therefore flat; they gradually increase in length 

from uppermost to lowermost, and taper to a 

blunt tip. Length of uppermost arm spines about 

half an arm segment long, overlapping and often 

extending beyond the rounded lower edge of the 

upper arm plate. Lowermost arm spines slightly 

overlap succeeding tentacle scales, with greater 

overlap on the first seven arm segments. The 

number of arm spines by segment are given in 

Table 28. 

Two tentacle scales are present except at the 

very tip of the arm where only one is seen. Outer 

tentacle scale substantially wider than inner, 

squarish, resembling a ninety degree quadrant of 

a circle, and overlapping the articular base of the 

lowermost arm spine. Inner tentacle scale approximately oval, about two and one-half to three 

times as long as broad, about half an arm segment 

long, and about one-half to one-third again as 

long as the outer tentacle scale. The distance 

between the inner edges of the inner tentacle 

scales of two sides of an arm is about one and 

one-half to one and two-thirds the length of one 

inner tentacle scale (Figure 178a). 

Coloration, grayish brown with yellow 

splotches, is shown in Plate 5 {top right). In alcohol 

yellow areas have turned white, and the brown 

has dulled on drying. The yellow splotches vary 

from nearly circular to oval and are somewhat 

smaller than the radial shields. Radial shields are 

the same brown as the ground color of the disc; 

however, the disc scales beneath the brown granules are only tinged with brown in the preserved 

specimen. Side arm plates are brown, under surface of disc white, under arm plates buff, and 

arm spines yellow-brown. There are no banding 

patterns on the arms. 

0

****mffffffffi * f i I # ?.? y 

FIGURE 178.—Comparison of arm tips in two species of 

Ophioderma: a, 0. anitae, USNM El 7645, dry; b. O. rubicundum, 

USNM El7683, dry. (Scale = 5 mm.) 

TABLE 28.—Number of arm spines by segments in specimens of Ophioderma anitae, new species 

Specimen 

HOLOTYPE 

USNM El7461 

PARATYPES 

USNM El7646 

USNM El7644 

USNM El7645 

USNM El7645 

USNM El7645 

USNM El7643 

USNM El7644 

USNM El7642 

1 

3 

3 

2 

2 

3 

3 

2 

3 

2 

2 

4 

3 

3 

3 

3 

4 

4 

3 

4 

3 

5 

5 

4 

4 

1 

5 

4 

5 

4 

4 

6 

6 

5 

5 

5 

6 

5 

5 

5 

5 

7 

7 

6 

6 

6 

7 

6 

6 

6 

6 

7 

7 

7 

6 

7 

8 

7 

7 

7 

7 

9 

8 

8 

7 

8 

9 

8 

8 

7 

8 

9 

9 

8 

9 

9 

8 

7 

8 

7 

9 

9 

9 

8 

9 

9 

9 

7 

8 

7 

10 

9 

9 

8 

9 

9 

9 

8 

8 

7 

11 

10 

9 

8 

9 

9 

9 

7 

8 

7 

12 

9 

9 

8 

9 

9 

8 

9 

7 

7 

Segment 

13 

8 

9 

8 

9 

9 

9 

8 

8 

7 

14 

9 

8 

8 

8 

9 

8 

8 

8 

7 

num 

15 

9 

9 

8 

9 

9 

8 

8 

8 

7 

her 

16 

9 

9 

8 

9 

9 

8 

8 

8 

7 

17 

9 

8 

8 

10 

9 

8 

8 

8 

7 

18 

9 

8 

7 

9 

9 

8 

9 

8 

7 

19 

9 

8 

8 

9 

9 

8 

8 

8 

7 

20 

9 

8 

7 

9 

9 

8 

8 

8 

7 

21 

9 

7 

7 

8 

9 

8 

8 

8 

7 

22 

8 

8 

7 

8 

8 

8 

8 

7 

7 

23 

9 

8 

7 

8 

9 

8 

8 

7 

6 

24 

9 

8 

7 

9 

8 

8 

8 

7 

7 

25 

8 

8 

7 

8 

9 

8 

8 

7 

7 

Disc diameter 

(mm) 

18.5 

17.3 

17.0 

16.3 

14.0 

13.7 

13.5 

12.0 

11.3 
NUMBER 12 401 

Paratypes: Disc diameters range from 11.3 to 

17.3 mm. In the smaller specimens the arms are 

disproportionately shorter, the outer genital slit is 

beside the fourth side arm plate, there can be 

only seven or eight oral papillae, the yellow markings are more commonly larger than the radial 

shields, often there is no evidence of a supplementary plate at the proximal corner of the oral 

shield, and the oral shields are more nearly as 

wide as high. 

COMPARISON WITH OTHE R SPECIES.—Ophioderma 

anitae keys to the 0. cinereum, 0. phoenium, and 0. 

rubicundum couplets of H. L. Clark's (1933) key, 

and to the 0. pallidum (Verrill)-O. phoenium couplet in Ziesenhenne's (1955) key. Ophioderma cinereum is distinguished from the present 

species by its subdivided uppe r arm plates; 0. 

phoenium is recognized by adoral shields covered 

with granules; 0. rubicundum is closest but is entirely different in color and details of morphology. 

In a specimen of 0. rubicundum from the same 

habitat (USNM El7683) with disc diameter 14.8 

mm the length of the first 15 segments from the 

mouth frame is 14.5 mm, whereas in a specimen 

of 0. anitae (USNM E17645) with disc diameter 

14.0 mm the length of the first 15 segments from 

the mout h frame is 15.3 mm (Figure 178). Further, the arm tips of 0. rubicundum are more slender 

and not as flattened as in 0. anitae (Figure 178). 

ETYMOLOGY.—Name d for Anita Pruzan. 

Ophioderma appressum (Say) 

PLATE 5: top right 

Ophiura appressa Say, 1825:151.—Lyman, 1865:34.—Rathbun, 1879:151.—Ives 1889:175.—Verrill, 1899a:6.—H. L. 

Clark, 1901:242.—A. H. Clark, 1939b: 10. 

Ophioderma virescens Liitken, 1859:86, pl. 1: fig. 4a-d. 

Ophioderma appressum.—Ljungman, 1866:304; 1871:615.— 

Lyman, 1882:9.—Koehler, 1913:353.—H. L. Clark, 1915: 

300; 1919:57, 72; 1933:40, 68; 1942:379.—A. H. Clark, 

1922:213; 1939a:452; 1954:377.—Boone, 1933:113, pis. 

66, 75b, 76b.—Engel, 1939:5, 9.—Caso, 1951:219, 222, 

243, 249, 254, figs. 15-18; 1953:222.—Ziesenhenne, 1955: 

188, 200.—Costa and Costa, 1962:3.—Parslow and Clark, 

1963:47.—Roa, 1967:294.—Boffi, 1972:319.—Devaney, 

1974:141, fig. 9.—Zeiller, 1974:107 [color fig.]. 

Ophioderma appressa.—Koehler, 1907:281; 1914:1,3, 147.—A. 

H. Clark, 1921:56: 1939b: 10.—Caso, 1961:176-178, figs. 

74, 75.—Tommasi, 1970:65. 

MATERIAL DEPOSITED.—USN M El767 6 (3 

spec); USN M El767 8 (2 spec); USN M El7680 

(2 spec). 

DISTRIBUTION.—Two specimens collected 

amon g the discarded conch shells at the foot of 

the pier on the lagoon side of Carrie Bow Cay 

(station 1). Very common on the reef flat where 

it was collected in the north and south Penicillus 

zones (stations 4 and 5), and in the coral-head/ 

coral-pavement zone (station 6). Collected just 

inside the reef crest (station 7) and on the reef 

crest in the rubbl e of the coral berm (station 8). 

REMARKS.—Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

141). 

Ophioderma brevicaudum Liitken 

Ophioderma brevicaudum Liitken, 1856a:8; 1859:94, pis. 1, 2: 

fig. 3a-c—H. L. Clark, 1915:300; 1919:57, 72; 1933:40, 

69.—A. H. Clark, 1922:213; 1939a:452, pl. 54: figs. 1, 2; 

1954:377.—Engel, 1939:5, 9.—John and Clark, 1954: 

140.—Ziesenhenne, 1955:188, 199.—Roa, 1967:294.—Devaney, 1974:141. 

Ophiura brevicauda.— Verrill, 1899a:5.—H. L. Clark, 1901: 

242. 

Ophioderma brevicauda.—Ljungman, 1866:303; 1871:615.— 

Koehler, 1907:281; 1914:1,4, 147.—A. H. Clark, 1921:56. 

MATERIAL DEPOSITED.—USN M El 7677 (2 

spec). 

DISTRIBUTION.—Ten specimens collected on the 

reef flat in the south Penicillus zone (station 5), in 

the coral rubble of the coral-head/coral-pavement zone (station 6), and from just inside the 

reef crest (station 7). On e specimen also collected 

on the reef crest in the rubble of the coral berm 

(station 8). 

REMARKS.—Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

141). 

Ophioderma brevispinum (Say) 

Ophiura brevispina Say, 1825:149.—Lyman, 1865:18.—Rathbun, 1879:151.—Ives, 1889:175.—Verrill, 1899a:4.— 

Brooks and Grave, 1899:83, pis. 1, 2: figs. 1-5. 

—H.L.Clark, 1901:242. 
402 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Ophiura olivaceum Ayres, 1852:134. 

Ophioderma serpens Liitken, 1859:86, pis. 1, 2: fig. 6a-c. 

Ophiura olivacea. —Liitken, 1859:96.—Grave, 1899:92, figs. 

1-5. 

Ophioderma olivaceum.—Ljungman, 1866:304. 

Ophioderma brevispina.—Ljungman, 1866:303; 1871:615.—Lyman, 1882:9.—Koehler, 1907:281; 1913:354; 1914:1, 4, 

147.—Fontaine, 1953:204.—Cherbonnier, 1959:261 — 

262, figs. 5a,b, 6a,b.—Tabb and Manning, 1961:566.— 

Parslow and Clark, 1963:48.—Tommasi, 1970:66. 

Ophioderma brevispinum.—H. L. Clark, 1915:300; 1919:57, 72; 

1933:40, 70; 1941:89.—Engle, 1939:5, 9.—A. H. Clark, 

1954:377.—Ziesenhenne, 1955:188, 198.—Roa, 1967:294, 

fig. 24.—Godcharles, 1971:28, 33.—Godcharles and Jaap, 

1973:57.—Stancyk, 1973:10. 

MATERIAL DEPOSITED.—USNM El 7649 (1 

spec). 

DISTRIBUTION.—One specimen was collected in 

the north Penicillus zone under rubble (station 4). 

REMARKS.—The specimen was orange in life 

but has lost all color in alcohol. This species is 

known to have a vitellaria larva (Brooks and 

Grave, 1899). 

Ophioderma cinereum Muller and Troschel 

Ophioderma cinereum Muller and Troschel, 1842:87.—Ljung 

man, 1866:304; 1871:615.—Koehler, 1913:354.—H. L 

Clark, 1915:301; 1918:337; 1919:57,72; 1933:40,71; 1942: 

379.—A. H. Clark, 1922:212; 1939a:452; 1954:377.-

Boone, 1933:115-116, pis. 68, 69.—Engel, 1939:5, 9.— 

Caso, 1951:219, 221, 243, 249, figs. 11-14; 1953:215, 222 

1961:157-162, figs. 61-64.—Zeisenhenne, 1955:187 

192.—da Costa and da Costa, 1962:2.—Thomas, 1962b 

65, fig. Ic—Parslow and Clark, 1963:48.—Roa, 1967 

295.—Tommasi, 1970:67.—Godcharles and Jaap, 1973 

57.—Devaney, 1974:143. 

Ophioderma antillarum Liitken, 1859:190, pl. 1: fig. la-c. 

Ophiura cmerea.— Lyman, 1865:27.—Verrill, 1868:367.— 

Rathbun, 1879:150.—Ives, 1889:175; 1890:331 1891: 

339.—Stearns, 189l:iv.—Verrill, 1899a:6.—H. L. Clark, 

1901:242. 

Ophioderma cinerea.—Koehler, 1907:281; 1914:1,6,9, 147.— 

A. H. Clark, 1921:57. 

Ophiocryptus hexacanthus H. L. Clark, 1915:64. 

MATERIAL DEPOSITED.—USNM El 7856 (9 

spec.) 

DISTRIBUTION.—Collected among the discarded 

conch shells at the foot of the pier on the lagoon 

side of Carrie Bow Cay (station 1) and just inside 

the reef crest (station 7). 

REMARKS.—Previously reported from Belize 

(Koehler, 1914:147), Turneffe Islands, Lighthouse Reef, and Glover's Reef (Devaney, 1974: 

143). 

Ophioderma guttatum Liitken 

PLATE 5: top right 

Ophioderma guttata Liitken, 1859:95, pl. 1: fig. 8a,b. 

Ophiura guttata.—iA. L. Clark, 1901:251. 

Ophiura guttatum.—H. L. Clark, 1915:301; 1919:57, 72. 

Ophioderma guttatum.—Ljungman, 1866:303.—H. L. Clark, 

1933: 40, 72.—Ziesenhenne, 1955:186, 189.—Thomas, 

1962b:65, fig. la.—Zeiller, 1974:108 [color fig.]. 

MATERIAL DEPOSITED.—USNM El 7684 (2 

spec); USNM El7685 (1 spec); USNM El7777 

(1 spec); USNM El7778 (1 spec). 

DISTRIBUTION.—Five specimens collected by 

Kjell Sandved just inside the reef crest (station 

7), and on the reef crest in the rubble of the coral 

berm (station 8). 

Ophioderma phoenium H. L. Clark 

PLATE 5: top right 

Ophioderma phoenium H. L. Clark, 1918:333, pl. 6: figs. 1, 2; 

1919:57, 72, pl. 3: fig. 1; 1933:40, 71.—Ziesenhenne, 1955: 

187, 192.—Devaney, 1974:144. 

MATERIAL DEPOSITED.—USNM El 7690 (1 

spec); USNM El7776 (1 spec); USNM El7781 

(1 spec). 

DISTRIBUTION.—Three specimens collected on 

the reef crest in the rubble of the coral berm 

(station 8). 

REMARKS.—Previously reported from Lighthouse Reef (Devaney, 1974:144). 

Ophioderma rubicundum Liitken 

FIGURE 1786; PLATE 5: top right 

Ophioderma rubicumdum Liitken, 1856a:8; 1859:192, pl. 1: fig. 

2a-c—Ljungman, 1866:304.—H. L. Clark, 1915:302; 

1919:57, 72; 1933:40, 71.—A. H. Clark, 1922:213; 1939a: 

452; 1954:377.—Ziesenhenne, 1955:187, 197.—Roa, 1967: 

295.—Devaney, 1974:144. 
NUMBER 12 403 

Ophiura rubicunda.— Verrill, 1899a:6.—H. L. Clark, 1901:242. 

Ophioderma rubicunda.—Koehler, 1907:282; 1914:1, 7, 148.— 

A. H. Clark, 1921:57. 

MATERIAL DEPOSITED.—USNM El 7683 (1 

spec); USNM El7687 (2 spec). 

DISTRIBUTION.—Collected by Kjell Sandved 

just inside the reef crest (station 7) and on the 

reef crest in the rubble of the coral berm (station 

8). Collected in the spur and groove zone of the 

inner fore reef from a coral bulk sample taken in 

10 to 12 m of water (station 10), from the outer 

ridge of the outer fore reef (station 12), and from 

the fore-reef slope at 26 to 28 m (station 13). 

REMARKS.—Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

144). 

Ophioderma squamosissimum Liitken 

Ophioderma squamosissimum Liitken, 1856a:8; 1859:92, pl. 1: 

fig. 7a,b.—Ljungman, 1866:304; 1871:615.—H. L. Clark, 

1915:302; 1918:335, pl. 4: fig. 1, pl. 6: figs. 3, 4; 1919:57, 

69, 72, pl. 3: fig. 2; 1933:72.—Ziesenhenne, 1955:187, 

189.—Thomas, 1962b:62, fig. lb.—Parslow and Clark, 

1963:27.—Godcharles and Jaap, 1973:57.—Zeiller, 1974: 

108 [color fig.]. 

MATERIAL DEPOSITED.—CBC 75.5.15-1 (1 

spec); CBC 75.5.24 (1 spec.) (not catalogued). 

DISTRIBUTION.—Found inside the sponge Pseudoceratina crassa (Hyatt) collected by K. Riitzler 

on the inner-reef slope near station 11, 20 m (15 

May 1975); also found by B. Spracklin on a 

lagoon patch reef, 1 km WSW of Carrie Bow 

Cay, 4 m (24 May 1975). 

REMARKS.—Specimen CBC 75.5.15-1 has a disc 

diameter of 7 mm and an arm length of approximately 26 mm measured from the edge of the 

disc. The upper arm plates form five rows. The 

radial shields are exposed and consist of wellformed single plates. The sixth arm segment is 

just outside the margin of the disc and bears five 

spines on the side arm plates. The oral shields are 

exposed. There are no supplementary oral shields. 

The adoral shields carry a few large granules. A 

translucent gelantinous cuticle coats the entire 

exterior including spines, granules and oral papillae. The color in alcohol is white. 

Specimen CBC 75.5.24 has a disc diameter of 

35 to 38 mm. The arms are approximately 7 mm 

thick at the edge of the disc. Arm length from the 

edge of the disc is roughly approximated as 135 

mm. There is no noticeable gelatinous cuticle. 

Supplementary oral shields present distal to the 

oral shields. Upper arm plates form seven rows. 

There are six spines on the ninth arm segment 

just outside the disc. The adoral shields are concealed by the oral shields. There are no single 

conspicuous plates readily identifiable as radial 

shields; instead, a large number of small plates 

bordered by granules occur in the general area of 

the arm bases and on the aboral interradial peripheral half of the disc. The central area of the 

aboral disc and the oral interradial areas are 

completely covered by granules similar to those 

present in Ophioderma guttatum. 

Ophioderma species 

FIGURE 179 

MATERIAL DEPOSITED.—USNM El 7788 (1 

spec); USNM El7789 (1 spec). 

DISTRIBUTION.—USNM El7788 collected on 

the reef flat under rocks in the north Penicillus 

zone (station 4); USNM El7789 on the reef flat 

under rocks in the south Penicillus zone (station 

5). 

REMARKS.—USNM El7788 has 15.8 mm disc 

diameter; upper arm plates entire; adoral shields 

bare; three small radial shields bare; 10 and 11 

blunt-tipped arm spines on arm segments eight 

through 16, lowermost arm spine longest. 

Although the specimen keys to Ophioderma rubicundum following Ziesenhenne (1955), it compares 

more favorably with 0. appressum in total morphology and habitat. The color is mottled greenish brown and, as also found by Devaney (1974) 

at Lighthouse Reef and Glover's Reef, almost all 

0. appressum at Carrie Bow Cay have adoral 

shields free of granules. The absence of granules 

on three radial shields leaves identification problematic. 
404 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 179.—Ophioderma sp., USNM El7788, dry. (Scale = 10 mm.) 

USNM El7789 agrees in all respects with the 

above except that all radial shields are covered 

by granules. 

Ophioderma species, juvenile 

FIGURE 180 

MATERIAL DEPOSITED.—USNM El 7686 (1 

spec). 

DISTRIBUTION.—Collected on the reef flat in the 

rubble of the coral-head/coral-pavement zone 

(station 6). 

REMARKS.—Disc diameter 3.2 mm; upper arm 

plates entire; radial shields and adoral shields 

covered by granules; disc granules do not extend 

onto any part of the arms; arm spines five, less 

than half an arm segment long, conical, sharply 

pointed, of equal length; arm tips flattened. 

The arm spines resemble those of both 0. brevicaudum and 0. brevispinum. The former, however, 

has relatively shorter arm segments and is said to 

pass through an Ophiocryptus stage with granules 

on the arms (A. H. Clark, 1939a); the latter has 

adoral shields free of granules. The upper surface 

of the disc is red with a central white spot, the 

lower surface white, and the arms banded red 

and white. Identification is problematic. 

Ophiurochaeta littoralis (Koehler) 

Ophiolimna littoralis Koehler, 1913:370, pl. 21: figs. 1-3; 1914: 

3, 96, 111. 

Ophiolreta littoralis.—H. L. Clark, 1915:216; 1919:56; 1933: 

34, 43.—A. H. Clark, 1954:377.—Parslow and Clark, 

1963:26. 

Ophiurochaeta littoralis.—Matsumoto, 1917:316-317.—Devaney, 1974:147, figs. 10-12. 
NUMBER 12 405 

FIGURE 180.—Ophioderma sp., juvenile, USNM El7686, dry. (Scale = 2 mm.) 

MATERIAL DEPOSITED.—USNM El 7795 (1 

spec); USNM El7806 (1 spec); USNM El7810 

(1 spec). 

DISTRIBUTION.—Collected in the spur and 

groove zone of the inner fore reef from a coral 

bulk sample taken in 10 to 12 m of water (station 

10), and from the fore-reef slope in 26 and 28 

meters of water (station 13). 

REMARKS.—Presence of young in the bursa of 

specimen USNM El7795 suggests but does not 

prove viviparity. Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

147). 

Family OPHIURIDAE 

Ophiolepis impressa Liitken 

Ophiolepis impressa Liitken, 1859:203, pl. 2: fig. 3a,b.—Ljungman, 1866:306.—Devaney, 1974:154, figs. 13-15. 

Ophiozona impressa.—Lyman, 1865:64.—Verrill, 1899a:8.— 

H. L. Clark, 1901:242; 1915:337; 1919:57, 72; 1933:41, 

73.—Koehler, 1907:289; 1914:2, 11, 150.—Matsumoto, 

1915:82; 1917:298, 389.—A. H. Clark, 1921:57; 1922:213; 

1939a:452; 1954:378.—Engel, 1939:5.—Fell, 1960:33.— 

Roa, 1967:296.—Tommasi, 1970:74. 

MATERIAL DEPOSITED.—USNM El 7848 (2 

spec); USNM El7852 (3 spec). 

DISTRIBUTION.—Collected on the back reef in 

the patch-reef zone of the study transect (station 

3); on the reef flat in the north and south Penicillus 

zones (stations 4 and 5), in the coral-head/coralpavement zone (station 6); just inside the reef 

crest (station 7), and on the reef crest in the 

rubble of the coral berm (station 8). One small 

specimen was extracted from a bulk sample from 

the outer ridge of the outer fore reef (station 12). 

REMARKS.—Previously reported from Lighthouse Reef and Glover's Reef (Devaney, 1974: 

154). 

Ophiolepis paucispina (Say) 

Ophiura paucispina Say, 1825:149. 

Ophiolepis paucispina.—Miiller and Troschel, 1842:90.—Liitken, 1859:204-206, pl. 2: fig. 2a,b.—Lyman, 1865:55.— 

Ljungman, 1866:306; 1871:618.—Verrill, 1868:367; 

1899b:373.—Rathbun, 1879:151.—H. L. Clark, 1901:251; 

1915:343; 1919:57, 72; 1933:41, 74; 1942:379.—Koehler, 

1907:287; 1913:355; 1914:2, 11, 149.—A. H. Clark, 1921: 

32, 44, 50, 58, 62; 1939a:452; 1954:378.—Engel, 1939:5, 

9.—Fontaine, 1953:205.—Parslow and Clark 1963:48.— 

Tommasi, 1970:73, 92.—Devaney, 1974:155. 

MATERIAL DEPOSITED.—USNM El 7740 (1 

spec); USNM El7760 (8 spec). 

DISTRIBUTION.—Twenty-nine specimens collected among the discarded conch shells at the 

foot of the pier on the lagoon side of Carrie Bow 

Cay (station 1). 

REMARKS.—The presence of arm tips of mini-
406 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ature young protruding through the genital slits 

in three deposited specimens probably indicates 

that this species is viviparous. Collected by Devaney (1974:155) at Lighthouse Reef. 

Conclusions 

The following species are reported for the first 

time from Belize: Astrophyton muricatum, Ophiomitrella glabra, Sigsbeia murrhina, Amphiura fibulata, A. 

stimpsonii, Axiognathus squamatus, Ophiostigma sp. 

(undescribed), Ophiactis algicola, Ophionereis squamulosa, Ophiocomella ophiactoides, Ophiopsila riisei, 

Ophioderma anitae, new species, 0. brevispinum, 0. 

guttatum, 0. squamosissimum. Only Ophioderma anitae 

and Ophiocoma paucigranulata can at this time be 

considered possibly endemic to this part of the 

Gulf of Honduras. The fauna is otherwise widely 

distributed throughout the Greater and Lesser 

Antilles (Parslow and Clark, 1963), as might be 

expected from the general westerly flow of water 

through the Carribean (Duncan et al., 1977; Metcalfet al., 1977). 

Although many species are found in all the 

major physiographic parts of the reef, nevertheless 

the rank-order distributions show certain qualitative and quantitative trends from lagoon to fore 

reef. Table 29 is an attempt to characterize the 

composition and facies of the ophiuroid fauna 

present in four main physiographic areas of the 

reef. The distribution of individual species over 

the study area has been given in the preceding 

systematic section. Because no quantitative sampling was undertaken, the actual numbers of 

specimens of any species collected are not meaningful. Very common species were not collected 

on every encounter, and the discovery of a previously unrepresented species usually initiated an 

attempt to secure additional specimens. It appears, however, that this distortion of actual and 

relative abundances has not greatly upset the 

rank order of abundances. My subjective impresTABL E 29.—Rank-orde r abundance s of ophiuroids within major physiographi c areas nea r 

Carri e Bow Ca y (station numbers in parentheses, species arrange d in each column in decreasing 

orde r of abundance , vertical bars connec t species represented equally) 

Lagoo n (1,2) Ree f flat, back reef (3-7) Ree f crest (8, 9) Fore reef (10-13) 

Ophiothrix oerstedi 

Ophiactis savignyi 

Ophiothrix angulata 

Ophiolepis paucispina 

Ophiocoma pumila 

Ophiocomella ophiactoides 

Axiognathus squamatus 

Amphiura stimpsonii 

Ophionereis reticulata 

Ophioderma cinereum 

Ophiocoma echinata 

Ophiostigma sp. 

Ophiocoma wendti 

Amphiura fibulata 

Ophioderma appressum 

Ophionereis squamulosa 

Ophioderma appressum 

Ophiocoma echinata 

Ophiolepis impressa 

Ophiocoma wendti 

Ophiocoma pumila 

Ophioderma rubicundum 

Ophiomyxa flaccida 

Ophioderma brevicaudum 

Ophiothrix oerstedi 

Ophionereis squamulosa 

Ophiocomella ophiactoides 

Ophionereis reticulata 

Ophioderma cinereum 

Ophioderma guttatum 

Ophiactis savignyi 

Ophioderma sp. 

Ophiothrix angulata 

Ophioderma brevispinum 

Ophioderma sp., juv . 

Ophiocoma wendti 

Ophiocoma paucigranulata 

Ophiocoma pumila 

Ophiocoma echinata 

Ophioderma appressum 

Ophioderma anitae 

Ophiothrix oerstedi 

Ophiolepis impressa 

Ophioderma phoenium 

Ophioderma guttatum 

Ophiomyxa flaccida 

Ophioderma rubicundum 

Ophiomitrella glabra 

Ophioderma brevicaudum 

Ophiocoma pumila 

Ophiothrix angulata 

Ophioderma rubicundum 

Ophiurochaeta littoralis 

Ophiopsila riisei 

Ophiactis savignyi 

Ophiothrix suensoni 

Ophiothrix oerstedi 

Sigsbeia murrhina 

Amphiura stimpsonii 

Ophiocoma echinata 

Ophiocoma wendti 

Ophionereis reticulata 

Ophiostigma sp. 

Astrophyton muricatum 

Ophiomyxa flaccida 

Ophiomitrella glabra 

Ophiocoma paucigranulata 

Ophiolepis impressa 

Ophiactis algicola 

Ophionereis squamulosa 

Ophioderma squamosissimum 
NUMBER 12 407 

sion is that the most commo n species in the field 

are the most commo n in the collection (and conversely for the least common), but Tabl e 29 

should only be considered an approximation for 

a numbe r of reasons. Subjective field impressions 

involve psychometric aspects such as size and 

visibility which may only partly correlate with 

actual numbers of individuals. Certain species 

may be very adept at escaping collection with the 

hand methods employed. Although these results 

should be used with caution, they seem worth 

presenting, because I have not found any other 

attempts at rank ordering the species of ophiuroids by general habitat for the West Indian 

region. Species are listed in the table in decreasing 

level of abundance , with a vertical ba r connecting 

those equally represented by numbers of specimens in the collection. 

T he ophiuroids found amon g discarded conch 

shells on the lagoon side of the island (station 1) 

are mainly small or minut e species. It is notable 

that of the minut e species three are fissiparous 

{Ophiactis savignyi, Ophiocomella ophiactoides, and 

Ophiostigma sp.) and two (probably three) are 

viviparous {Axiognathus squamatus, Amphiura stimpsonii, and Ophiolepis paucispina). Th e species of 

ophiuroids found on the reef flat (stations 4-6), 

including those from the inner reef crest zone 

(station 7), an d the patch reef zone (station 3) are 

dominated by the families Ophiodermatidae , 

Ophiocomidae and Ophiurida e and the principal 

habitat is beneath coral, coral rock, or dead shells. 

The species of ophiuroids found on the reef crest, 

principally in the rubbl e of the coral berm (station 8), but also just seaward (station 9), are 

dominated by the families Ophiocomida e and 

Ophiodermatidae . It is likely that Ophiocoma paucigranulata is significantly overrepresented in the 

reef crest collection because once discovered, it 

was deliberately sought. Th e species of ophiuroids 

found on the fore reef—the spur and groove zone 

(station 10), the trough (station 11), the outer 

ridge (station 12), and also the accessible part of 

the fore reef slope (station 13)—are notable for 

the strong representation of epizootic species not 

found shallower, especially Astrophyton muricatum, 

Sigsbeia murrhina, Ophiothrix suensoni, Ophiopsila riisei, and Ophiurochaeta littoralis. Still, there is sufficient crevice habitat to support the presence of 

species of Ophiocoma, Ophionereis, and Ophiolepis, 

which are found principally unde r coral rock in 

shallower water. 

Although the reproductive biology of the 

ophiuroids of Carrie Bow Cay needs further extensive investigation, the discovery of young in 

the bursae of Ophiurochaeta littoralis and Ophiolepis 

paucispina and the confirmation of fissiparity in 

West Indian Ophiostigma deserve comment at this 

time. Th e presence of young in genital bursae 

suggests but does not prove that these ophiuroids 

are viviparous. Th e counter example was provided by Smith (1938) in the case of Ophiothrix 

fragilis, which has a planktotrophic ophiopluteus 

larva. Smith found very young post larvae clinging to the oral interbrachial arm spines and 

slightly older post larvae present in the bursae of 

a small percentage of the adult population. Similarly, A. H. Clark (1939a) found young post 

larvae comparably located on the spines of 

Ophiothrix hartfordi from Puerto Rico. Th e young 

post larvae of Ophiurochaeta littoralis and Ophiolepis 

paucispina may somehow preferentially settle from 

the plankton onto the adults, but the mechanism 

for such action is difficult to identify. Th e subsequent presence of young post larvae in bursae 

may be interpreted (in accord with Smith) as the 

result of crevice-seeking behavior. 

This possible strategy seems less applicable to 

Ophiolepis paucispina than to Ophiurochaeta littoralis 

because a relatively large proportion of the ophiuroids comparabl e in size with 0. paucispina in the 

habitat of discarded conch shells practice direct 

recruitment through either viviparity or fissiparity, as noted above. This aspect of the minut e 

ophiuroids of this habitat is highlighted not only 

by the possibility that 0. paucispina is viviparous, 

but also by the presence of exclusively fissiparous 

specimens of Ophiostigma sp. A. M. Clark 

(1967a) did not include Ophiostigma in her list of 

fissiparous ophiuroids, presumably because she 

tentatively referred the fissiparous specimen of 

Ophiostigma recorded by H. L. Clark (1942) from 
408 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Bermuda to Ophiocomella ophiactoides (see Parslow 

and Clark, 1963). In my opinion, however, the 

generic identity of the present fissiparous specimens is certain, although they seem to represent 

an undescribed species rather than Ophiostigma 

isacanthum. 

Literature Cited 

Agassiz, L. 

1835. Prodrome d'une monographic des radiaires ou 

echinodermes. Memoires de la Societe des Sciences 

Naturelles de Neuchatel, 1:168-199. 

Ayres, W. O. 

1852. Descriptions of Two New species of Ophiuridae. 

Proceedings of the Boston Society of Natural History, 4: 

133-135, 248-250. 

Bernasconi, Irene 

1926. Una ofiura vivipara de Necochea. Anales del Museo 

Nacional de Historia Natural "Bernardino Rivadavia,'' 

Buenos Aires, 34:145-153, 4 plates. 

Boffi, E. 

1972. Ecological Aspects of Ophiuroids from the Phytal 

of S. W. Atlantic Ocean Warm Waters. Marine 

Biology, 15:316-328. 

Boone, Lee 

1933. Scientific Results of Cruises of the Yachts Eagle 

and Ara, 1921-1928, William K. Vanderbilt, 

Commanding: Echinodermata. Bulletin of the Vanderbilt Marine Museum (Huntington), 4:68-164, 

plates 25-102. 

Brito, I. 

1960. Os ofiuroides do Rio de Janeiro, Parte 1: Ophiothrichidae, Ophiochitonidae e Ophiactidae. Universidade do Brasil Faculdade Nacional de Filosofia Centra 

de Estudos Zoologicos, 6:1-4, 3 figures. 

Brooks, W. K , and C. Grave 

1899. Ophiura brevispina. Memoirs of the National Academy of 

Sciences, 8:79-100, 3 plates. 

E. 

Contribucion al conoscimiento de los ofiuroides de 

Mexico, I: Algunas especies litorales. Anales del 

Instituto Biologica de la Universidad Nacional de Mexico, 

22:219-312, illustrated. 

Estado actual de los conocimientos acerca de la 

fauna de los equinodermos de Mexico. In Memoria 

del Congreso Cientifico Mexicano, VII, pages 209-222, 

12 figures. 

Los equinodermos de Mexico. 380 pages, 124 figures, 20 plates. Thesis, Facultad de Ciencias, Universidad Nacional de Mexico. 

Cherbonnier, G. 

1959. Echinodermes de la Guyane francaise, 3e note. 

Caso, M. 

1951. 

1953. 

1961. 

Bulletin du Museum National d'Histoire Naturelle, 31: 

261-265, 3 figures. 

Clark, A. H. 

1921. Report on the Ophiurans Collected by the Barbados-Antigua Expedition from the University of 

Iowa in 1918. University of Iowa Studies in Natural 

History, 9(5):29-64. 

1922. The Ophiurans of the Island of Curacao. Bijdragen 

tot de Dierkunde, 22:209-213. 

1939a. Echinoderms of the Smithsonian-Hartford Expedition, 1937, with Other West Indian Records. 

Proceedings of the United States National Museum, 86: 

441-456, plates 53, 54. 

1939b. Echinoderms (Other than Holothurians) Collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, 98(11): 1-18, 5 plates. 

1954. Echinoderms (Other than Holothurians) of the 

Gulf of Mexico. Fishery Bulletin of the Fish and 

Wildlife Service, 89(55):373-379. 

Clark, A. M. 

1953. A Revision of the Genus Ophionereis (Echinodermata, Ophiuroidea). Proceedings of the Zoological 

Society of London, 123(l):65-94, 3 plates. 

1967a. Variable Symmetry in Fissiparous Asterozoa. Symposium of the Zoological Society of London, 20:143-157. 

1967b. Notes on the Family Ophiothricidae (Ophiuroidea). Annals and Magazine of Natural History, series 

13, 9:637-655, illustrated. 

Clark, H. L. 

1901. The Echinoderms of Porto Rico. United States Fish 

Commission Bulletin for 1900, 2:231-263, plates 14-

17. 

1904. The Echinoderms of the Woods Hole Region. 

Bulletin, United States Fish Commission, 22:545-576, 

14 plates. 

1914. Growth-Changes in Brittle-Stars. Papers from the 

Tortugas Laboratory of the Carnegie Institution of Washington, 5(7):91-126, 3 plates. 

1915. Catalogue of Recent Ophiurans: Based on the 

Collection of the Museum of Comparative Zoology. Memoirs of the Museum of Comparative Zoology at 

Harvard College, 25(4): 165-376, 20 plates. 

1918. Brittle Stars, New and Old. Bulletin of the Museum 

of Comparative Zoology at Harvard College, 62:265-

338, 8 plates. 
NUMBER 12 409 

1919. The Distribution of the Littoral Echinoderms of 

the West Indies. Papers from the Department of Marine 

Biolqgy of the Carnegie Institution of Washington, 13(3): 

49-74, 3 plates. 

1921. The Echinoderm Fauna of Torres Strait: Its Composition and Its Origin. Department of Marine Biology 

of the Carnegie Institution of Washington, 10: 223 

pages, 38 plates. [Carnegie Institution of Washington 

Publication, 214.] 

1933. A Handbook of the Littoral Echinoderms of Porto 

Rico and the Other West Indian Islands. New York 

Academy of Sciences Scientific Survey of Porto Rico and 

the Virgin Islands, 16(1): 1-147, 7 plates. 

1939. Two New Ophiurans from the Smithsonian-Hartford Expedition, 1937. Proceedings of the United States 

National Museum, 86:415-418. 

1941. Reports on the Scientific Results of the Atlantis 

Expeditions to the West Indies, under the Joint 

Auspices of the University of Havana and Harvard University: The Echinoderms (Other than 

Holothurians). Memonas de la Sociedad Cubana de 

Historia Natural "Felipe Poey" 15:1-154, 10 plates. 

1942. The Echinoderm Fauna of Bermuda. Bulletin of the 

Museum of Comparative Zoology at Harvard College, 

89(8): 367-391, 1 plate. 

Coe, W. R. 

1912. Echinoderms of Connecticut. Connecticut State Geological and Natural History Survey Bulletin, 19:1-152, 

32 plates. 

Costa, H. R. da, and L. S. da Costa 

1962. Sobre as especies brasileiras do genero Ophioderma 

(Echinodermata, Ophiuroidea). Universidade do 

Brasil, Faculdade Nacional de Filosofia, Centro de Estudos Zoologicos, Avulso, 16:1-4, 1 plate, 

da Costa. See Costa, H. R. da 

Davis, W. P. 

1966. Observations on the Biology of the Ophiuroid 

Astrophyton muricatum. Bulletin of Marine Science, 

16(3):435-444. 

Delle Chiaje, S. 

1829. Memoria sulla storia et analomia degli animali senze 

vertebre del regno di Napoli. Volume 5. [Not seen], 

de Loriol. See Loriol, P. de 

Devaney, D. M. 

1970. Studies on Ophiocomid Brittlestars, I: A New 

Genus (Clarkcoma) of Ophiocominae with a Reevaluation of the Genus Ophiocoma. Smithsonian 

Contributions to Zoology, 51:1-41. 

1974. Shallow-Water Echinoderms from British Honduras, with a Description of a New Species of 

Ophiocoma (Ophiuroidea). Bulletin of Marine Science, 

24(1): 122-164, illustrated. 

Doderlein, L. 

1911. Beitrage zur Naturgeschichte Ostasiens: Uber japanische und andere Euryalae. Abhandlungen der 

Bayenschen Akademie der Wissenschaften, supplement, 

2(5): 1-123, 9 plates. 

1927. Indopacifische Euryalae. Abhandlungen der Bayerischen Akademie der Wissenschaften, 31(6): 1-105, 10 

plates. 

Duncan, C. P., D. K. Atwood, J. R. Duncan, and P. N. 

Froelich 

1977. Drift Bottle Returns from the Caribbean. Bulletin 

of Marine Science, 2 7 (3): 580-586. 

Engel, H. 

1939. Echinoderms from Aruba, Curacao, Bonaire and 

Northern Venezuela. In Zoologische Ergebnisse 

einer Reise nach Bonaire, Curacao und Aruba im 

Jahre 1930, No. 27. Capita Zoologica, 8(4): 1-12. 

Fell, H. B. 

1960. Synoptic Keys to the Genera of Ophiuroidea. 

Zoology Publications from Victoria University of Wellington, 26:1-44. 

1962. A Revision of the Major Genera of Amphiurid 

Ophiuroidea. Transactions of the Royal Society of New 

Zealand (Zoology), 2(1): 1-26, plate 1. 

Fontaine, A. R. 

1953. The Shallow-Water Echinoderms of Jamaica, Part 

II: The Brittle-Stars (Class Ophiuroidea). Natural 

History Notes, Natural History Society of Jamaica, 60: 

197-206. 

Forbes, E. 

1841. A History of British Starfish and Other Animals of the 

Class Echinodermata. 267 pages. London: John Van 

Voorst. 

Fricke, H.-W. 

1968. Beitrage zur Biologie der Gorgonenhaupter Astrophyton muricatum (Lamarck) und Astroboa nuda (Lyman) (Ophiuroidea, Gorgonocephalidae). 107 

pages. Ph.D. Thesis, Mathematisch-Naturwissenschaftliche Fakultat der Freien Universitat Berlin. 

Godcharles, M. F. 

1971. A Study of the Effects of a Commercial Hydraulic 

Clam Dredge on Benthic Communities in Estuarine Areas. State of Florida Department of Natural 

Resources, Technical Series, 64:1-51 pages. 

Godcharles, M. F., and W. C. Jaap 

1973. Fauna and Flora in Hydraulic Clam Dredge Collections from Florida West and Southeast Coasts. 

Florida Department of Natural Resources Marine Research Laboratory, Special Scientific Report, 40:1-89. 

Grave, C. 

1898. Embryology of Ophiocoma echinata, Agassiz, Preliminary Note. Johns Hopkins University Circulars, 18:6-

7, illustrated. 

1899. Notes of the Development of Ophiura olivacea Lyman. Zoologischer Anzeiger, 22:92-96, illustrated. 

Grube, A. E. 

1857. Diagnosen einiger neuen Echinodermen. Archiv fiir 

Naturgeschichte, 23:340-344. 
410 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Hendler, G. 

1975. Adaptational Significance of the Patterns of 

Ophiuroid Development. American Zoologist, 15(3): 

691-715. 

Ives, J. E. 

1889. Catalogue of the Asteroidea and Ophiuroidea in 

the Collection of the Academy of Natural Sciences 

of Philadelphia. Proceedings of the Academy of Natural 

Sciences of Philadelphia for 1889, 41:169-179. 

1890. Echinoderms from the Northern Coasts of Yucatan and the Harbor of Vera Cruz. Proceedings of the 

Academy of Natural Sciences of Philadelphia for 1890, 

42:317-340, plates 6-8. 

1891. Echinoderms from the Bahama Islands. Proceedings 

of the Academy of Natural Sciences of Philadelphia for 

1891, 43:337-341, plate 16. 

John, D. D., and A. M. Clark 

1954. The "Rosaura" Expedition, 3: The Echinodermata. Bulletin of the British Museum (Natural History), 

Zoology, 2(6): 139-162, plate 6. 

Koehler, R. 

1907. Revision de la collection des ophiures du Museum 

d'Histoire Naturelle de Paris. Bulletin Scientifique de 

la France et de la Beligique, 41:279-351, plates 10-

14. 

1913. Ophiures. Zoologische Jahrbucher Supplement, 11(3): 

351-380, plates 20, 21. 

1914. A Contribution to the Study of the Ophiurans of 

the United States National Museum. United Stales 

National Museum Bulletin, 84:1-173, 18 plates. 

Lamarck, J. B. 

1816. Histoire naturelle des animaux sans vertebres, pre'sentant 

les caracteres, generalaux et particuliers de ces animaux, 

leur distribution, leur classes, leur families, leur genres et 

la citation synonymique des principales especes qui s'y 

rapportent. 1st edition, volume 2, 568 pages. Paris: 

Verdiere. 

Leach, W. E. 

1815. Zoological Miscellany. Volume 2, 154 pages, 120 

plates. 

Ljungman, A. V. 

1866. Ophiuroidea viventia hue usque cognita enumerat. Ofversigt af Kongl. Velenskaps-Akademiens Forhandlingar, 9:303-336. 

1871. Fdrtechning ofver uti Vestindien af Dr. A. Goes 

samt under Korvetten Josefinas Expedition i Atlantiska Oceanen samlade Ophiurider. Ofversigt af 

Kongl. Vetenskaps-Akademiens Fdrhandlinger Stockholm, 

1871:615-658. 

Loriol, P. de 

1893. Echinodermes de la Baie d'Amboine. Revue Suisse 

de Zoologie, 1:359-426, plates 13-15. 

Ludwig, H. 

1882. Verzeichniss der von Prof. E. van Beneden an der 

Kiiste von Brasilien gesammelten Echinodermen. 

Memoires Couronne's et Memoires des Savants Etrangers 

Public's par L'Academie Royale des Sciences, des Lettres, 

et des Beaux-Arts de Belgique, 44:1:26. 

Liitken, C. F. 

1856a. Bidrag til Kundskab om Slangestijernerne, II: Oversigt over de Vestindiske Ophiurer. Videnskabelige 

Meddelser fra Dansk Naturhistorisk Forening i Kobenhaven, 1856:1-26. 

1856b. Bidrag til Kundskab om Slangestijernerne, III: 

Bidrag til Kundskab om Ophiurerne ved CentralAmerikas Vestkyst. Videnskabelige Meddelser fra 

Dansk Naturhistorisk Forening i Kobenhaven, 1856:20-

26. 

1859. Additamenta ad historiam Ophiuridarum, Anden 

Afdeling. Kongelige Dansk Videnskabernes Selskabs 

Skrifter, Femte Raekke, Naturviddenskabelig og mathematisk Afdeling, 5:75-271, plates 1-5. 

1869. Additamenta ad historiam Ophiuridarum: Beskrivende og kritiske Bidrag til Kundskab om Slangestjernerne, Tredie Afdeling. Kongelige Danske Videnskabernes Selskabs Skrifter, Femte Raekke, Naturvidenskabelig og mathematisk Afdelung, 8:24-109. 

Liitken, C. F., and Th. Mortensen. 

1899. Reports on an Exploration off the West Coasts of 

Mexico, Central and South America, and off the 

Galapagos Islands, in Charge of Alexander Agassiz, by the U.S. Fish Commission Steamer "Albatross," during 1891, Lieut. Commander Z. L. Tanner, U.S.N., Commanding, XXV: The Ophiuridae. Memoirs of the Museum of Comparative Zoology at 

Harvard College, 23:97-208, 23 plates. 

Lyman, Th. 

1860. Descriptions of New Ophiurans Belonging to the 

Smithsonian Institution and to the Museum of 

Comparative Zoology at Cambridge. Proceedings of 

the Boston Society of Natural History, 7:193-252. 

1865. Ophiuridae and Astrophytidae. In Illustrated Catalogue of the Museum of Comparative Zoology at 

Harvard College, Volume 1. Memoirs of the Museum 

of Comparative Zoology at Harvard College, 1(1): 1-200, 

2 plates. 

1869. Preliminary Report on the Ophiuroidae and Astrophytidae Dredged in Deep Water between 

Cuba and the Florida Reef. Bulletin of the Museum 

of Comparative Zoology at Harvard College in Cambridge, 

l(10):309-354. 

1875. Zoological Results of the Hassler expedition, II: 

Ophiuridae and Astrophytidae, Including Those 

Dredged by the Late Dr. William Stimpson. In 

Illustrated Catalogue of the Museum of Comparative Zoology at Harvard College, Volume 8. 

Memoirs of the Museum of Comparative Zoology at Harvard College, 4(11): 1-34, 5 plates. 
NUMBER 12 411 

1878. Reports on the Operations of the U. S. Coast 

Survey Steamer "Blake": Ophiurans and Astrophytons. Bulletin of the Museum of Comparative Zoology 

at Harvard College in Cambridge, 5(9):217-238, 3 

plates. 

1882. Report on the Ophiuroidea. In Report on the Scientific Results of the Voyage of H.M.S. Challenger during 

the Years 1873-76 under the Command of Captain George 

S. Nares, R.N., F.R.S., and Captain Frank Tourle 

Thomson, Zoology, 5(14): 1-386, 48 plates. Edinburg: 

Her Majesty's Stationery Office. 

1883. Reports on the Results of Dredging, under the 

Supervision of Alexander Agassiz, in the Caribbean Sea in 1878-79, and along the Atlantic Coast 

of the United States during the Summer of 1880, 

by the U. S. Coast Survey Steamer "Blake": Report on the Ophiuridae. Bulletin of the Museum of 

Comparative Zoology at Harvard College in Cambridge, 

10(6):227-287, 8 plates. 

1888. Ophiurans. In A. Agassiz, Three Cruises of the United 

States Coast and Geodetic Survey Steamer "Blake" in the 

Gulf of Mexico, in the Caribbean Sea, and along the 

Atlantic Coast of the United States, from 1877 to 1880, 

2:109-116, figures 338-403. Boston and New 

York: Houghton, Mifflin and Co. 

Marktanner-Tureretscher, G. 

1887. Beschreibung neuer Ophiuriden und Bemerkungen zu bekannten. Annalen des K. K. Nalurhistonschen Hofmuseums, 2:291 -316. 

Martens, E. von 

1870. Die Ophiuriden des indischen Oceans. Archiv fur 

Naturgeschichte, 36:245-262. 

Martin, R. B. 

1968. Aspects of the Ecology of Axiognathus squamata 

(Echinodermata, Ophiuroidea). Tane, the Journal of 

the Auckland University Field Club, 14:65-81. 

Matsumoto, H. 

1915. A New Classification of the Ophiuroidea. Proceedings of the Academy of Natural Sciences of Philadelphia, 

67:43-92. 

1917. A Monograph of Japanese Ophiuroidea Arranged 

according to a New Classification. Journal of the 

College of Science, Imperial University of Tokyo, 38:1-

408, 7 plates. 

Metcalf, W. G , M. C. Stalcup, and D. K. Atwood 

1977. Mona Passage Drift Bottle Study. Bulletin of Marine 

Science of the Gulf and Caribbean, 27:586-591. 

Millott, N. 

1953. A Remarkable Association between Ophionereis reticulata (Say) and Harmothoe lunulata (Delle Chiaje). 

Bulletin of Marine Science of the Gulf and Caribbean, 3: 

96-99. 

Mortensen, Th. 

1921. Studies of the Development and Larval Forms of Echinoderms. 261 pages, 102 plates. Copenhagen: G.E.C. 

Gad. 

1936. Echinoidea and Ophiuroidea. Discovery Reports, 12: 

199-348, 9 plates. 

Muller, J., and F. H. Troschel 

1842. System der Asteriden. 134 pages, 12 plates. Braunschweig: Friedrich Vieweg und Sohn. 

Nutting, C. C. 

1895. Narrative and Preliminary Report of Bahama Expedition. Bulletin from the Laboratories of Natural 

History of the State University of Iowa, 3:1-251. 

Parslow, R. E., and A. M. Clark 

1963. Ophiuroidea of the Lesser Antilles. In Studies on the 

Fauna of Curacao and other Caribbean Islands, 15:24-

50, illustrated. 

Pawson, D. L. 

1978. The Echinoderm Fauna of Ascension Island, 

South Atlantic Ocean. Smithsonian Contributions to 

the Marine Sciences, 2:1-31, 11 figures. 

Pearson, J.F.W. 

1936. Studies of the Life Zones of Marine Waters adjacent to Miami, I: The Distribution of the Ophiuroidea. Proceedings of the Florida Academy of Sciences, 

1:66-72. 

Rathbun, R. 

1879. A List of the Brazilian Echinoderms, with Notes 

on Their Distribution, etc. Transactions of the Connecticut Academy of Arts and Sciences, 5:139-158. 

Roa, E. Z. de 

1967. Contribucion al estudio de los equinodermos de 

Venezuela. Acta Biologica Venezuelica, Universidad 

Central de Venezuela, Facultad de Ciencias, 5:267-333, 

illustrated. 

Say, T. 

1825. On the Species of the Linnaean Genus Asterias, 

Inhabiting the Coast of the United States. Journal 

of the Academy of Natural Sciences at Philadelphia, 5: 

141-154. 

Smith, J. E. 

1938. Occurrence of Young Ophiothrix fragilis within the 

Genital Bursa of the Adult. Nature, 141:554. 

Stancyk, S. E. 

1973. Development of Ophiolepis elegans (Echinodermata: 

Ophiuroidea) and Its Implications in the Estuarine Environment. Marine Biology, 21(1): 7-12. 

Stearns, F. S. 

1891. List of the Asteroidea, Ophiuroidea, Echinoidea, 

and Crustacea, at Present in the Private Collection 

of Frederick Stearns, Detroit, Michigan, U.S.A., 

to Which Is Appended a Paper on "Echinoderms 

and Arthropods from Japan" by J. E. Ives, viii + 

14 pages, 6 plates. Published by the author. 

Tabb, D. C , and R. B. Manning 

1961. A Checklist of the Flora and Fauna of Northern 
412 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Florida Bay and Adjacent Brackish Waters of the 

Florida Mainland Collected during the period 

July, 1957 through September, 1960. Bulletin of 

Marine Science of the Gulf and Carribean, 11(4):552-

649, 8 figures. 

Thomas, L. P. 

1962a. The Shallow Water Amphiurid Brittle Stars (Echinodermata, Ophiuroidea) of Florida. Bulletin of 

Marine Science of the Gulf and Caribbean, 12(4):623-

694, illustrated. 

1962b. Two Large Ophiodermatid Brittle Stars New to 

Florida. Quarterly Journal of the Florida Academy of 

Sciences, 25(l):65-69, illustrated. 

1966. A Revision of the Tropical American Species of 

Amphipholis (Echinodermata: Ophiuroidea). Bulletin of Marine Science, 16(4):827-833. 

1973. Western Atlantic Brittle Stars of the Genus Ophionereis. Bulletin of Marine Science, 23(3):585-599, 

illustrated. 

Tommasi, L. R. 

1965. Alguns Amphiuridae (Ophiuroidea) do litoral de 

Sao Paulo e Santa Catarina. Contribuicoes Avulsas 

do Instituto Oceanografico, Universidad de Sao Paulo, 

Series Oceanografia-Biologica, 8:1-9, 5 figures. 

1970. Os ofiuroides do Brasil e de regios Vizinhas. Contribuicoes Avulsas do Instituto Oceanografico, Universidad 

de Sao Paulo, Series Oceanografia-Biologica, 20:1-146, 

46 plates. 

1971. Equinodermes do Brasil, I: Sobre algumas especies 

novas e outras pouco conhecidas, para o Brasil. 

Boletim do Instituto Oceanographico, Sao Paulo Universidade, 20:1-21, illustrated. 

Verrill, A. E. 

1867. Notes on the Radiata in the Museum of Yale 

College, with Descriptions of New Genera and 

Species, No. 3: On the Geographical Distribution 

of the Echinoderms of the West Coast of America. 

Transactions of the Connecticut Academy of Arts and 

Sciences, 1:323-351. 

1868. Notes on the Radiata in the Museum of Yale 

College with Descriptions of New Genera and 

Species, No. 4: Notice of the Corals and Echinoderms Collected by Prof. C. F. Hart, at the Abrolhos Reefs, Province of Bahia, Brazil, 1867. Transactions of the Connecticut Academy of Arts and Sciences, 

1(2):351-371. 

1899a. Report on the Ophiuroidea collected by the Bahama Expedition in 1893. Bulletin from the Laboratories of Natural History of the Slate University of Iowa, 

5(1): 1-86, 8 plates. 

1899b. North American Ophiuroidea, Part I: Revision 

of Certain Families and Genera of West Indian 

Ophiurans. Part II: A Faunal Catalogue of the 

Known Species of West Indian Ophiurans. Transactions of the Connecticut Academy of Arts and Sciences, 

10(2):301-386, plates 42, 43. 

von Martens. See Martens, E. von 

Zeiller, W. 

1974. Tropical Marine Invertebrates of Southern Florida and the 

Bahama Islands. New York, London, Sydney, Toronto: John Wiley and Sons. 132 pages. 

Ziesenhenne, F. C. 

1955. A Review of the Genus Ophioderma M. & T. In 

Essays in the Natural Sciences in Honor of Captain Allan 

Hancock on the Occasion of His Birthday July 26, 1955, 

pages 185-201. Los Angles: University of Southern California Press. 
Shallow-Water Crinoidea (Echinodermata) 

from Carrie Bow Cay, Belize 

D. Bradford Macurda, Jr. 

ABSTRACT 

Four species of shallow-water crinoids occur on 

the barrier reef off Carrie Bow Cay, Belize: Nemaster discoidea, N. rubiginosa, Ctenantedon kinziei, 

and Analcidometra armata. The distribution of these 

animals within the reef environment is determined by water movement and depth. The number of color variants of the two species of Nemaster 

is as high as found at most localities in the West 

Indies. The discovery of Ctenantedon kinziei extends 

considerably the range of this species. 

Introduction 

Although several species of crinoids have long 

been known from the Caribbean area (see, for 

example, Clark, 1921), the shallow-water crinoids 

of the West Indies were a virtually unknown part 

of the reef fauna until D. L. Meyer began to 

study them in the late 1960s. Work by Meyer and 

myself in the 1970s revealed the ubiquity of these 

animals from the Bahamas to the South American coast and from Central America to Barbados. 

Detailed studies have been published for the Bahamas (Macurda, 1973; 1975), Jamaica (Meyer, 

1973a), and Colombia (Meyer and Macurda, 

1976). More general accounts for the Caribbean 

are found in Meyer (1973b), for the West Indies 

in Macurda and Meyer (1977). The overall distribution of shallow- and deep-water crinoids of 

D. Bradford Macurda, Jr., Museum of Paleontology, University of 

Michigan, Ann Arbor, Mich. 48109. Mailing address: 10260 

Westheimer, Suite 110, Houston, Texas 77042. 

the West Indies was recently reviewed on the 

basis of extensive new collections made during 

the 1960s and 1970s (Meyer et al., 1978). 

Eight species of shallow-water crinoids belonging to five genera are found in the Caribbean. 

Nemaster is represented by three species: N. discoidea (Carpenter), N. grandis Clark, and N. rubiginosa 

(Pourtales); Comactinia by two: C. echinoptera 

(Muller) and C. meridionalis meridionalis (Agassiz); 

and the following genera by single species: Analcidometra armata (Pourtales), Ctenantedon kinziei 

Meyer, and Tropiometra carinata (Lamarck). David 

L. Meyer (pers. comm.) identified Neocomatella sp. 

from one shallow-water station in Panama but no 

one has found a representative of this genus again 

above 50 m. Tropiometra carinata, in the Caribbean, 

is found only in the southern and eastern parts, 

but it occurs elsewhere in the southwestern Atlantic. Shallow-water Comactinia meridionalis meridionalis is known only from Panama and the Carolinas. Nemaster grandis occurs in the western and 

central Caribbean. The remaining five species are 

found in reef environments throughout the West 

Indies. All eight species are illustrated in Macurda and Meyer (1977). 

ACKNOWLEDGMENTS.—I wish to thank B. W. 

Spracklin and K. M. Muzik for collecting, and 

C. G. Messing for identifying two specimens of 

Analcidometra armata that were found off Carrie 

Bow Cay subsequent to my visit. I am also indebted to D. L. Meyer for extensive discussions 

of crinoid taxonomy and ecology. This research 

was conducted under National Science Foundation Grant GB-36439. 

413 
414 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Carrie Bow Cay Crinoids 

Dredging does not permit accurate determination of the taxonomic composition and relative 

abundance of crinoids in reef environments because of the cryptic habits of many species. Divers 

using SCUBA must make transects for direct 

observation and counts, and break apart coral 

rock to find cryptic species, such as Ctenantedon 

kinziei and Comactinia echinoptera. Most of the following data for the Carrie Bow Cay area were 

obtained from 11 dives on the fore reef east of 

South Water Cay, Carrie Bow Cay, and Curlew 

Bank (see Riitzler and Macintyre, herein, Figure 

2), between 24 and 29 April 1973. Four species of 

crinoids are present on this section of the Belizean 

barrier reef: Nemaster discoidea, N. rubiginosa, Ctenantedon kinziei, and Analcidometra armata. The first 

two are far more abundant than the latter two. 

Analcidometra, a small crinoid that lives on gorgonians (primarily Pseudopterogorgia spp.), was not 

seen during my visit but was later collected twice 

on the fore reef east of Carrie Bow Cay. Another 

nearby record of this species is from Isla Roatan, 

Bay Island, Honduras, reported by Macurda and 

Meyer (1977). These authors also found Comactinia echinoptera and Nemaster grandis in the Bay 

Islands; neither species has yet been obtained in 

Belize. 

The profiles surveyed for crinoids run perpendicular to the barrier reef. There is an inner fore 

reef gradually deepening from the reef crest to 13 

m; the bottom then slopes into a sand trough (23 

m), rises again to an outer ridge (13 m), and then 

drops at an angle of 40°-50° along the fore-reef 

slope to 50 m before becoming vertical (Riitzler 

and Macintyre, herein: 9, Figure 5). The distance 

from the reef crest to the sand trough is 300 m, 

the remainder of the profile covers an additional 

100 m. 

Both Nemaster species are abundant on the forereef slope and on the top of the outer ridge but 

not on its shoreward face, the sand trough, or the 

deeper part of the inner reef slope. They become 

abundant again in the spur and groove zone of 

the inner fore reef between 10 and 15 m, but are 

not found shallower than this. The depth distribution of these species reflects their need of moderate water motion without direct wave surge. 

Commensals include ophiuroids, myzostomes, 

and shrimps. 

Nemaster discoidea and N. rubiginosa are rheophobic crinoids that hide the calyx beneath a coral 

or within a crevice and then extend some of the 

arms into the surrounding water in order to catch 

microscopic particles. A variety of substrates is 

used to hide the disc. Pinnules are regularly arrayed in a tetrad; the arms do not form a regular 

TABLE 30.—Depth distribution of color varieties oi Nemaster discoidea and N. rubiginosa on outer 

ridge and fore-reef slope off Curlew Bank and South Water Cay (counts are based on one 

transect at each location, approximately 2 m wide and conducted over a 45 min dive period; 

first number of each pair of values is for Curlew Bank) 

Color variety 

Nemaster discoidea 

Dark green 

Gold 

Yellow with black stripe 

N. rubiginosa 

Orange with black stripe 

Orange with yellow-tipped 

pinnules 

Orange with white-tipped 

pinnules 

10-15 

2;0 

19;23 

10;4 

2;2 

Depth rar 

15-20 

5;0 

6;1 

1;0 

20-25 

3;1 

ige (m) 

25-30 

0;9 

3;2 

1A3 

0;1 

30-35 

i;i 

l;3 

1;0 

35-40 

0;7 

Totals 

1;10 

7;13 

8;3 

26;25 

10;4 

3;2 
NUMBER 12 415 

TABLE 31.—Arm length and arm number of color varieties 

of Nemaster discoidea and N. rubiginosa from Carrie Bow Cay 

Color variety 

Nemaster discoidea 

Dark green 

Gold 

Yellow with black 

N. rubiginosa 

Orange with black 

stripe 

stripe 

Orange with yellow-tipped 

pinnules 

Orange with white-tipped 

pinnules 

Average 

arm 

length 

(mm) 

127 

117 

130 

141 

84 

70 

Average 

arm 

number 

17 

18 

19 

21 

21 

21 

Sample 

size 

4 

11 

9 

41 

24 

10 

fan as in rheophilic crinoids (see Macurda and 

Meyer, 1977, figs. 1, 2). Arm lengths are commonly asymmetric owing to restricted space for 

some of the arms to grow. Nemaster rubiginosa, 

predominant in shallow water, is gradually replaced by N. discoidea with depth. Several persistent, easily recognizable color varieties of each are 

known and also present off Carrie Bow Cay (Tables 30, 31). There are three distinct color varieties 

of Af. discoidea: a form with dark green arms, a 

form with gold arms, and a form with yellow 

arms with an intermittent black stripe. All have 

black-tipped pinnules. The dark green variety is 

found on the fore-reef slope between 18 and 40 m 

(maximum observed range is larger than transect 

data given in Table 30). The gold form ranges 

from 13 m on top of the outer ridge (very rare) to 

40 m depth on the fore-reef slope; it is the most 

abundant variety. The unusual yellow form with 

black stripes occurs on the fore-reef slope between 

13 and 28 m. The arms of this species are always 

extended into the surrounding water. 

Nemaster rubiginosa, the most abundant crinoid, 

occurs primarily on the outer ridge and in the 

spur and groove zone (Plate 4: bottom right). It 

appears in at least seven color varieties, of which 

only three are common (Table 30): orange with 

black stripe, orange with yellow-tipped pinnules, 

and orange with white-tipped pinnules. The 

black-striped form is the most common one and 

is particularly abundant on the outer ridge. It is 

the only variety to extend its range down the forereef slope, to 33 m, although it is rare below 16 

m. The forms with yellow and white pinnule tips 

are more common at the spur and groove zone 

than on the outer ridge. The yellow-tipped variety is the more abundant but also the more 

cryptic of the two. The arms of N. rubiginosa are 

stouter than those of N. discoidea and may reach 

300 mm in length, although 200 mm was the 

longest observed off Belize. There, this species is 

mostly confined to heads of the coral Montastrea 

annularis (Ellis and Solander), unlike Panama 

where it occurs among Acropora cervicornis (Lamarck). 

Ctenantedon kinziei is rare (five individuals observed) but its discovery in Belize is important 

because this species was unknown until Meyer 

(1972) described it from the southern Caribbean. 

We have since discovered it throughout the West 

Indies. In Belize, it occurs from on top of the 

outer ridge and down the fore-reef slope, from 13 

to 25 m, completely hidden within coral heads 

such as are formed by the species of Montastrea. 

Two specimens of Analcidometra armata were 

discovered on gorgonians after my survey of 1973. 

One was found on Munceopsis sp. on the fore-reef 

slope in 20 m (B. W. Spracklin, May 1975), the 

second on Gorgonia mariae Bayer in the low-relief 

spur and groove zone in 11 m (K. M. Muzik, 

April 1979). 

Conclusion 

My investigations revealed that only four of 

the five most common Caribbean shallow-water 

crinoids are present at Carrie Bow Cay. Similar 

work should be conducted at other localities to 

ascertain the possible presence of additional species before the results of this study can be accepted 

as a definitive reflection of crinoid diversity on 

the Barrier reef of Belize. 
416 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Literature Cited 

Clark, A. H. 

1921. Report on the Crinoids Collected by the BarbadosAntigua Expedition in 1918. University of Iowa 

Studies in Natural History, 9(5):3-28. 

Macurda, D. B., Jr. 

1973. Ecology of Comatulid Crinoids at Grand Bahama 

Island. Hydro-Lab Journal, 2:9-24. 

1975. The Bathymetry and Zoogeography of ShallowWater Crinoids in the Bahama Islands. Hydro-Lab 

Journal, 3:5-24. 

Macurda, D. B., Jr., and D. L. Meyer 

1977. Crinoids of West Indian Coral Reefs. In S. H. 

Frost, M. P. Weiss, and J. B. Saunders, editors, 

Reefs and Related Carbonates—Ecology and Sedimentology. Studies in Geology, 4:231-237, 4 figures. Tulsa, Oklahoma: The American Association of Petroleum Geologists. 

Meyer, D 

1972. 

1973a. 

1973b. 

Meyer, D 

1976 

Meyer, D 

1978 

Ctenantedon, a New Antedonid Crinoid Convergent 

with Comasterids. Bulletin of Marine Science, 22:53-

66. 

Distribution and Living Habits of Comatulid Crinoids near Discovery Bay, Jamaica. Bulletin of 

Marine Science, 23:244-259. 

Feeding Behavior and Ecology of Shallow Water 

Unstalked Crinoids (Echinodermata) in the Caribbean Sea. Marine Biology, 22:105-129. 

. L., and D. B. Macurda, Jr. 

Distribution of Shallow-Water Crinoids near 

Santa Marta, Colombia. Mitteilungen, Instituto Colombo-Aleman Investigaciones Cientificas, 8:141-156. 

. L., C. G. Messing, and D. B. Macurda, Jr. 

Zoogeography of Tropical Western Atlantic Crinoidea (Echinodermata). Bulletin of Marine Science, 

28:412-441. 
Chemical Defense in Tropical Marine Algae 

James N. Norris 

and William Fenical 

ABSTRACT 

Chemical and taxonomic studies of the benthic 

algae of Carrie Bow Cay, Belize, have revealed 

that species of certain families of the Rhodophyta, 

Phaeophyta, and Chlorophyta produce unique or 

unusual secondary compounds. Analysis of the 

structural organic chemistries of these natural 

products shows that the substances produced consist largely of halogenated and non-halogenated 

terpenoids including monoterpenoids (Cio compounds), sesquiterpenoids (C15 compounds), and 

diterpenoids (C20 compounds), and larger compounds of mixed acetate-mevalonate origin. 

These secondary compounds, hypothesized to be 

feeding deterrents developed as defenses against 

herbivores, were examined for antibiotic activity 

and toxicity. Fish toxicity experiments using Eupomacentrus leucostictus and these compounds revealed sublethal to lethal effects. One compound, 

elatol, from Laurencia obtusa was found to inhibit 

sea urchin egg development totally. Field observations indicate that marine algae having these 

secondary metabolites are not eaten by many 

herbivores, and biological activity testing suggests 

that certain of these compounds may be responsible. In some cases, specialized grazers have apparently co-evolved to tolerate these potential 

chemical "deterrents," and may in turn use them 

in their own defense against predation. 

Introduction 

Land plants have evolved elaborate morphological and chemical defense mechanisms against 

James N Norris, Department of Botany, National Museum of Natural 

History, Smithsonian Institution, Washington, D.C. 20560; and 

William Fenical, Institute of Marine Resources, Scripps Institution 

of Oceanography, Lajolla, Calif 92093. 

herbivores (Fraenkel, 1959; Whittaker and Feeney, 1971). Since Janzen (1973) put forth the 

working hypothesis that the primary role of secondary compounds in higher plant vegetation 

and seeds is defense against herbivores and microorganisms, it has been generally recognized that 

these natural products may either serve as feeding 

deterrents or attractants in terrestrial plant-animal interactions (see, for example, Harbourne, 

1977), or function as allelopathic chemicals 

(Muller, 1970) or antibiotics. The same evolutionary pressures responsible for the many biologically active compounds found in terrestrial vegetation have been predicted to have parallels in 

marine vegetation (Whittaker and Feeney, 1971; 

Kittredge, 1976). Predaceous fishes and invertebrate animals have been important forces acting 

as agents of natural selection in the evolution of 

protective mechanisms (Bakus, 1964; 1966; 1969). 

The world-wide distribution of herbivorous 

marine fishes shows a concentration, both in species diversity and biomass, on tropical reefs (Hiatt 

and Strasburg, 1960). Herbivorous fishes generally dominate fish communities on tropical reefs 

(Bardach, 1959; Hiatt and Strasburg, 1960); and 

the number and biomass of both vertebrate and 

invertebrate herbivores are two of the striking 

features of Caribbean coral reefs (Ogden, 1976). 

Marine algae have developed several defenses 

in response to herbivory (see for example, Paine 

and Vadas, 1969; Vadas, 1977; 1979). Algal species may deter herbivores in one or more of the 

following ways: (1) by having a resistant or unpalatable physical structure (for example, the 

calcareous nature of some Rhodophyta, Chloro417 
418 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

phyta, and Phaeophyta, or the tough texture of 

certain Phaeophyta), or a morphology that makes 

it difficult for the herbivore to attach to feed (see 

Hay, 1981); (2) by residing in habitats that are 

cryptic or unfavorable for herbivore feeding (Ogden et al., 1973; Adey and Vassar, 1975), such as 

in crevices where they are inaccessible, in areas of 

high surf or strong surge where herbivores cannot 

attach themselves for feeding, or in areas where 

herbivores are subject to increased predation; (3) 

by having heteromorphic life histories, with a fast 

growing stage and a resistant stage (Lubchenco 

and Cubit, 1980); (4) by living in association with 

toxic or unpalatable algae (see plant defense 

guilds of Atsatt and O'Dowd, 1976); (5) by being 

unpredictable in occurrence in time or space (for 

example, ephemeral species, or species having 

rare or patchy distribution) (Littler and Littler, 

1980); and/or (6) by producing secondary metabolites (Fenical, 1975), ranging from unpalatable 

to toxic, as chemical defense against herbivores. 

Since the pioneering work of Pratt et al. (1951) 

reporting antibacterial activity of various seaweed extracts, several investigators have demonstrated the antibiotic activity of extracts from 

some tropical algae (for example, Burkholder et 

al., 1960; Olesen et al., 1964; Sieburth, 1964; 

Burkholder, 1973; Bhakuni and Silva, 1974; Nunez and Serpa Sanabria, 1975). Studies of Boyd 

et al. (1966) on the effects of selected tropical 

algae on human erythrocytes found that the extracts from six brown algae agglutinated blood 

groups O and A. More recently, Targett and 

Mitsui (1979) studied the effects of aqueous extracts from tropical algae on fish erythrocyte 

hemolysis and fish mortality, and Targett (1979) 

developed a behavioral bioassay (gastropod tentacle withdrawal) to test extracts from marine 

algae for biological activity. In another study, 

both seasonal variability and locality differences 

of the tropical algal extract's antibiotic activity 

against selected pathogenic bacteria was observed 

(Almodovar, 1964). Hornsey and Hide (1974) 

found similar seasonal variation in their screening 

experiments for antimicrobial activity of British 

marine algae. Sieburth (1968) offered some ecological interpretation of the role of algal antibiotics in marine algal-algal and algal-animal interactions. Unfortunately, there have been few 

quantitative studies on the diets of marine herbivores, and our knowledge of feeding behavior 

and food preferences of tropical herbivores is still 

too inadequate to be conclusive. What is known 

on tropical fish diets can be found in Randall 

(1967), while sea urchin diets have been summarized by Lawrence (1975). 

In this paper we provide an overview of interactions between tropical reef algae and herbivores 

in the Caribbean, with reference to our identification of specific compounds from Belize reef 

algae. On the basis of the antimicrobial activity 

and fish toxicity of these compounds we present 

hypotheses of their ecological role in reef algalanimal interactions. 

Over the past four years at Carrie Bow Cay, 

off the coast of Belize, we have investigated the 

systematics of the benthic algae as well as the 

chemical nature of their secondary compounds. 

Some of the Chlorophyta, Phaeophyta, and Rhodophyta synthesized unusual secondary products 

which in the laboratory showed biological activity 

against selected micro-organisms and a reef fish. 

Many of the relatively abundant algae that were 

accessible to potential herbivores were found to 

contain these biologically active substances. We 

hypothesize that these compounds have been selected for by the intense grazing pressure on 

tropical marine algae and that they function to 

minimize population losses by inhibiting or repelling herbivores. Our present analysis indicates 

that tropical reef algae are chemically defended 

from many generalist herbivores, while some specialist herbivores have evolved a physiological 

tolerance to these natural products. 

ACKNOWLEDGMENTS.—We thank our colleagues 

who have discussed aspects of this research with 

us. Particularly helpful were the reviews of I. 

Abbott, W. Adey, S. Brawley, K. Bucher, D. 

Cheney, J. Cubit, M. Hay, R. Houbrick, M. 

Littler, and R. Vadas, and the editoral comments 

of V. Macintyre. We especially thank R. Jacobs 

(University of California, Santa Barbara) for providing the data from his experiment for Figure 

182. " 
NUMBER 12 419 

Materials and Methods 

Field trips were mad e to Carrie Bow Cay on 

the barrier reef of Belize (16°48'N; 88°05'W), 

during the spring (March-May ) of 1976, 1977, 

and 1978 (for a review of the biological and 

geological features of Carrie Bow Cay, see Riitzler 

and Macintyre , herein: 9). During each expedition, benthic macro-algae were collected for subsequent studies on their systematics and chemistry. Homogeneous collections were identified and 

separated into specimens for chemical analysis 

and vouchers for taxonomic verfication; the former were preserved in isopropanol, the latter in 

4% Formalin-seawater. 

Voucher specimens have been deposited in the 

U.S. National Herbarium, Smithsonian Institution. Th e following algal species were chemically 

investigated in this study. Chlorophyta: Caulerpa 

cupressoides (Vahl) C. Agardh, C. mexicana (Sonder) 

Kiitzing, C. racemosa (Forsskal) J. Agardh, C. serrulata (Forsskal) J. Agardh emend B0rgesen, C. 

sertularioides (Gmelin) Howe , C. verticillata J. 

Agardh, Rhipocephalus phoenix (Ellis and Solander) 

Kiitzing, and Udotea flabellum (Ellis and Solander) 

Howe. Phaeophyta : Dictyota bartayresii Lamouroux, Stypopodium zonale (Lamouroux) Papenfuss, 

Sargassum polyceratim var. ovatum (Collins) Taylor, 

and Turbinaria turbinata (Linnaeus) Kuntze . Rhodophyta: Liagora farinosa Lamouroux, Asparagopsis 

taxiformis (Delile) Trevisan; Ochtodes secundiramea 

(Montagne) Howe , Laurencia caraibica Silva, and 

L. obtusa (Hudson) Lamouroux. 

T h e biological activity of the secondary compounds produced by the marine algae was investigated by nutrient agar plate disc assay methods 

in the laboratory of W. Fenical, and by fish 

toxicity experiments. Antibacterial activity was 

examined on the basis of inhibition oi Staphylococcus aureus Rosenbach, Bacillus subtilis (Ehrenberg) 

Cohn, and Escherichia coli (Migula) Castellani and 

Chalmers, and antifungal activity was assessed 

against the huma n pathogen Candida albicans (Robin) Berkhout. Toxicity to fish was tested using 

Eupomacentrus leucostictus (Muller and Troschel) 

with serial dilutions of the alga's natura l compounds in seawater (as outlined by Sun and 

Fenical, 1979), and the crude extract. In each 

treatment (three replicates), either the extract or 

the pure compoun d was added directly to the 

seawater with the aid of an ethanol dispersant; 

controls were untreated seawater an d seawater 

containing only the ethanol. Fish mortality 

within one hour was considered to indicate a 

toxic compound, and sublethal effects, such as 

loss of equilibrium and respiratory stress were 

noted as "strong" or "mild. " Results of the antimicrobial and ichthyotoxic experiments are summarized in Tabl e 32. 

Results and Discussion 

CHLOROPHYTA.—Of the benthic algae in the 

vicinity of Carrie Bow Cay (Norris and Bucher, 

herein: 167), the green algae are perhaps the most 

abundan t in biomass, having representatives 

throughout the diverse intertidal and subtidal 

habitats. Thre e families predominate , the Codiaceae with species of Avrainvillea and Codium, 

Udoteaceae with Halimeda, Penicillus, Rhipocephalus, and Udotea, and the Caulerpaceae with 

species of its monotypic genus, Caulerpa. 

T h e species we investigated from these families 

produced unusual secondary metabolites. Of six 

species of Caulerpa from Carrie Bow Cay, C. cupressoides, C mexicana, C racemosa, C. serrulata, C. 

sertularioides, an d C verticillata, all but C. mexicana 

contained the known compoun d caulerpin (Figure 181a, compound I). This substance was originally isolated from C racemosa, C. serrulata, an d 

C. sertularioides by Aguilar-Santos and Doty 

(1968). In recent studies of C. taxifolia (Vahl) C. 

Agardh, Maiti and Thomson (1977) re-examined 

the structure of caulerpin (Aguilar-Santos, 1970) 

and found it to be a derivatized indole dimer. 

We found that species of Caulerpa, Halimeda, 

Penicillus, Rhipocephalus, and Udotea also produce 

terpenoids of rather complex structure. Although 

it is difficult to analyze man y of these compounds 

thoroughly because of their instability, we have 

isolated from Rhipocephalus phoenix two sesquiterpenoids, rhipocephenal and rhipocephalin, designated as compounds II and III respectively 

(Figure 1816). These compounds typify the struc-
420 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 32.—Compounds identified from tropical benthic algae of Carrie Bow Cay, and results 

of biological activity tests (compound designations identified in Figures 181, 182; 0 = no 

response, + = positive response, NT = not tested) 

Species 

CHLOROPHYTA 

Caulerpa cupressoides 

C. racemosa 

C. serrulata 

C. sertularioides 

C. verticillata 

Rhipocephalus phoenix 

R. phoenix 

PHAEOPHYTA 

Sargassum polyceratinum var. ovatum 

Turbinaria turbinata 

Dictyota bartayresii 

Stypopodium zonale 

RHODOPHYTA 

Asparagopsis taxiformis 

Ochtodes secundiramea 

0. secundiramea 

Liagora farinosa 

Laurencia caraibica 

L. obtusa 

Compound 

I 

I 

I 

I 

I 

II 

III 

IV 

IV 

V 

VI 

VII 

VIII 

IX 

X 

XI 

XII 

Antimicrobial 

activity 

0 

0 

0 

0 

0 

+ 

+ 

+

3 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

+

4 

+ 

Toxicity 

to fish 

NT 

+ ' 

NT 

NT 

NT 

+

2 

+ 

NT 

NT 

0 

+ 

NT 

NT 

NT 

+ 

NT 

NT 

1

 Lewin, 1970

 2 Sun and Fenical, 1979 3 Glombitza, 1977 4 Izac, 1979 

tural types we have detected in the Caulerpaceae 

and in some species of the Codiaceae and Udotaceae. Compounds II and III were ichthyotoxic 

at the 2 and 10 /xg/ml levels, respectively, against 

Eupomacentrus leucostictus; when these compounds 

were placed in food offered to this fish it was not 

eaten (Sun and Fenical, 1979). We are continuing 

studies with Halimeda species, Udotea flabellum, U. 

conglutinata (Ellis and Solander) Lamouroux, Penicillus capitatus Lamark, and Avrainvillea longicaulis 

(Kiitzing) Murray and Boodle, each of which 

appears to contain unique and biologically active 

secondary metabolites. 

Our field observations indicate that the tropical algae of these three families are avoided by 

most herbivores, at least by the "generalists" 

(sensu Emlen, 1973). Few quantitative studies 

have been done on food preferences of tropical 

fish and invertebrates. Those dealing with the 

herbivores and green algae we studied (see Lawrence, 1975; Ogden and Lobel, 1978) noted that 

the urchin Echinometra lucunter (Linnaeus) near St. 

Croix, Virgin Islands, does not eat Caulerpa and 

Halimeda, even though they are available in small 

amounts on substrate (Juliana and Ambrosetti, 

1974; Ogden, 1976) and as drift (D. Abbott et al., 

1974). Rates of feeding by Lytechinus variegatus 

(Lamarck) varied in laboratory experiments, with 

Caulerpa being eaten least rapidly when offered 

alone and most rapidly when offered with five 

other marine plants; Halimeda was consistently 

the least preferred alga (Lawrence, 1975). Lowe 

and Lawrence (1976) suggest that L. variegatus in 

Florida may be feeding preferentially on detrital 

seagrasses. Similarly, Vadas et al. (in press) 

showed the preferred diet of field populations of 

L. variegatus was detrital Thalassia, while siphonaceous greens were largely not eaten (except by 

one population in which approximately 30% of 

their diet consisted of these algae). Lowe and 

Lawrence (1976) suggested a mixed diet might be 

superior in nutrients for L. variegatus. 
NUMBER 12 421 

CHO 

CHO 

OAc 

VI VII •COOH 

AA/ =V

=

\ =/ 

f 

* 

VIII 

CHBrXI 

FIGURE 181.—Structures of biologically active metabolites isolated from Carrie Bow Cay marine 

algae: a, caulerpin (compound I) from species of Caulerpa; b, sesquiterpenoids, rhipocephenal 

(II) and rhipocephalin (III), from Rhipocephalus phoenix; c, polyphenol (IV) from Sargassum and 

Turbinaria; d, pachydictyol A (V) from Dictyota bartayresii; e, stypotriol (VI) from Stypopodium 

zonale; fi acetylene containing lipid (VII) from Liagora farinosa; g, bromoform (VIII) from 

Asparagopsis taxiformis; h, polhyalogenated monoterpenoids, ochtodene (IX) and ochtodial (X), 

from Ochtodes secundiramea; i, allolaurinterol (XI) from Laurencia caraibica. 

Lewis (1958) suggested that Tripneustes ventricosus (Lamarck) refused Halimeda because of its 

calcified structure, but we suggest this rejection 

may also be due to the chemical component of 

Halimeda. Green algae not eaten by the urchin 

Diadema antillarum Philippi included Penicillus 

(Ogden et al., 1973; Ogden, 1976) and Halimeda 

(Ogden, 1976). Feeding preferences of sea urchins 
422 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

have been summarized by Lawrence (1975) and 

studied by Vadas (1977), Larson et al. (1980), 

and Vadas et al. (in press). 

Of the Chlorophyta in fish-feeding trials, Earle 

(1972) observed that Codium and Halimeda were 

rapidly consumed, only portions of Caulerpa and 

Udotea were eaten, and Penicillus and Avrainvillea 

were largely not eaten. Similarly, Mathieson et 

al. (1975) found that Avrainvillea nigricans Descaisne was mostly not eaten, that most fish, except 

the parrotfish, did not eat Halimeda, that only the 

Grey Angelfish consumed Caulerpa, and that Penicillus capitatus and Udotea conglutinata remained 

uneaten at the end of their feeding trials. Tsuda 

and Bryan (1973) noted that the siganid fishes 

readily consumed Caulerpa during feeding trials. 

It is difficult to generalize about tropical herbivores and their feeding habits and preferences on 

the basis of such scant information. Of the greens 

we studied, however, most are apparently not 

eaten by many herbivores in habitat. 

A few herbivores apparently can physiologically cope with, or perhaps even selectively eat, 

algae having chemical defenses. Sacoglossan opisthobranchs are a notable exception to the general avoidance of Caulerpa and members of the 

Codiaceae as a food source. In fact, bivalved 

sacoglossans are restricted to Caulerpa, while nonshelled sacoglossans have been reported in association with a variety of algae that include several 

species of Codiaceae (MacNae, 1954), and a few 

other Chlorophyta and Vaucheria (Chrysophyta) 

(see Kay, 1968). 

The association between these algae and sacoglossans in the Caribbean has been reported from 

Puerto Rico (Warmke and Almodovar, 1963), 

and in the Pacific from Japan (Hamatani, 1972) 

and Fiji (Burn, 1966). The distribution of sacoglossans appears dependent on the range of their 

food source, Caulerpa (Burn, 1966), with members 

of the family Juliidae reflecting the distribution 

oi Caulerpa throughout the Indo-West Pacific, the 

Caribbean, the Mediterranean and Victoria, Australia (Kay, 1968). 

Specialist herbivores were relatively common 

at Carrie Bow Cay. Found on Caulerpa racemosa 

were shelled sacoglossans, Lobiger souverben 

(Fischer) and Oxynoe species, and the non-shelled 

sacoglossans, Elysia cauzae (Er. Marcus), Elysia 

species, and Volvatella species (the latter two and 

the Oxynoe may represent undescribed species, J. 

R. Lance, pers. comm.). 

Some sacoglossans not only ingest intact chloroplasts from Caulerpa and utilize their products 

(see Taylor, 1968; Greene, 1974; Trench, 1975), 

but ingest some of the alga's secondary metabolites as well. Caulerpicin and caulerpin were identified from Elysia panamensis Pilsbry and Olsson 

(Doty and Aguilar-Santos, 1970) collected off 

Baja California feeding on Caulerpa sertularioides. 

This sacoglossan can secrete a milky mucus that 

is toxic to fish (Lewin, 1970). The biological 

activity of caulerpin (Figures 181a, compound I) 

and caulerpicin (Doty and Aguilar-Santos, 1966) 

was reported by Doty and Aguilar-Santos (1970) 

to be toxic to, and physiologically active in, mice 

and rats, respectively. 

While Caulerpa, Halimeda, Udotea, Penicillus, Rhipocephalus, and Avrainvillea have evolved secondary 

compounds that deter a wide variety of herbivores, the sacoglossans have evolved a physiological mechanism /or tolerating the chemical deterrents produced by the algae, and thus have become "specialists" feeding on a restricted but 

exclusive diet. It also appears that these specialists 

deter predation upon themselves by concentrating these compounds. More than three times the 

amount of caulerpicin and more than twice the 

amount of caulerpin were found concentrated in 

the sacoglossan Elysia panamensis than were found 

in the algal host, Caulerpa (Doty and AguilarSantos, 1970). 

PHAEOPHYTA.—The abundant brown algae at 

Carrie Bow Cay belong to two families, the Sargassaceae and the Dictyotaceae. Species of Sargassum and Turbinaria (Sargassaceae) are apparently not eaten by some herbivores, at least in 

areas where these browns are common. 

In the Caribbean, Sargassum and Turbinaria 

sometimes grow in areas of the reef where herbivores may have difficulty reaching them or remaining attached to feed. Also, in preliminary 
NUMBER 12 423 

subtidal feeding trials and translocation experiments of Sargassum in Caribbean Panama (J. 

Cubit and M. Hay, pers. comm.) and Islas San 

Bias (J. Cubit, pers. comm.), these algae were 

readily consumed, particularly by kyphosid 

fishes. These observations suggest that herbivory 

may limit the lower level of distribution of certain 

Sargassum species. It is also possible that increased 

predation or other factors may keep these fish 

from the reef where Sargassum grows; we also 

suggest that members of this family may not be 

eaten owing to their inherent high concentrations 

of polyphenolic substances based upon the polymerization of phloroglucinol (Figure 18 k, compound IV). Types of biological activity shown by 

polyphenols include antibacterial (Conover and 

Sieburth, 1964) and antilarval (Conover and Sieburth, 1966). These polyphenols may also be 

ecologically important in plant-plant interactions; growth of certain phytoplankton and macroalgae may be inhibited (McLachlan and Craigie, 1964) or enhanced (Ragan et al., 1980) by 

polyphenols. 

The presence and absence of these compounds 

in the Sargassaceae have been reviewed by Glombitza (1977). Pfeffer (1963) suggested that the 

inability of fish to digest Sargassum was due to its 

tannins (Ogino, 1963). From stomach analysis of 

tropical reef fish, Randall (1967) concluded Sargassum was eaten only by a few larger herbivorous 

and omniverous fishes—Kyphosus incisor (Cuvier 

and Valenciennes), K. sectatrix (Linnaeus), Pomacanthus arcuatus (Linnaeus), P. para (Bloch), and 

Melichthys niger (Bloch)—and that these kyphosids 

and M. niger feed on drifting algae. Earle (1972) 

suggested it may be eaten by certain fishes, and 

W. H. Adey (pers. comm.) reports that the bucktooth parrotfish, Sparisoma radians (Cuvier and 

Valenciennes), readily consumes Sargassum. Atkinson et al. (1973) found that Sargassum, though 

available, was not present in the guts oi Diadema 

antillarum (see also Lawrence, 1975), while another 

sea urchin, Paracentrotus lividus (Lamarck) has been 

recorded as feeding on Sargassum (Lawrence, 

1975). Sammarco et al. (1974) observed lush 

growth of Tubinaria on a patch reef after they 

removed Diadema; a diet of Turbinaria also provided the best growth for Tripneustes of the algae 

tested in St. Croix (J. Ogden, R. Vadas, and S. 

Miller, pers. comm.). 

The common browns at Carrie Bow Cay belong to the Dictyotaceae; members of Dictyota, 

Dictyopteris, Lobophora, Padina, and Stypopodium are 

represented. Although Padina and Lobophora species are not noticeably grazed in the vicinity of 

Carrie Bow Cay, we have not found in them the 

unusual organic compounds that are present in 

other tropical members of the family; therefore, 

structure or some other factor may be important, 

such as increased predation upon potential herbivores, or possibly water soluble compounds or 

secondary compounds below our level of detection may be present. Interestingly, a Pacific species of Lobophora from Palau was found to contain 

unusual compounds (Fenical, in progress). 

Of the several species of Dictyota occurring in 

the vicinity of Carrie Bow Cay (Norris and 

Bucher, herein: 167), we investigated the most 

predominant one, D. bartayresii, which grows 

among corals in the spur and groove area of the 

barrier reef. Recognized in the field by its marked 

iridescence, D. bartayresii in our chemical studies 

was shown to contain a series of toxic bicyclic 

diterpenoids, the major component being the 

compound pachydictyol A (Figure 18 Id, compound V), previously described by Hirschfeld et 

al. (1973) from another Dictyotaceae genus, Pachydictyon (for review of Dictyotaceae diterpenoid 

synthesis see McEnroe et al., 1977). Although 

some sea urchins have been noted to feed on some 

species oi Dictyota (Atkinson et al., 1973; Abbott 

et al., 1974; Lawrence, 1975), as yet none are 

known to feed specifically on D. bartayresii. This 

alga has been recorded in stomach contents of 

three species offish (Randall, 1967; Earle, 1972). 

It is also possible that pachydictyol A may not 

function to deter urchin or fish predators, but 

might be an example of a compound that acts 

against certain micro-organisms. This metabolite's toxicity to certain fungi suggests it could be 

defense against microbial pathogens (Fenical, unpublished data). 
424 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Another predominant brown alga at Carrie 

Bow Cay is Stypopodium zonale, which grows primarily in open, shallow areas (4-10 m depths) 

where it is accessible to grazers. Frequently to 20 

cm in length and conspicuous in its environment, 

S. zonale is apparently not eaten by most herbivores. When freshly collected S. zonale is placed in 

cool seawater for 10 hours, the water becomes 

dark brown, apparently from release of pigments 

and secondary compounds. These substances 

were toxic to the herbivorous damsel fish, Eupomacentrus leucostictus, found at Carrie Bow Cay, 

and exhibited antimicrobial activity in preliminary testing. Ethanol extracts of S. zonale were 

equally toxic to damsel fish at levels of approximately 3 /ig/ml. The toxic components have been 

found to consist of a mixture of several related 

C27 compounds derived from a mixed biosynthesis 

of diterpenoid and acetate precursors. One of 

these compounds, the triol stypotriol (Figure 18 k, 

compound VI), has been isolated and structurally 

defined (Gerwick and Fenical, 1980). We are not 

aware of published records of S. zonale as a food 

source for fishes or urchins. As in the green algasacoglossan relationship, certain mollusks may 

have co-evolved as specialist predators of S. zonale. 

Recent collections of this alga from the Florida 

Keys contained large numbers of the sea hare 

Petalifera petalifera Ranger (Fenical, unpublished 

data), but it was not observed on any other algae 

and may represent another specialist with a preferred or exclusive food source. 

RHODOPHYTA.—Although they have the largest 

number of species in the environs of Carrie Bow 

Cay, the red algae are generally smaller in size 

and individual species are less abundant than 

either the green or brown algal species. Red algae 

that are apparently avoided by herbivores at 

Carrie Bow Cay belong to four families: Helminthocladiaceae, Bonnemaisoniaceae, Rhizophyllidaceae, and Rhodomelaceae. Members of 

the latter three families are known to produce 

elaborate halogenated, often toxic, metabolites 

(Fenical, 1975). With the recent discovery of 

unique compounds in Liagora farinosa (Paul and 

Fenical, 1980), the Helminthocladiaceae was 

added to the expanding list of families known to 

produce secondary compounds. 

The moderately calcified red, Liagora farinosa 

(Helminthocladiaceae), was seasonally abundant 

in the spur and groove habitats of the barrier reef 

at Carrie Bow Cay. We noted unusual compounds present on TLC plates, and recently Paul 

and Fenical (1980) described the major metabolite, an unusual acetylene containing lipid (Figure 

181/, compound VII), which occurs along with 

several minor, related compounds in this species. 

These compounds were observed to be toxic 

against Eupomacentrus leucostictus at the 5-8 jug/ml 

range in seawater. We did not observe any grazers 

on L. farinosa. 

Despite its fine, delicate structure, Asparagopsis 

taxiformis (Bonnemaisoniaceae) does not seem to 

be utilized as a food source. Asparagopsis species 

produce unique toxins, the major metabolite 

being the noxious compound bromoform (Figure 

181^, compound VIII) (McConnell and Fenical, 

1976). Several halogenated acetones are also produced by A. taxiformis (compounds of this type 

have been produced synthetically and used as 

tear gas). 

Asparagopsis was not eaten by the fish Acanthurus 

triostegus (Linnaeus) (Randall, 1961), nor has it 

been reported as being grazed by sea urchins 

(Lawrence, 1975). We suggest it is not eaten by 

most herbivores because of its toxic metabolites. 

These compounds inhibited the growth of all 

micro-organisms tested, and they were severe lacrymators. Interestingly, though toxic or unpalatable to most grazers, it is consumed by man, for 

Asparagopsis taxiformis is the favorite seaweed food 

of the Hawaiians (Abbott and Williamson, 1974). 

Falkenburgia hillebrandii (Ardissone) Falkenberg, 

the alternate sporophytic stage in the life history 

oi Asparagopsis taxiformis (Chihara, 1961), has also 

been collected at Carrie Bow Cay. This morphologically different stage also appears to be avoided 

by grazers. In other studies, McConnell and Fenical (unpublished data) have found Falkenburgia 

from the North Atlantic also to contain bromoform. We suspect that the Carrie Bow Cay Falkenburgia, which is generally not eaten by herbi-
NUMBER 12 425 

vores, also produces this compound. Fish have 

not been observed to feed on this stage of Asparagopsis (Earle, 1972), but it was found in 1% of 

Diadema antillarum guts by Atkinson et al. (1973). 

Since that was such a small sample, it may have 

indirectly been consumed while feeding primarily 

on other turf-forming algae. This possibility, however, will have to be further investigated. 

Another red alga containing toxic secondary 

compounds is Ochtodes secundiramea of the Rhizophyllidaceae. Ochtodes secundiramea grows in shallow water inside the reef crest at South Water 

Cay, in areas of strong wave agitation and surge. 

This alga contains one major and one minor 

polyhalogenated monoterpenoid, octodene and 

ochtodial (Figure 18IA, compounds IX and X) 

(McConnell and Fenical, 1978). Compound IX, 

the major metabolite, is strongly antibiotic and 

made up over 50% of the total organic extract of 

the alga. During our field studies on freshly collected specimens, we observed iridescence from 

the large "glandular cells" or "Driisenzellen" 

(Kylin, 1956) present in the cortex of 0. secundiramea (Joly and Ugadim, 1966). Whether these 

refractive bodies are the site of halogenation in 

0. secundiramea is uncertain, although we observed 

these bodies were no longer iridescent after the 

methanol extraction and they appeared to be 

"empty" on examination with a Wild M5 stereomicroscope. We are not aware of any published 

accounts of Ochtodes as a food source; we did not 

find it being grazed during our daytime observations in Belize. 

Three species of the cosmopolitan genus Laurencia (Rhodomelaceae) from Carrie Bow Cay 

contained interesting secondary metabolites. This 

genus is known to produce a complex variety of 

halogen-containing compounds (Fenical and 

Norris, 1975; Fenical, 1975). We conducted chemical studies on the two abundant species, Laurencia 

caraibica and L. obtusa (identified following Taylor, 

1960:626), at Carrie Bow Cay. The third species, 

L. intricata Lamouroux, was not abundant enough 

for further study, even though an initial TLC 

indicated the presence of secondary compounds. 

Laurencia caraibica Silva grows sympatrically 

with L. obtusa in areas of intense water motion, 

generally along the outer reef crest. This alga 

produces quantities of the antibiotic cuparane 

derivative allolaurinterol (Figure 18h, compound 

XI), a compound which has also been isolated as 

a minor component of L. subopposita (J. Agardh) 

Setchell from California (Wratten and Faulkner, 

1977) and L. filiformis (C. Agardh) Montagne 

from the east coast of Australia (Kazlauskas et 

al., 1976). Izac and Sims (1979) discovered a 

unique iodinated sesquiterpene in L. caraibica, the 

first report of such a compound and of iodine 

being found in the genus. We found the same 

major metabolite in this species, and did not 

observe this alga being eaten by fish or urchins. 

Laurenica obtusa was found in large quantities 

along the Carrie Bow Cay reef crest (at least 

during April and May). We found this alga to 

contain more than 3% (dry weight) of the cytotoxic chamigrene derivative elatol (Figure 182, 

compound XII), a compound originally isolated 

from the Australian species L. elata (C. Agardh) 

Hooker and Harvey (Sims et al., 1974). Elatol has 

subsequently been found to be moderately antibiotic, but more interestingly this compound 

shows exceptional toxicity against fertilized sea 

urchin eggs, and totally inhibits cell division (FigHO

 elato 1 

100 

2 4 8 16 

Bath Concentration (pg/ml) 

FIGURE 182.—Dose-response curve of elatol (compound XII) 

from Laurencia obtusa, against fertilized eggs of the sea urchin, 

Stronglyocentrotus purpuratus (the seven markers represent individual experiments). 
426 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ure 182). The dose-response curve of elatol with 

fertilized eggs from Stronglylocentrotus purpuratus 

(Stimpson), a California urchin, shown in Figure 

182, has been provided by R. Jacobs. The LD50 

of elatol is estimated at about 7 jug/ml bath 

concentration. Laurenica obtusa is not grazed by 

Diadema (Ogden, 1976), and is avoided by most 

herbivores. It is interesting to speculate on the 

ecological role of compounds such as elatol, which 

are now known to inhibit, or, at certain levels, to 

kill developing sea urchin eggs. We hypothesize 

that if such compounds are released or secreted 

in the vicinity of Laurencia stands, they could 

inhibit sea urchin egg development in situ, and 

thus give the alga a selective advantage against 

the settlement and development of potential predators in their vicinity. 

In view of the biologically active constituents 

from Laurencia, it is not surprising that the alga 

has few predators. The sea hares (Anaspidea), 

particularly some species of Aplysia, have specific 

feeding preferences for Laurencia. In our field studies in the Gulf of California, California, the Galapagos Islands and now in Belize, we have observed this grazer-alga relationship between Laurencia and Aplysia. During our spring studies at 

Carrie Bow Cay, we typically found A. dactylomela 

(Ranger) grazing on L. obtusa on the fore-reef 

crest. It appears that some Aplysia species may be 

dependent on Laurencia, at least during certain 

stages of their life cycle, as larvae and juveniles. 

Culture studies on A. californica Cooper (Kriegstein et al., 1974) indicate that settling and metamorphosis of the free-swimming larval stage are 

enhanced by the presence of Laurencia pacifica. 

Stallard and Faulkner (1974) showed that Aplysia californica concentrated the metabolites of Laurencia pacifica in their digestive glands. We have 

found quantities of elatol, identified from L. obtusa 

in our study, in the digestive glands of A. dactylomela. It has been proposed that the concentration 

of the algal metabolites by the sea hare provide 

a selective advantage against potential predators 

(Stallard and Faulkner, 1974; Kittredge et al., 

1974). Although the defense role of this concentration needs to be tested, known predators of 

Aplysia are few, and opisthobranchs in general are 

avoided as a food source by Caribbean fish (Randall, 1967). Opisthobranch gastropods used in 

feeding-acceptability tests with selected fish by 

Thompson (1960) were almost invariably refused. 

Recently, it was suggested that juveniles of the 

tropical bridled burrfish, Chilbmycterus antennatus 

(Cuvier), in Caribbean Panama, are Batesian 

mimics of Aplysia dactylomela and perhaps avoid 

predation by mimicking the shape and coloration 

of the "unpalatable sea hare" (Heck and Weinstein, 1978). We suggest that the avoidance could 

be due to the concentration of elatol by A. dactylomela from Laurencia obtusa. 

Summary and Conclusions 

Our studies of Belize macro-algae, indicate that 

members of specific algal families—the Caulerpaceae, Codiaceae, Udoteaceae, Dictyotaceae, 

Sargassaceae, Helminthocladiaceae, Bonnemaisoniaceae, Rhizophyllidaceae, and Rhodomelaceae—produce unusual secondary compounds. 

The species we studied are not eaten by many 

herbivorous fishes or sea urchins. These herbivores and the coral reef algae on which they feed 

appear to represent a co-evolved system of defense 

and counterdefense. Chemical defense appears 

successfully to deter the majority of herbivores; 

however, some specialist grazers have co-evolved 

physiological mechanisms that enable them to 

tolerate or possibly even select for some of these 

same algae. In some specific animal-plant interactions, for example, saccoglossan-CWfrjfra and 

Aplysia-Laurencia, the mollusks not only exclusively select certain algal species for food and 

show preference for others but they also concentrate the alga's secondary compounds as defense 

against being preyed upon. Thus, it appears the 

chemical deterrent against most herbivores may 

have become a chemical attractant to these specialist species. Published evidence and our own 

observations, though scant, indicate most herbivorous reef fishes are highly selective in the algae 

they consume. In tropical reef algal-animal relationships, there is strong evolutionary interaction 

between herbivores and algae, based on the alga's 

defenses—structure (Lubchenco, 1978), productivity and growth form (Littler, 1980), life history 

strategy (Lubchenco and Cubit, 1980), habitat, 
NUMBER 12 427 

and chemistry—and the herbivore's feeding 

habits (preference for, or not eating of, algae and 

the peculiarities of their digestive physiology). 

Ehrlich and Raven (1965) first attributed to 

secondary compounds of plants a key role in 

determining the pattern of insect-plant co-evolution. As suggested by Feeny (1975), we may be 

witnessing an "evolutionary arms race" in which 

the algae must deploy part of their metabolic 

budget on defense against herbivory, and the 

herbivores must devote a portion of their assimilated energy on ways to locate their algal food, 

know which species to avoid, and on developing 

counter measures to tolerate the alga's chemical 

defense (see for example feeding preference studies of Vadas, 1977; Larson et al., 1980; for additional interpretation of "evolutionary arms race" 

theory, see Atsatt and O'Dowd, 1976). 

The subject of tropical reef algal-animal interactions presents a myriad of problems to be investigated and questions to be answered. Recognizing the speculative nature of this study we 

concur with Janzen (1977) that there is "quite 

enough hypothetical biology on the books" and 

we now need quantitative ecological and behavioral studies "on the pragmatics" of tropical 

plant-animal interactions. 

Literature Cited 

Abbott, D. P., I. A. Abbott, G. Juliana, J. Amborsetti, J. C. 

Ogden, and K. Warner 

1974. Drifting Macroscopic Plants as Sources of Food 

for the Sea Urchin Echinometra lucunter at Boiler 

Bay, St. Croix. In D. P. Abbott, J. C. Ogden, and 

I. A. Abbott, editors, Studies on the Activity Pattern, Behavior, and Food of the Echinoid Echinometra lucunter (Linnaeus) on Beachrock and Algal 

Reefs at St Croix, U.S. Virgin Islands. West Indies 

Laboratory, Special Publication, 4:78-86. St. Croix, 

U.S. Virgin Islands. 

Abbott, I. A., and E. H. Williamson » 

1974. Limu: An Ethnobotanical Study of Some Edible Hawaiian 

Seaweeds. 21 pages. Lawai, Kauai, Hawaii: Pacific 

Tropical Botanical Garden. 

Adey, W. H., and J. M. Vassar 

1975. Colonization, Succession and Growth Rates of 

Tropical Crustose Coralline Algae (Rhodophyta, 

Cryptonemiales). Phycologia, 14(2):55-69. 

Aguilar-Santos, G. 

1970. Caulerpin, a Red Pigment from Green Algae of 

the Genus Caulerpa. Journal of the American Chemical 

Society, section c, 92:842. 

Aguilar-Santos, G , and M. S. Doty 

1968. Chemical Studies on Three Species of the Marine 

Algal Genus Caulerpa. In H. D. Freudenthal, editor, 

Proceedings of the Third Conference on Food and Drugs 

from the Sea, pages 173-176. Marine Technology 

Society. 

Almodovar, L. R. 

1964. Ecological Aspects of Some Antibiotic Algae in 

Puerto Rico. Botanica Marina, 6:143-146. 

Atkinson, C , S. Hopley, L. Mendelsohn, and S. Yacowitz 

1973. Food Studies on Diadema antillarum on a Patch 

Reef, St. Croix, U.S. Virgin Islands. In J. C. 

Ogden, D. P. Abbott, and I. A. Abbott, editors, 

Studies on the Activity and Food of the Echinoid 

Diadema antillarum Philippi on a West Indian Patch 

Reef. West Indies Laboratory, Special Publication, 9: 

65-80. St. Croix, U.S. Virgin Islands. 

Atsatt, P. R., and D. J. O'Dowd 

1976. Plant Defense Guilds. Science, 193:24-29. 

Bakus, G J. 

1964. The Effects of Fish Grazing on Invertebrate Evolution in Shallow Tropical Waters. Allan Hancock 

Foundation Publications, Occasional Paper, 27:1-29. 

1966. Some Relationships of Fishes to Benthic Organisms on Coral Reefs. Nature, 210(5033):280-284. 

1969. Energetics and Feeding in Shallow Marine Waters. International Review of General and Experimental 

Zoology, 4:275-369, New York: Academic Press, 

Inc. 

Bardach, J. E. 

1959. The Summer Standing Crop of Fish on a Shallow 

Bermuda Reef. Limnology and Oceanography, 4(1): 

77-85. 

Bhakuni, D. S., and M. Silva 

1974. Biodynamic Substances from Marine Flora. Botanica Marina, 17:40-51. 

Boyd, W. C , L. R. Almodovar, and L. G. Boyd 

1966. Agglutinins in Marine Algae for Human Erythrocytes. Transfusion, 6(l):82-83. 

Burkholder, P. R. 

1973. The Ecology of Marine Antibiotics and Coral 

Reefs. In O. A. Jones, and R. Endean, editors, 
428 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Biology and Geology of Coral Reefs, pages 117-282. 

New York: Academic Press. 

Burkholder, P. R., L. M. Burkholder, and L. R. Almodovar 

1960. Antibiotic Activity of Some Marine Algae of 

Puerto Rico. Botanica Marina, 2:149-156. 

Burn, R. 

1966. The Opisthobranchs of a Caulerpin Microfauna 

from Fiji. Proceedings of the Malacological Society of 

London, 37:45-65. 

Chihara, M. 

1961. Life Cycle of the Bonnemaisoniaceous Algae in 

Japan, 1. Science Reports of the Tokyo Kyoiku Daigaku, 

section B, 10(153): 121-154, plates 1-6. 

Conover, J. T., and J. McN. Sieburth 

1964. Effects of Sargassum Distribution on Its Epibiota 

and Antibacterial Activity. Botanica Marina, 6:147— 

157. 

1966. Effect of Tannins Excreted from Phaeophyta on 

Planktonic Animal Survival in Tide Pools. In E. 

G. Young and J. L. McLachlan, editors, Proceedings 

of the Fifth International Seaweed Symposium, pages 

99-100. Oxford: Pergamon Press. 

Doty, M. S., and G. Aguilar-Santos 

1966. Caulerpicin, a Toxic Constituent of Caulerpa. Nature, 211(505 2): 990. 

1970. Transfer of Toxic Algal Substances in Marine 

Food Chains. Pacific Science, 24(3):351-355. 

Earle, S. A. 

1972. The Influence of Herbivores on the Marine Plants 

of Great Lameshur Bay, with an Annotated List 

of Plants. In B. B. Collette and S. A. Earle, editors, 

Results of the Tektite Program: Ecology of Coral 

Reef Fishes. Natural History Museum, Los Angeles 

County, Science Bulletin, 14:17-44. 

Ehrlich, P. R., and P. H. Raven 

1965. Butterflies and Plants: A Study in Coevolution. 

Evolution, 18:586-608. 

Emlen, J. M. 

1973. Ecology, an Evolutionary Approach. 493 pages. Reading, Mass.: Addison-Wesley Publishing Company, 

Inc. 

Feeny, P. 

1975. Biochemical Coevolution between Plants and 

Their Insect Herbivores. In L. E. Gilbert, and P. 

H. Raven, editors, Co-evolulion of Animals and Plants, 

pages 3-19. Austin: University of Texas Press. 

Fenical, W. 

1975. Halogenation in the Rhodophyta: A Review. Journal of Phycology, 11:245-259. 

Fenical, W., and J. N. Norris 

1975. Chemotaxonomy in Marine Algae: Chemical Separation of Some Laurenica Species (Rhodophyta) 

from the Gulf of California. Journal of Phycology, 11: 

104-108. 

Fraenkel, G. S. 

1959. The Raison d'Etre of Secondary Plant Substances. 

Science, 129:1466-1470. 

Gerwick, W. H., and W. Fenical 

1980. Ichthyotoxic and Cytotoxic Metabolites of the 

Tropical Brown Alga, Stypopodium zonale. Journal of 

Organic Chemistry, 46:22-27. 

Glombitza, K. W. 

1977. Highly Hydroxylated Phenols of the Phaeophyceae. In D. J. Faulkner and W. H. Fenical, editors, 

Marine Natural Products Chemistry, pages 191-204. 

New York: Plenum Press. 

Greene, R. W. 

1974. Sacoglossans and Their Chloroplast Endosymbionts. In W. B. Vernberg, editor, Symbiosis in the 

Sea, pages 21-27. Columbia, South Carolina: University of South Carolina Press. 

Hamatani, I. 

1972. A New Species of Volvatella Pesae 1860 Found in 

the Caulperin Microfauna in the Province of Kaii, 

Middle Japan. Publications of the Seto Marine Biological Laboratory, 21(1): 13-20. 

Harbourne, J. B. 

1977. Introduction to Ecological Biochemistry, xiv + 243 

pages. London: Academic Press. 

Hay, M. E. 

1981. The Functional Morphology of Turf-forming 

Seaweeds: Persistence in Stressful Marine Habitats. Ecology, 62(3):739-750. 

Heck, K. L., and M. P. Weinstein 

1978. Mimetic Relationship between Tropical Burrfishes and Opisthobranchs. Biolropica, 10(1):78—79. 

Hiatt, R. W., and D. W. Strasburg 

1960. Ecological Relationships of the Fish Fauna on 

Coral Reefs of the Marshall Islands. Ecological 

Monographs, 30(1):65-127. 

Hirschfeld, D. R., W. Fenical, G.H.Y. Lin, R. M. Wing, and 

P. C. Radlick 

1973. Pachydictyol A, a Unique Diterpene Alcohol from 

the Brown Seaweed Pachydictyon coriaceum. Journal 

of the American Chemical Society, 95:4049. 

Hornsey, I. S., and D. Hide 

1974. The Production of Antibiotic Compounds by British Marine Algae, I: Antibiotic Producing Marine 

Algae. British Journal of Phycology, 9:353-361. 

Izac, R. R. 

1979. Secondary Metabolites from the Red Alga Laurencia. Ph.D. dissertation. University of California, 

Riverside. 

Izac, R. R., and J. J. Sims 

1979. Marine Natural Products, 18: Iodinated Sesquiterpenes from the Red Algal Genus Laurencia. Journal of the American Chemical Society, 101:6136-6137. 
NUMBER 12 429 

Janzen, D. L. 

1973. Community Structure of Secondary Compounds. 

In T. Swain, editor, Chemistry in Evolution and Systematics, pages 529-538. London: Butterworths. 

1977. Promising Directions of Study in Tropical AnimalPlant Interactions. Annals of the Missouri Botanical 

Garden, 64:706-736. 

Joly, A. B., and Y. Ugadim 

1966. The Reproduction of Ochtodes secundiramea (Montagne) Howe (Gigartinales, Rhizophyllidaceae). 

Boletim do Instituto Oceanografico, Universidad de Sao 

Paulo, 15(l):55-64. 

Juliana, G , and J. Ambrosetti 

1974. Feeding and Diet in Echinometra lucunter. In D. P. 

Abbott, J. C. Ogden, and I. A. Abbott, editors, 

Studies on the Activity Pattern, Behavior, and 

Food of the Echinoid Echinometra lucunter (Linnaeus) on Beachrock and Algal Reefs at St. Croix, 

U.S. Virgin Islands. West Indies Laboratory, Special 

Publication, 4:65-77. St. Croix, U.S. Virgin Islands. 

Kay, E. A. 

1968. A Review of the Bivalved Gastropods and a Discussion of Evolution within the Sacoglossa. In V. 

Fretter, editor, Studies in the Structure, Physiology and Ecology of Molluscs. Symposia of the Zoological Society of London and the Malacological Society of 

London, 22:109-134. New York: Academic Press, 

Inc. 

Kazlauskas, R., P. T. Murphy, R. J. Quinn, and R. J. Wells 

1976. New Laurene Derivatives from Laurencia filiformis. 

Australian Journal of Chemistry, 29:2533. 

Kittredge, J. S. 

1976. Behavioral Bioassays and Biologically Active 

Compounds. In H. H. Webber and G D. Ruggieri, 

editors, Proceedings of the Fourth Conference on Food 

and Drugs from the Sea, pages 467-475. Marine 

Technology Society. 

Kittredge, J. S., F. T Takahashi, J. Lindsey, and R. Lasker 

1974. Chemical Signals in the Sea: Marine Allelochemics and Evolution. Fishery Bulletin, 72(1): 1-11. 

Kriegstein, A. R., V. Castellucci, and E. R. Kandel 

1974. Chemical Signals in the Sea: Marine Allelochemics and Evolution. Fishery Bulletin, 72(1): 1-11. 

Kriegstein, A. R., V. Castellucci, and E. R. Kandel 

1974. Metamorphosis oi Aplysia californica in Laboratory 

Culture. Proceedings of the National Academy of Sciences, USA, 71(9):3654-3658. 

Kylin, H. 

1956. Die Gattungen der Rhodophyceen. Frontispiece + xv 

+ 660 pages. Lund: C.W.K. Gleerups. 

Larson, B. R., R. L. Vadas, and M. Keser 

1981. Feeding and Nutritional Ecology of the Sea Urchin, Strongylocentrotus drobachiensis, in Maine, 

U.S.A. Marine Biology, 59:49-62. 

Lawrence, J. M. 

1975. On the Relationships between Marine Plants and 

Sea Urchins. In H. Barnes, editor, Oceanography and 

Marine Biology, Annual Review, 1975, 13:213-286. 

London: George Allen and Unwin, Ltd. 

Lewin, R. A. 

1970. Toxic Secretion and Tail Autonomy by Irritated 

Oxynoe panamensis. Pacific Science, 24:356-359. 

Lewis, J. B. 

1958. The Biology of the Tropical Sea Urchin, Tripneustes 

esculentus Leske in Barbados, British West Indies. 

Canadian Journal of Zoology, 36:607-621. 

Littler, M. M. 

1980. Morphological Form and Photosynthetic Performances of Marine Macroalgae: Tests of a Functional Form Hypothesis. Botanica Marina, 22:161-

165. 

Littler, M. M., and D. S. Littler 

1980. The Evolution of Thallus Form and Survival 

Strategies in Benthic Marine Macroalgae: Field 

and Laboratory Tests of a Functional Form 

Model. The American Naturalist, 116(1):25—44. 

Lowe, E. F., and J. M. Lawrence 

1976. Absorption Efficiencies of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea) for Selected 

Marine Plants. Journal of Experimental Marine Biology 

and Ecology, 21:223-234. 

Lubchenco, J. 

1978. Plant Species Diversity in a Marine Intertidal 

Community: Importance of Herbivore Food Preference and Algal Competitive Abilities. The American Naturalist, 112:23-39. 

Lubchenco, J. L., and J. E. Cubit 

1980. Effects of Herbivores on Heteromorphology in 

Some Marine Algae. Ecology, 61(3):676-687. 

McConnell, O. J., and W. Fenical 

1976. Halogen Chemistry of the Red Alga Asparagopsis. 

Phytochemistry, 16:367. 

1978. Ochtodene and Ochtodiol: Novel Polyhalogenated Cyclic Monoterpenes from the Red Seaweed, 

Ochtodes secundiramea. Journal of Organic Chemistry, 

43:4238-4341. 

McEnroe, F. J., R. J. Robertson, and W. Fenical 

1977. Diterpenoid Synthesis in Brown Seaweeds of the 

Family Dictyotaceae. In W. Fenical and D. J. 

Faulkner, editors, Marine Natural Products Chemistry, 

pages 179-189. New York: Plenum Press. 

McLachlan, J., and J. S. Craigie 

1964. Algal Inhibition by Yellow Ultraviolet-Absorbing 

Substances from Fucus vesiculosus. Canadian Journal 

of Botany, 42:287-292. 

MacNae, W. 

1954. On Four Sacoglossan Molluscs New to South Africa. Annates of the Natal Museum, 13:51-64. 
430 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Maiti, B. C , and R. H. Thomson 

1977. Caulerpin. In D. J. Faulkner and W. H. Fenical, 

editors, Marine Natural Products Chemistry, pages 

159-164. New York: Plenum Press. 

Mathieson, A. C , R. A. Fralick, R. Burns, and W. Flashive 

1975. Phycological Studies during Tektite II, at St. John 

U.S.V.I. In S. A. Earle and R. J. Lavenberg, 

editors, Results of the Tektite Program: Coral 

Reef Invertebrates and Plants. Natural History Museum, Los Angeles County, Science Bulletin, 20:77-103. 

Muller, C. H. 

1970. Phytotoxins as Plant Habitat Variables. Recent 

Advances in Phytochemistry, 3:106-121. 

Nunez, G , and L. M. Serpa Sanabria 

1975. Effectos producidos por extractos de algas marina 

macroscopicas sobre el crecimiento bacteriano. 

Acta Biologia Venezuela, 9(1): 121-134. 

Ogden, J. C. 

1976. Some Aspects of Herbivore-Plant Relationships on 

Caribbean Reefs and Seagrass Beds. Aquatic Botany, 2:103-116. 

Ogden, J. C , R. A. Brown, and N. Salesky 

1973. Grazing by the Echinoid Diadema antillarum Philippi: Formation of Halos around West Indian 

Patch Reefs. Science, 182:715-717. 

Ogden, J. C , and P. S. Lobel 

1978. The Role of Herbivorous Fishes and Urchins in 

Coral Reef Communities. Environmental Biology of 

Fishes, 3(l):49-63. 

Ogino, C. 

1962. Tannins and Vacuolar Pigments. In R. Lewin, 

editor, Physiology and Biochemistry of Algae, pages 

437-443. New York: Academic Press, Inc. 

Olesen, P. E., A. Maretzki, and L. A. Almodovar 

1964. An Investigation of Antimicrobial Substances 

from Marine Algae. Botanica Marina, 6:224-232. 

Paine, R. T., and R. L. Vadas 

1969. Calorific Values of Benthic Marine Algae and 

Their Postulated Relationship to Invertebrate 

Food Preference. Marine Biology, 4:79-86. 

Paul, V., and W. Fenical 

1980. Toxic Acetylene Containing Lipids from the Red 

Alga Liagora farinosa Lamouroux. Tetrahedron Letters, 1980(21):3327-3330. 

Pfeffer, R. 

1963. The Digestion of Algae by Acanthurus sandricensis. 

47 pages. Masters thesis, University of Hawaii, 

Honolulu. 

Pratt, R. H., H. Mutner, G. M. Gardner, Y. Sha, and Y. 

Dufrenoy 

1951. Report on Antibiotic Activity of Seaweed Extracts. 

Journal of the American Pharmaceutical Association, 40: 

575-579. 

Ragan, M. A., C. M. Ragan, and A. Jensen 

1980. Natural Chelators in Seawater: Detoxification of 

Zn2+ by Brown Algal Polyphenols. Journal of Experimental Marine Biology and Ecology, 44:261-267 

Randall, J. E. 

1961. Overgrazing of Algae by Herbivorous Marine 

Fishes. Ecology, 42:812. 

1967. Food Habits of Reef Fishes of the West Indies. 

Studies in Tropical Oceanography, 5:655-897. 

Sammarco, P. W., J. S. Levinton, and J. C. Ogden 

1974. Grazing and Control of Coral Reef Community 

Structure by Diadema antillarum Philippi (Echinodermata: Echinoidea): A Preliminary Study. Journal of Marine Research, 32:47-53. 

Sieburth, J. McN. 

1964. Antibacterial Substances Produced by Marine Algae. Developments in Industrial Microbiology, 5:124-

134. 

1968. The Influence of Algal Antibiosis on the Ecology 

of Marine Microorganisms. In M. R. Droop, and 

E. J. Ferguson Wood, editors, Advances in Microbiology of the Sea, pages 63-94. New York: Academic 

Press. 

Sims, J. J., G.H.Y. Lin, and R. M. Wing 

1974. Elatol, a Halogenated Sesquiterpene Alcohol from 

the Red Alga Laurencia elata. Tetrahedron Letters, 

1974:3487-3490. 

Stallard, M. O., and D. J. Faulkner 

1974. Chemical Constituents of the Digestive Gland of 

the Sea Hare Aplysia californica, I: Importance of 

Diet. Comparative Biochemistry and Physiology, 49B: 

25-36. 

Sun, H., and W. Fenical 

1979. Rhipocephenal and Rhipocephalin, Novel Linear 

Sesquiterpenoids from the Tropical Green Alga 

Rhipocephalus phoenix. Tetrahedron Letters, 1979(8): 

685-688. 

Targett, N. M. 

1979. Gastropod Tentacle Withdrawal: A Screening 

Procedure for Biological Activity in Marine Macroalgae. Botanica Marina, 22:543-545. 

Targett, N. M., and A. Mitsui 

1979. Toxicity of Subtropical Marine Algae Using Fish 

Mortality and Red Blood Cell Hemolysis for 

Bioassays. Journal of Phycology, 15:181-185. 

Taylor, D. L. 

1968. Chloroplasts as Symbiotic Organelles in the Digestive Gland oi Elysia viridis (Gastropoda: Opisthobranchia). Journal of the Marine Biological Association, 

United Kingdom, 48:1-15. 

Taylor, W. R. 

1960. Marine Algae of the Eastern Tropical and Sub-tropical 

Coasts of the Americas, ix + 870 pages. Ann Arbor: 

University of Michigan Press. 

Thompson, T. E. 

1960. Defensive Adaptations in Opisthobranchs. Journal 

of the Marine Biological Association, United Kingdom, 

39:123-134. 
NUMBER 12 431 

Trench, R. K. 

1975. Of "Leaves that Crawl": Functional Chloroplasts 

in Animal Cells. In D. H. Jennings and D. L. Lee, 

editors, Symbiosis, pages 229-265. Cambridge University Press. 

Tsuda, R. T., and P. G. Bryan 

1973. Food Preferences of Juvenile Siganus rostralus and 

S. spinus in Guam. Copeia, 1973:604-606. 

Vadas, R. L. 

1977. Preferential Feeding: An Optimization Strategy 

in Sea Urchins. Ecological Monographs, 47:337-371. 

1979. Seaweeds: An Overview—Ecological and Economic Importance. Experienta, 35(4):429-432. 

Vadas, R. L., T. Fenchel, and J. C. Ogden 

In press. Ecological Studies on the Sea Urchin, Lytechinus 

variegatus, and the Algal-Seagrass Community of 

the Mosquito Keys, Nicaragua. Aquatic Botany. 

Warmke, G. L., and L. R. Almodovar 

1963. Some Associations of Marine Mollusks and Algae 

in Puerto Rico. Malacologia, 1(2): 163-177. 

Whittaker, R. H., and P. P. Feeny 

1971. Allelochemics: Chemical Interactions between 

Species. Science, 171:757-770. 

Wratten, S. J., and D. J. Faulkner 

1977. Metabolites of the Red Alga Laurencia subopposita. 

Journal of Organic Chemistry, 43:3343. 
Life History of the Hydromedusa 

Stomotoca pterophylla Haeckel 

and Its Ichthyoparasitic Hydroid 

Ronald J. Larson 

ABSTRACT 

Hydroids of the genus Hydrichthys Fewkes, 1887, 

were found parasitizing about 30% of the leptocephalid fish larvae collected at night from a 

shallow reef flat at Belize, Central America. Little 

or no specificity was shown for either host or site 

of attachment. Medusa e released from these hydroids and reared in the laboratory for a period 

of three weeks were identified as Stomotoca pterophylla Haeckel, 1879. Th e hydroid stage of this 

medusa was hitherto unknown. Because S. pterophylla medusae are epipelagic, the leptocephalids 

probably were parasitized in the open ocean 

rather than near shore. Th e generic name Stomotoca L. Agassiz, 1862, predates Hydrichthys Fewkes, 

1887, and therefore has taxonomic priority. Hydroids of Hydrichthys {= Stomotoca) species are 

highly variable in their morphology. As a result, 

it is doubtful that all of the five previously described species are valid. 

Introduction 

Hydroids are known to occur on a wide variety 

of substrates, including plants and animals. 

Fewkes (1887) described Hydrichthys mirus attached to Seriola zonata (Cuvier), a teleost fish. 

Since the hydranths lacked tentacles, Fewkes 

(1888) suspected that the hydroid might be a true 

parasite. Later, Warren (1916) reported another 

species of Hydrichthys, H. boycei, and substantiated 

Ronald J. Larson, Department of Biology, University of Victoria, 

British Columbia, Canada V8W 2Y2. 

its suspected parasitic habits by noting that its 

polyps fed on the dermis of fish. Th e worldwide 

appearanc e on a variety of host fishes has been 

described for three additional species of this genus. Because of the parasitic specializations and 

highly variable morphology of the hydroids, in 

addition to our lack of knowledge concerning the 

medusae of the genus Hydrichthys Fewkes, 1887, 

(= Stomotoca L. Agassiz, 1862) the systematic position and the validity of Hydrichthys spp. are 

obscure. This pape r is the first description of the 

life cycle of Stomotoca pterophylla Haeckel, 1879. 

Aspects of its unique life history and the taxonomy of "Hydrichthys" (= Stomotoca) spp. hydroids 

are discussed. 

ACKNOWLEDGMENTS.—I wish to thank D. G. 

Smith, Marine Biomedical Institute, University 

of Texas at Galveston, for identifying the leptocephalus larvae, and K. S. Larson, Smithsonian 

Institution, Washington, D.C , for editing and 

typing the manuscript. 

Materials and Methods 

Hydroids and medusae were collected at Carrie 

Bow Cay, located on the barrier reef of Belize, 

Central America, in March and April 1977 and 

1978, and November 1978. Medusa e and leptocephalus larvae with attached hydroids were obtained by suspending a 30 cm diameter, 220 jum 

mesh plankton net, at night, from a pier over the 

reef flat and allowing water currents to carry the 

specimens into the net. Wate r depth here varies 

from 0 to 0.5 m; the bottom consists of coral 

433 
434 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

rubble, living coral, coarse Halimeda spp., sand, 

and Thalassia testudinum Banks ex Konig. About 

0.6 km seaward of Carrie Bow Cay is the shelf 

edge, where water depth reaches more than three 

hundred meters. 

Newly released medusae were obtained by 

placing leptocephali with hydroids in small bowls 

of sea water overnight. These medusae were kept 

in bowls; the sea water was changed at least once 

daily, generally after feeding. For feeding, medusae were transferred, via pipette, to watch 

glasses. When possible, each specimen was individually fed to repletion once or twice daily. 

Initially, medusae were fed boiled egg yolk and 

Artemia salina (Linnaeus) nauplii; after one week, 

small pieces of gonad from the echinoid Lytechinus 

variegatus (Lamarck) and bits of lobster and fish 

muscle were used. The third week, the diet consisted of live medusae, Liriope sp. and Phialidium 

sp., and chaetognaths. 

All material is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. 

Results 

Leptocephali were found only at night on the 

reef flat. During each 30-minute plankton tow, 

10 to 20 or more larvae were collected. The larvae 

had an adult-like head indicating an advanced 

stage (Figure 183^) and when kept in the laboratory they metamorphosed into juveniles after 

three to 14 days. 

Hydroid colonies were found on a significant 

number of leptocephalids. Of 101 larvae examined, 35 had attached hydroid colonies. In all, 10 

species of larvae were collected: Ahlia egmontis 

(Jordan), Myrophis punctatus Liitken, Myrophis platyrhynchus Breder, Albula vulpes Linnaeus, Arisoma 

balearicum (De la Roche), Chilorhinus suensoni Liitken, Moringula edwardsi (Jordan and Bollman), 

Flops saurus Linnaeus, Ophichthus sp., and 

POphichthidae; only the first four species mentioned were parasitized by hydroids. Ahlia egmontis 

and M. punctatus were the most abundant in the 

collections and also accounted for the majority of 

the hydroid colonies. Since most of the other six 

species were represented by only one or two specimens, the degree of parasitism could not be 

determined. By contrast, C. suensoni was represented by eight specimens that were all free from 

infection, indicating that this species may be able 

to avoid the hydroid. 

The presence of hydroids on all areas of the 

body points to a lack of specificity for attachment 

sites. Generally, only a single colony was found 

per fish; however, several had two, and one had 

four on different areas of the fish. 

LIFE HISTORY OF Stomotoca pterophylla HAECKEL.—Description of Polyps: Hydranths protrude 

from naked basal anastomotic hydrorhizal plates, 

1 to 4 mm in diameter, that are attached to the 

dermis of the eel larvae (Figure 183</). Polyps are 

elongate, 0.5 to 4.0 mm in length, without tentacles, and number 10 to 20 or more per colony. 

Medusae buds occur on nearly every colony but 

their number and location depend on the polyp 

size. Twenty five or more occur over most of the 

lateral branches of the large, 2 to 4 mm long, 

hydranths (Figure 183c). Smaller polyps, 0.5 to 

2.0 mm, on the proximal portion of the hydranth 

generally show few sessile buds (Figure 183a,£). 

Medusae buds first appear as globular outgrowths 

from the polyps, later elongating and then developing a pair of triangular tentacle rudiments. In 

life, polyps are reddish, medusa buds are unpigmented except for the yellowish endodermal region of the manubrium and the tentacle rudiments. From these characters the hydroids are 

identified as belonging to the genus Hydrichthys 

Fewkes, 1887. 

Description of Medusae: The bell-shaped umbrella 

of medusae reared from Hydrichthys sp. polyps 

measures 0.5 to 0.7 mm in height and is slightly 

narrower than tall (Figure 184a). Cnidae are 

scattered over the exumbrella. The cruciform 

manubrium occupies about one-half the diameter 

of the subumbrellar cavity. Lips are simple, not 

folded. Two opposite tentacles originate from 

flattened triangular perradial marginal bulbs. 

Two remaining perradial bulbs are wart-like and 

without tentacles. Four smaller wart-like bulbs 

occur interradially. Vascular canals are narrow. 

The gastrodermis of the manubrium and the 
NUMBER 12 435 

lmm 

FIGURE 183.—Stomotoca pterophylla hydroids: a-c, hydranths with developing medusa buds (a,b 

with several buds, c with numerous buds); d, metamorphosizing leptocephalid Myrophis punctatus 

with S. pterophylla hydroid colony attached to left side (hy = hydranths, mb = medusa bud). 

marginal bulbs is reddish-orange in color, other 

structures are unpigmented. 

Newly released Stomotoca pterophylla medusae 

collected directly from the reef flat are morphologically identical to those released in the laboratory. During subsequent growth, the most notable changes are the appearance of an apical 

projection, absolute and relative increase in manubrium length, formation of gonads, and increase 

in the number of marginal warts. At 3 mm bell 

height (7 days), the umbrella develops an apical 

projection (Figure 184£), the manubrium extends 

to the velum, and 12 to 18 marginal bulbs become 

evident. At 5 mm height (9-14 days) the apical 

projection becomes more prominent, the manubrium originates from a slight peduncle, the 

mouth stretches beyond the velum, the lips fold 

slightly, adradial gonads appear in 2 to 5 transverse folds on the proximal portion of the manubrium, and 16 to 30 marginal warts become 

apparent (Figure 184c). Already at this stage the 

two opposite tentacles, peduncle, and folded adradial gonads are diagnostic of the genus Stomotoca 

L. Agassiz, 1862. At 8 mm bell height (14-17 

days) the peduncle enlarges, and adradial gonads 

add folds, even secondary folds on the largest 

specimens (Figure 184</), and the lips also plicate. 

The gastrodermis of the manubrium, tentacle 

bulbs, and marginal warts is brownish. After 21 

days of development, the total observation time, 

only two of the 25 specimens have bell heights of 

10 mm. At this time, the bell shape appears quite 

variable from specimen to specimen owing to 

differential resorption of the mesoglea during pe-
436 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

0.5 mm 

FIGURE 184.—Stomotoca pterophylla medusae: a, newly released, about 0.5 mm bell height; b, 

approximately 7 days old, about 3 mm high; c, approximately 14 days old, 5 mm high, gonads 

appear as folds on manubrium; d, front view of folded gonad of 17-day old, 8 mm high medusa. 

riods when the medusae would not feed. Because 

of the presence of secondary gonad folds, these 

medusae are diagnosed as Stomotoca pterophylla 

Haeckel, 1879. 

Behavior of Polyps and Medusae: Although hydranths are typically inactive, sporadic serpentine 

motion, bending, turning, and mouth opening 

were observed. During feeding, hydranths bend 

over touching the epidermis of the host leptocephalus, and open the mouth wide, trumpet-like, 

over a small area of the fish. Cnidae inside the 

mouth may attach the polyps's circular lip to the 

larvae during feeding. Damage to surrounding 

fish tissues was observed in all cases, as indicated 

by numerous minute pits and surface irregularities surrounding the hydroid colony. Hydranths 

removed from their hosts were incapable of capturing live Artemia salina nauplii, but would eat 

dead nauplii. 

Stomotoca pterophylla medusae actively pulsate 

except when feeding. Since the medusae lack 

sufficient propulsion to maintain themselves off 

the bottom of the observation dish, they may, in 

the open ocean, rely on water turbulence to help 
NUMBER 12 437 

stay adrift. The long filiform tentacles extending 

to 15 or more times the bell height are probably 

used to reduce sinking as well as to capture prey. 

When food is touched to one or both tentacles 

they contract and bend inward; the mouth is 

then projected outwards by contraction of the 

umbrella, the food is grasped by the lips, and 

swimming pulsations cease. In newly released 

medusae, it is sometimes necessary to place the 

food inside the umbrella cavity to produce this 

feeding behavior. Usually if a medusa keeps 

swimming, the food is dropped. Specimens that 

repeatedly reject food show signs of starvation, 

that is, resorption of umbrellar mesoglea. 

Discussion 

Previously, the hydroid stages of Stomotoca spp. 

were unknown. Strong (1925) obtained planulae 

from Stomotoca atra L. Agassiz medusae from Puget 

Sound. These planulae failed to settle even 

though a variety of substrates was offered; Strong 

suggested that the hydroid might be substrate 

specific. Because £. atra and S. pterophylla medusae 

are taxonomically separated on only a few minor 

criteria (see Bigelow, 1909), undoubtedly their 

hydroid generations are similar. Thus the hydroid 

stage of S. atra probably occurs parasitically on 

fishes. Strong (1925) also found in the Puget 

Sound region a hydroid of the family Pandeidae, 

Perigonimus repens (Wright) commonly attached to 

the marine fish Hypsagonus quadricornis Cuvier and 

Valenciennes. Although the medusa stage for P. 

repens is unknown, it is unlikely that this is the 

hydroid of S. atra medusae because it is morphologically very different from the hydroid of S. 

pterophylla. 

Table 33 lists the host fishes on which "//ydnchthys" spp. and Ichthyocodium sarcotretis Jungersen, a closely related species, have been reported. 

Many of these species, for instance, Cyclothone 

signata Garman, Ceratias holboelli Kr0yer, Diaphes 

theta Eigenmann and Eigenmann, Lampanyctus leucopsarus (Eigenmann and Eigenmann), Tarletonbeania crenularis (Jordan and Gilbert), and Benthosema glacialis (Reinhart), are pelagic. Others have 

long-lived larvae, like Acanthurus triostegus sandvicensis Streets, which was found to be planktonic 

for 2 V2 months (Randall, 1961). 

Myrophis punctatus, the most heavily parasitized 

leptocephalid found during this study, has a pelagic larva that lives offshore for as long as 6 to 

14 months before moving inshore and metamorphosizing (M. Leiby, pers. comm., 1979). This 

long pelagic life may be the reason why it is 

parasitized more often than benthic or inshore 

species without pelagic larvae, in light of the 

medusa stages of the hydroids also being pelagic. 

TABLE 33.—Previous reports of the hydroids "Hydrichthys" and Ichthyocodium, and their host fishes 

Hydroid 

Hydrichthys boycei Warren 

Hydrichthys cyclothonis Damas 

Hydrichthys minis Fewkes 

Hydrichthys pacificus Miyashita 

Hydrichthys pietschi Martin 

Hydrichthys sp. 

Ichthyocodium sarcotretis Jungersen 

?Ichthyocodium sarcotretis Jungersen 

Host fish 

Ambassis safgha (Forskal) 

Chaetodon lunula (Lacepede) 

Mugil sp. 

Cyclothone signata Garman 

Seriola zonata (Cuvier) 

Acanthurus triostegus sandvicensis Streets 

Chaetodon sp. 

Ceratias holboelli Kr0yer 

Diaphus theta Eigenmann and Eigenmann 

Lampanyctus leucopsarus (Eigenmann and E 

Tarletonbeama crenularis (Jordan and Gilbei 

Benthosema glacialis (Reinhart) 

Sebastes mentella Travin 

? Sebastes fasciatus Storer 

Sebastes mentella Travin 

genmann) 

•t) 

Reference 

Warren, 1916; Millard, 1975 

Damas, 1934 

Fewkes, 1887 

Randall, 1961 

Miyashita, 1941 

Martin, 1975 

McCormick et al., 1967 

Jungersen, 1911 

Templeman, 1973 

Jones, 1966 
438 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

In the Atlantic, most Stomotoca pterophylla medusae 

have been collected in oceanic regions—the Florida current, Gulf Stream, and in the mid-Atlantic 

(see Kramp, 1959)—rather than in neritic waters. 

At Carrie Bow Cay, numerous juvenile S. pterophylla medusae were collected, but no mature 

specimens were found. Hence it is probable that 

the leptocephali captured at Carrie Bow Cay 

were parasitized during their lengthy larval period in the open ocean. 

In conjunction with showing the genera Hydrichthys and Stomotoca to be synonymous, it is 

essential to determine the taxonomic status of the 

species in Hydrichthys. Unfortunately, hydroids in 

this genus are separated on the basis of highly 

variable morphological characters, for instance, 

height and diameter of colony, extent of branching, and location of medusa buds. Rather than 

being genetic, these criteria probably depend on 

size of host, age of colony, and other ontogenetic 

factors. The genus Ichthyocodium Jungersen is morphologically similar to the genus Hydrichthys, 

which was also described on the basis of characters that are likely to be affected ontogenetically. 

Since the hydroids of the genera Hydrichthys and 

Ichthyocodium are specialized, owing to their parasitic existence, they have few morphological characters, which seem to vary with environmental 

conditions. Consequently, it may be impossible to 

separate members of these genera solely on polyp 

morphology. Host specificity may also be of limited taxonomic value, since this study shows that 

many hosts are parasitized. Although Ichthyocodium sarcotretis seems to parasitize only ichthyoparasitic copepods, McCormick et al. (1967) found 

Hydrichthys sp. attached to both copepods and 

fish. They, too, suggested that host specificity 

may not be taxonomically useful. 

Comparison of medusae from these species may 

prove to be the most reliable means of identification. Where known, the medusae buds of all 

species of Hydrichthys and Ichthyocodium sarcotretis 

are similar (Fewkes, 1888; Jungersen, 1911; Warren, 1916; Damas, 1934; Miyashita, 1941; Jones, 

1966; McCormick et al., 1967; Templeman, 

1973). Juvenile medusae are known for only two 

species of these hydroids, H. mirus and H. boycei. 

Fewkes (1888) reported that the juvenile medusa 

of H. mirus has a cruciform manubrium and two 

tentacles, but that it later develops a circular 

manubrium and four tentacles. This report led 

Mayer (1910) to postulate that this medusa might 

belong to the genus Sarsia. Warren (1916) found 

that the juvenile H. boycei medusa resembles newly 

released H. mirus but, doubting that it could 

develop more tentacles, he separated the two 

species. The fully developed medusa of H. mirus 

is different in many respects from the medusa of 

Stomotoca pterophylla reported herein. It is possible 

that Fewkes accidently introduced Sarsia sp. medusae into his cultures and mistook them for the 

advanced medusae of H. mirus. 

Clearly, much confusion exists about the validity of hydroids referred to the genus Hydrichthys. 

The true identity of those supposed species awaits 

examination of their life histories. Yet, there is 

little doubt that the genus Hydrichthys Fewkes, 

1887, is synonymous with Stomotoca L. Agassiz, 

1862. Accordingly, the new combinations are as 

follows: Hydrichthys boycei Warren, 1916 = Stomotoca boycei (Warren, 1916); Hydrichthys cyclothonis 

Damas, 1934 = Stomotoca cyclothona (Damas, 1934); 

Hydrichthys mirus Fewkes, 1887 = Stomotoca mira 

(Fewkes, 1887); Hydrichthys pacificus Miyashita, 

1941 = Stomotoca pacifica (Miyashita, 1941); Hydrichthys pietschi Martin, 1975 = Stomotoca pietschi 

(Martin, 1975). 

Literature Cited 

Agassiz, L. 

1862. Contributions to the Natural History of the United Slates 

of America. Volume 4, vii -I- 383 pages, 16 plates. 

Boston. 
NUMBER 12 439 

Bigelow, H. B. 

1909. Reports on the Scientific Results of the Expedition 

to the Eastern Tropical Pacific, in Charge of 

Alexander Agassiz, by the U.S. Fish Commission 

Steamer "Albatross," from October, 1904, to 

March, 1905, Lieutenant Commander L. M. Garrett, U.S.N. Commanding, XVI: Medusae. Memoirs of the Museum of Comparative Zoology, Harvard 

College, 37:243 pages. 

Damas, H. 

1934. Hydrichthys cyclothonis (nov. sp.), hydroide parasite 

du poisson, Cyclothone signata (Garman). Bulletin du 

Muse'e Royal d'Histoire Naturelle Belgique, 10(7): 1-10. 

Fewkes, J. W. 

1887. A Hydroid Parasite on a Fish. Nature, 36:604-605. 

1888. On Certain Medusae from New England. Bulletin 

of the Museum of Comparative Zoology, Harvard College, 

13(7):209-240. 

Haeckel, E. 

1879. Das System der Medusen. In Monographic der 

Medusen, I. Medicimsch-Naturwissenschaftliche Gesellschafl, Denkschriften (Jena), 1:1-360, atlas of 40 

plates. 

Jones, D. H. 

1966. A Gymnoblastic Hydroid Occurring on Sphyrion 

lumpi (Kr0yer). Annual Magazine of Natural History, 

13(9): 173-181. 

Jungersen, H.F.E. 

1911. On a New Gymnoblastic Hydroid (Ichthyocodium 

sarcotretis) Epizoic on a New Parasitic Copepod 

(Sarcotretes scopeli) Infesting Scopelus glacialis Rhdt. 

Videnskabelige Meddelelser Dansk Naturhistorisk Forening i Kj0benhavn, 64:1-33. 

Kramp, P. L. 

1959. The Hydromedusae of the Atlantic Ocean and 

Adjacent Waters. Dana-Report, 46:1-283. 

Martin, W. E. 

1975. Hydrichthys pietschi, New Species, (Coelenterata) 

Parasitic on the Fish Ceratias holboelli. Bulletin of the 

Southern California Academy of Sciences, 74(1): 1-5. 

Mayer, A. G. 

1910. The Medusae of the World. In Hydromedusae, I 

and II. Carnegie Institution of Washington, Publication, 

109(1-2): 498 pages. 

McCormick, J. M., R. M. Laurs, and J. E. McCauley. 

1967. A Hydroid Epizoic on Myctophid Fishes. Journal 

of the Fisheries Research Board of Canada, 24(9): 1985— 

1989. 

Millard, N.A.H. 

1975. Monograph on the Hydroida of Southern Africa. 

Annals of the South African Museum, 68:1-513. 

Miyashita, Y. 

1941. On the Occurrence of a New Hydrichthys in the 

Pacific Coast of Japan. Annotationes Zoologicae Japonenses, 20:151-153. 

Randall, J. E. 

1961. A Contribution to the Biology of the Convict 

Surgeonfish of the Hawaiian Islands Acanthurus 

triostegus sandvicensis. Pacific Science, 15(2) :215—272. 

Strong, L. H. 

1925. Development of Certain Puget Sound Hydroids 

and Medusae. Publications of the Puget Sound Biological Station, 3(75):383-393. 

Templeman, W. 

1973. First Records of the Gymnoblastic Hydroid, Ichthyocodium sarcotretis, on the Copepod, Sphyrion lumpi, 

from Redfish of the Northwest Atlantic. Journal of 

the Fisheries Research Board of Canada, 30:1655-1660. 

Warren, E. 

1916. On Hydrichthys boycei, a Hydroid Parasitic on 

Fishes. Annals of Durban Museum, 1:172-187. 
Variation in Growth Forms of the Reef Coral 

Montastrea annularis (Ellis and Solander): 

A Quantitative Evaluation of Growth Response 

to Light Distribution Using Computer Simulation 

Richard R. Graus 

and Ian G Macintyre 

ABSTRACT 

A new computer simulation model is used to 

evaluate the relationship between the various 

growth forms of small isolated colonies oi Montastrea annularis (Ellis and Solander) and submarine 

light distribution. In this model (Model 2) calcification rate as a function of light intensity is 

calculated from detailed measurements of skeletal 

growth rate and skeletal density in colonies collected from various depths off Carrie Bow Cay, 

Belize, and one large colony from Discovery Bay, 

Jamaica. A previous model (Model 1) was based 

on laboratory studies of photosynthesis rates and 

calcification in response to light intensity (Graus 

and Macintyre, 1976). 

Although these two models represent independent approaches to the understanding of coral 

morphogenesis, the simulation outputs are essentially similar and replicate the morphotypic variations of M. annularis observed in increasing water 

depths off Carrie Bow Cay. These two models 

together corroborate the hypothesis that light 

intensity and distribution provide the major ecological control for the skeletal morphogenesis of 

M. annularis. 

Introduction 

Numerous colonies of the Caribbean coral Montastrea annularis (Ellis and Solander) collected beRichard R. Graus and Ian G. Macintyre, Department of Paleobiology, 

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 

tween 1972 and 1979 from various reef settings 

and depths at Carrie Bow Cay, Belize, have 

provided considerable morphological information 

about skeletal growth and its intraspecific variability. Colonies of M. annularis transplanted from 

shallow water to deep water and vice versa helped 

us to identify skeletal characteristics that are light 

responsive. The modification of these characteristics in the transplanted colonies and their variability among in situ colonies suggest that ecophenotypic factors account for differences in 

growth form over the environmental range of this 

species, principally the ecophenotypic response of 

M. annularis to light. New data on light response 

have allowed us to propose a second model for 

skeletal growth that corroborates the hypothesis 

generated from an earlier model that was based 

on experiments of light-mediated photosynthesis 

and calcification of M. annularis (see Graus and 

Macintyre, 1976; Graus, 1977). 

ACKNOWLEDGMENTS.—We wish to thank M. R. 

Carpenter, R. Larson, B. Spracklin, W. L. Mahood, and R. G. Zingmark for their field assistance. P. Dustan generously provided several coral 

specimens from Discovery Bay, Jamaica, and D. 

L. Meyer provided one specimen from Galeta 

Reef, Panama. We especially thank W. T. Boykins for his assistance in the laboratory and G. 

Capone for his help with computer graphics and 

other programming problems. 

441 
442 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Methods 

Between 1972 and 1979 colonies of Montastrea 

annularis were collected from various reef locations 

off Carrie Bow Cay over a depth of 0.5 to 30.0 m. 

Many of these colonies were stained—some annually over a period of five years—with Alizarin 

Red-S to provide an accurate time marker for 

measuring growth rate. Stain lines correlated with 

cyclic density bands in x-radiographs (Macintyre 

and Smith, 1974), which were also used to measure growth. 

The effects of ambient radiance distribution on 

growth form were tested by transplanting colonies 

from shallow to deep locations on the reef and 

vice versa (details given in Table 34). First, however, each colony was stained with Alizarin RedS and nails were hammered into the skeleton 

vertically at the crest and horizontally in the 

direction of North in order to establish the original orientation at the new site. The colony was 

removed with a hammer, cemented to a concrete 

block, transplanted, left for three years, and then 

collected. 

Two-dimensional light fields were measured in 

the solar plane around several in situ and transplanted colonies at depths from 0.5 to 20.0 m by 

means of an underwater radiometer-photometer 

(United Detector Technology 40X Opto-Meter). 

Measurements were recorded on clear days at 30° 

intervals from 0° (up) to 180° (down), east and 

west of vertical, while the sun was near the zenith. 

Although we did not conduct a comprehensive 

survey of light at other azimuths and solar altitudes, we computed an attenuation coefficient 

that enabled us to correlate our Belizean data 

with the more extensive radiance data from Curacao (Roos, 1967). 

Morphological measurements were obtained 

from vertical skeletal slabs (approximately 0.5 cm 

thick) cut through the crest of each colony. The 

slabs were x-rayed and each negative made into 

a positive contact print on which morphological 

measurements were made directly. Consecutive 

dark and light density bands prominent in the xradiographs represent annual skeletal growth in 

most colonies (Figure 185). Care must be taken, 

however, to differentiate between these pairs of 

bands and other bands caused by local and regional noncyclic environmental fluctuations. 

Skeletal growth rate was determined on the basis 

of the length of a corallite within a given band 

(since corallites normally grow perpendicular to 

the growth surface). This distance was measured 

with vernier calipers. 

Skeletal growth rate is known to vary with 

orientation of the corallites, which can be determined by measuring the corallite growth angle. 

This angle is defined as the interior angle between 

a corallite's axis of growth and the vertical direction. We measured this angle with a straight edge 

and a goniometer. 

Skeletal density was measured in rectangular 

slabs (approximately 3X3X 1 cm) cut from (a) 

the crest and flanks of 9 specimens of Montastrea 

annularis from Carrie Bow Cay and (b) 12 posiTABLE 34.—Transplant conditions for colonies of Montastrea annularis; all at Carrie Bow Cay 

reef except colonies 5 and 6 (P. Dustan's 3-year transplant colonies from Jamaica); colony 8 

rotated 65° at same depth 

Colony 

number 

1 

2 

3 

4 

5 

6 

7 

8 

Original 

depth (m) 

1 

1 

1 

1 

1 

45 

45 

1 

Transplant 

depth (m) 

14 

14 

21 

21 

31 

15 

15 

1 

Transplant 

location 

Outer ridge 

Outer ridge 

Inner-reef slope 

Inner-reef slope 

Fore-reef slope 

Jamaica 

Jamaica 

Back reef 

Transplant 

date 

Jun 1975 

Jun 1975 

Apr 1975 

Apr 1975 

Jun 1975 

Aug 1972 

Aug 1972 

Apr 1975 

Collection 

date 

Mar 1978 

Mar 1978 

Mar 1978 

Mar 1978 

Mar 1978 

Aug 1975 

Aug 1975 

Mar 1978 
NUMBER 12 443 

FIGURE 185—Variations of coral growth patterns of Montastrea annularis with respect to water 

depth along the Carrie Bow Cay transect as shown by the five morphotypes recognized in this 

study: a, hemisphere, 1 m; b, tapered hemisphere, 10 m; c, column, 22 m; d, flared column, 25 

m; e, plate, 30 m; colonies are in life orientation, except for the plate (e), which originally 

dipped 30° to the left. 
444 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

tions spanning the entire range of growth angles 

on a colony of M. annularis from a depth of 5 m 

off Jamaica. An analytical balance was used to 

weigh the slabs both in air {Wa) and suspended 

in water {Ww). The weight in water was recorded 

within seconds of immersion in order to minimize 

seepage into the skeleton. The percentage of error 

for this procedure was thereby kept to less than 

1%. The weights in air and in water together with 

the density of water {Pi = 1.000 g/cm3) were used 

to calculate skeletal density {Ps) from Archimedes' 

formulas as follows: 

Ps = Pi 

Wa 

{Wa - Ww) 

Intercorallite spacing was measured on five 

colonies from depths of 0.5 to 30.0 m off Carrie 

Bow Cay and on one colony from a depth of 5 m 

off Jamaica. Photographic studies were based on 

enlarged 8 X 10 in (20 X 25 cm) prints of small 

sections of the surface of Carrie Bow Cay specimens, both at the crest and lower flanks of the 

colony. In the 30 m plate morphotype, however, 

sections at the center and outer edge were photographed. The specimen from Jamaica was photographed at 12 positions over the entire range of 

corallite growth angles. Owing to curvature of 

the surface of colonies, only 10 to 15 corallites at 

the center of each print were oriented perpendicular to the film and therefore only those could be 

measured without a distortion of distance. The 

distance between the centers of each neighboring 

pair of these central corallites was measured with 

vernier callipers. 

Results 

MORPHOLOGICAL VARIABILITY.—The massive 

variety of Montastrea annularis has been classified 

into three morphotypes on the basis of variations 

that occur at different depths over the entire 

range of this species: hemispheric, 0-5 m; columnar, 5-25 m; and platy, >25 m (Macintyre and 

Smith, 1974; Graus and Macintyre, 1976; Graus, 

1977). Our recent measurements of specimens 

from Carrie Bow Cay indicate that these three 

morphotypic stages are actually part of a continuous gradient that changes with depth. This gradient is described below in terms of five colonies 

that reflect distinct morphotypic stages. 

Hemisphere (Figure 185a): The colony from a 

depth of 1 m in the rubble and pavement zone 

off Carrie Bow Cay shows 10 years of growth 

from its fractured base. Its predominant morphological features are radiating corallites and the 

almost concentric, paired, high and low density 

bands representing annual growth. The rate of 

skeletal growth was found to be highest at the 

crest, averaging 0.83 cm/y for the life of the 

colony. The rate is nearly constant for angles of 

corallite growth up to 60°, but gradually diminishes thereafter and reaches 0.24 cm/yr at the 

maximum angle of 141°. This pattern repeats 

annually, gradually developing the form of the 

corallum. This growth pattern also influences the 

directions of corallite growth. At the crest, where 

the growth rate is nearly constant, all corallites 

except for those lying along the vertical axis 

radiate linearly away from each other. Because of 

differential growth rate along the flanks, corallites 

curve away from the vertical until the polyps 

eventually die out at the lower periphery. 

Interior budding occurs in regions of greatest 

corallite divergence (for example, the crest and 

raised humps). Peripheral budding occurs along 

the lower flanks at the free edge of the epitheca, 

and thus gradually extends the flanks to the 

substratum. 

Tapered Hemisphere (Figure 1856): This specimen from a depth of 10 m in the high-relief spur 

and groove zone resembles the hemispheric forr., 

in its high, relatively uniform growth rate (0.93 

cm/y) over the broad region of the crest. However, the maximum corallite growth angle at 110° 

is lower than that of the hemispheric form. The 

tapered form expands upward rapidly and outward gradually; because the maximum angle of 

corallite growth is greater than 90°, the mature 

form becomes a hemisphere that tapers like an 

inverted cone toward a smaller base. 

Column (Figure 185<r; Plate 4: center left): This 

specimen from a depth of 22 m on the inner-reef 

slope has a growth history of 24 years. Maximum 

skeletal growth rate at the crest averages 0.63 
NUMBER 12 445 

cm/y, which is lower than in either the hemisphere or tapered hemisphere stages. The growth 

rate diminishes beginning at a 10° angle of corallite growth and reaches a minimum rate of 0.18 

cm/y at an angle of 95°. The high differential 

growth rate between the crest and flanks produces 

strong curvature and divergence of corallites and 

thus creates room for new buds. Almost all corallites in the colony originate in an area that was 

an earlier crest of the colony. The geometry of 

corallite growth paths and the shape of the entire 

colony are similar to the axial growth pattern of 

branching corals (for example, species of Stylophora, Pocillopora, Porites, and Dendrogyra). Peripheral budding at the lower flanks forms epithecal 

extensions of the lower flanks, or skirts that become separated from the main skeleton. Buds 

initially grow parallel to the surface of the skirt, 

but within a year or two these buds turn around 

to a nearly perpendicular orientation. The skirts 

develop a concave upward form after several 

years because each new generation of buds turns 

a few additional degrees toward the vertical. Because these buds are not locked in on all sides by 

other corallites, they have more freedom in their 

direction of growth and can thereby influence the 

shape and orientation of the skirt. Although the 

colony has become distinctly columnar after 24 

years of growth, its annual bands indicate that 

the earliest stage was hemispherical and that the 

present columnar form developed gradually. 

Flared Column (Figure 185</): This colony from 

25 m indicates a maximum growth rate at the 

crest of 0.64 cm/y, which is similar to that of the 

columnar form, but here the maximum angle of 

corallite growth (60°) is much lower than for the 

columnar form. As in that form, peripheral budding in the flared column leads to the formation 

of skirts. The growth habit of this stage produces 

a broadly conical colony that is peaked at the 

crest and flared at the base. 

Plate (Figure 185<?): This colony from 30 m on 

the fore-reef slope shows 20 years of growth from 

the base. Although the colony's orientation is 

horizontal in this illustration, it was inclined 30° 

to the left during its life on the reef. Its average 

growth rate is 0.35 cm/y but at the maximum 

vertical angle (53°) growth rate is only 0.06 cm/ 

y. Although interior budding is not evident in the 

x-radiograph of this colony, it has been observed 

in raised areas of this plate. Peripheral epithecal 

budding is dominant, however, and new corallites 

form in the same manner as they do on the skirts 

of the column and flared column morphotypes. 

GENERAL TRENDS IN MORPHOLOGY WITH 

DEPTH.—Maximum Growth Rate and Maximum 

Growth Angle: Data (Figures 186, 187) from 38 

colonies of Montastrea annularis on the maximum 

annual growth rate and maximum angle of corallite growth—the primary factors controlling 

form—indicate that the relationships with depth 

are probably continuous and linear. The least 

squares regression line in Figure 186 predicts a 

maximum skeletal growth rate of 0.9739 cm/y at 

0 m depth and a minimum skeletal growth rate 

of 0 cm/y (that is, the cessation of growth) at 51 

m. The regression line in Figure 187 predicts a 

135.2° maximum angle of corallite growth at 0 

m and a minimum angle of 0° at 50.6 m depth. 

The predicted maximum depth limit of about 50 

m for both trends is about 10% greater than the 

45 m maximum depth said to be normal for the 

species (Goreau and Wells, 1967). 

As expected, maximum skeletal growth rate 

and maximum angle of corallite growth correlate 

positively and significantly with each other (r = 

0.689, p < 0.01). Because the trends are covariant, 

morphotypes can be classified in terms of only 

one variable, that is, maximum angle of corallite 

growth: 

Morphotype 

Hemisphere 

Tapered hemisphere 

Column 

Flared column 

Plate 

Range of maximum angles (c 

120-180 

105-120 

75-105 

45-75 

0-45 

Despite the statistically significant linear depth 

trends, dispersion about each of the calculated 

lines is notable. At any given depth, specimens 

vary in both maximum growth rate and maximum angle of corallite growth as well as in 

growth form. These parameters also vary from 

year to year within a single colony (Figure 185). 
446 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

' 

1.2-

1.0-

0.8-

0.6-

0.4-

0 2 -

0 -

• 

• • 

• 

1 

• 

• 

• 

• 

® 

• 

1 

• 

1 1 

• • v« s N ^ 

• 

I 1 1 

^ \ l 

i i r -1 

10 20 25 30 35 40 45 

Water Depth (m) 

5 0 5 5 

FIGURE 186.—Maximum skeletal growth rate at the crest of colonies oi Montastrea annularis as a 

function of water depth, all data from Carrie Bow Cay transect except values from 45 m, 

Dancing Lady Reef, Jamaica (open circles with dots = duplicate values; regression equation is 

Y= -0.019Z + 0.9739; linear correlation coefficient, r = -0.729, P < 0.01). 

160-

140-

t- 120-

100-

Water Depth (m) 

FIGURE 187.—Maximum corallite growth angle (with respect to vertical) as a function of water 

depth for colonies oi Montastrea annularis, all data from Carrie Bow Cay except the 45 m value, 

Jamaica (regression equation is Y = —2.700^ + 135.156; linear correlation coefficient, r = 

-0.872, P< 0.01). 
NUMBER 12 447 

Skeletal Density: The increase in skeletal density with depth is depicted in Figure 188. The 

regression equation predicts a minimum skeletal 

density of 1.5336 g/cm3 at the surface that increases to 2.0856 g/cm3

 at a depth of 30 m. No 

relationship between skeletal density and corallite 

growth angle could be clearly established. In some 

colonies, density increases from the crest to the 

flanks but in others it does not. Our inability to 

demonstrate this effect—which is readily observable in x-radiographs (Figure 185) as an increase 

in the ratio of dark area (high density) to light 

area (low density)—may be due to the comparatively large size of the skeletal slabs being considered. Slabs taken from the lower flanks of a colony 

generally include portions of skeleton from a previous crest. This material probably masks any 

differences in density between crest and flanks. If 

significantly smaller slabs were used, however, 

measurement error due to water leakage into the 

skeleton would be unacceptably magnified. 

Inter corallite Spacing: The increased intercorallite spacing at the crest with increasing depth is 

plotted in Figure 189. The trend in mean spacing 

concurs with Dustan's data (1975), although our 

results are not directly comparable with his because of unit differences. Our values of minimum 

spacing are based on single measurements for 

each colony; when a new interior bud is present, 

these values represent the distance between the 

new bud (<12 septa) and an adjacent mature 

corallite (24 septa). Values of maximum spacing 

are also based on single measurements. An interesting relationship was observed between minimum and maximum intercorallite spacing in certain colonies depending on whether a newly 

2.2 -y 

2.1 

2.0-1 

nf 

E 

o 

r i.9 

Q 

s 1.8 

1.7-

1.5 

10 15 20 25 

Water Depth (m) 

30 35 

— i 

40 

FIGURE 188.—Skeletal density as a function of water depth for colonies oi Montastrea annularis 

(circles = single-value data from Carrie Bow Cay, squares = depth-specific averages from St. 

Croix (Baker and Weber, 1975); regression equation is Y = 0.0184Z + 1.5336; linear correlation 

coefficient, r = 0.790, P < 0.01). 
448 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

S 0.5-

JG 0.4 -

Water Depth (m) 

FIGURE 189.—Intercorallite spacing at the crest oi Montastrea annularis colonies as a function of 

water depth; similar trends of linear equations for minimum (squares), mean (triangles), and 

maximum (dots) spacing suggests that interior budding is a space-controlled phenomenon 

(correlation coefficients: minimum, r = 0.950, P< 0.01; mean, r = 0.916, P< 0.01; maximum, 

r = 0.866, P< 0.05). 

erupted bud was present in the neighborhood of 

corallites. Neighborhood is taken to mean the 

area that encloses all nearest neighbor corallites. 

If a new bud was present, the distance between it 

and one of the mature corallites in this neighborhood would invariably be the minimum intercorallite spacing for the entire region. If a bud had 

not yet erupted, however, one of the distances 

between two mature corallites within the same 

neighborhod would be the maximum spacing for 

the region. This observation suggests that new 

buds originate in the largest intercorallite spaces, 

and that budding occurs only when sufficient 

space is available (that is, when the space exceeds 

a certain maximum distance between corallites). 

According to the calculated trend lines of Figure 

189, this distance criterion is met when the ratio 

of maximum/minimum intercorallite spacing is 

approximately 1.9. This ratio remains nearly constant even though both the maximum and minimum values increase significantly with depth. 

The linear increase in mean intercorallite spacing with depth is mimicked by the increase in 

mean corallite spacing with corallite growth angle 

on Dustan's colony from a depth of 5 m off 

Jamaica. The correlation coefficient is r = 0.888, 

P< 0.01. The least square regression equation, Y 

= 0.00077Z + 0.3136, predicts spacing to be a 

minimum of 0.31 cm at the crest and to increase 

to a maximum of 0.42 at 140°. Thus, intercorallite spacing varies with increasing corallite growth 

angle in the same way that it does with increasing 

depth. 

TRANSPLANTATION EXPERIMENTS.—Deep-water 

colonies of Montastrea annularis transplanted to a 

depth of 1 m died before they could be collected. 
NUMBER 12 449 

Because the time of death could not be ascertained, these colonies were discarded. Replacements for these samples were provided by Dustan 

who lent us two three-year transplants from 

depths of 45 m to 15 m on Dancing Lady Reef, 

Jamaica. 

All transplanted colonies surviving the experiment showed growth modifications of the type 

and degree expected for in situ colonies at the 

depths of transplantation. Figure 190 shows the 

changes in maximum skeletal growth rate among 

the transplanted specimens. Where the depth of 

transplantation exceeds the depth of origin, the 

maximum growth rate decreases, the new rate 

being in proportion to the new depth; where the 

depth of transplantation is less than the depth of 

origin, the maximum growth rate increases in 

similar fashion. The growth rates of transplanted 

colonies moved from 1 to 14 m are approximately 

the same as the growth rates of colonies moved 

from 45 to 15 m. Thus, the depth of origin as well 

as the direction of transplantation probably have 

little influence on the growth rate at the transplant site. Although the growth rates of transplanted colonies are consistently below the predicted trend for in situ colonies, this result may 

not necessarily be significant. Colonies 1, 4, 5, 

and 8 have growth rates below the trend at their 

depth of origin, so it is not surprising to find them 

below it at the depth of transplantation. The 

maximum value of colony 3 could not be measured at the crest owing to nail damage of the 

polyps, which had to regenerate before normal 

growth could ensue (Figure 192a). Also, all colonies may have experienced a lag in growth during 

a period of shock immediately following the transplantation. This period of shock has been observed in other transplant experiments (Dustan, 

1975) and may have contributed to the demise of 

our transplants from deep to shallow water. Colony 8, which was rotated (turned over) 65° and 

cemented at the same depth, grew at about the 

1.2 -, 

1.0-

o 0.8 

CL 

1 0.6 

o 

1 0.4 

w 

CO 

i 0.2 

0.0 

10 15 20 25 

- 1 — 

30 

— i — 

35 40 

— i — 

45 50 

Water Depth (m) 

FIGURE 190.—Systematic changes in maximum skeletal growth rate at the crest for colonies of 

Montastrea annularis reciprocally transplanted for 3 years at different depths; solid lines with 

arrows indicate direction of transplant and change in growth rate over transplant interval; 

colony 8 was cemented in rotated position (65° to original orientation) at the same depth; 

colonies 6, 7 are from 3-year transplant data from Jamaica (courtesy P. Dustan); dashed line 

is the regression equation of the maximum skeletal growth rate vs. depth of in situ colonies 

(Figure 186). 
450 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

same maximum rate at the new crest as it had at 

the old crest. 

Changes in maximum angle of corallite growth 

among the transplanted corals are depicted in 

Figure 191. Where the depth of transplantation 

exceeds the depth of origin, colonies have a reduced angle of growth almost proportional to the 

depth of transplantation. This reduction is caused 

by a gradual die-back of peripheral polyps (Figure 192a). On the other hand, colonies transplanted to shallower depths show an increase in 

maximum corallite angle, which is caused by 

peripheral budding from the epitheca (Figure 

192tV).The colonies may be said to be developing 

flanks, and given enough time they would probably become tapered hemispheres. 

The rotated colony shifted its entire annual 

growth pattern approximately 65° in the direction of the new vertical axis (Figure 193). New 

polyps appearing on the left periphery resembled 

those in colonies transplanted from deep to shallow water. No die-back occurred at the right 

periphery but the growth rate was considerably 

reduced. 

Similar reorientation of growth pattern following rotation is evident in an accidentally overturned colony found by David L. Meyer in Panama (Figure 194). After being overturned, the 

crest shifted to the new vertical position, new 

polyp growth extended the lower left flank, and 

polyps on the right flank died back. Polyps on the 

left side of the new crest accelerated their growth 

whereas polyps on the right side decelerated their 

growth. Polyps between the old and new crest 

developed a sigmoid growth path after their position changed from being on the left of the old 

I60-. 

140-

sr 120-

< loose-

's 60 -

4 0 -

2 0 -

0 i— 

10 

- 1 — 

15 20 25 

Water Depth (m) 

30 35 40 45 

—i 

50 

FIGURE 191.—Systematic changes in maximum corallite growth angle (with respect to vertical) 

for reciprocally transplanted colonies of Montastrea annularis; all data from Carrie Bow Cay 

transect except colonies 6 and 7, which were transplanted off Jamaica; for the rotated colony 

8, 8T is the angle relative to the new crest; dashed line is the same regression equation for in 

situ colonies (Figure 187). 
NUMBER 12 451 

crest to the right of the new crest. 

Transplantation-induced changes also took 

place in skeletal density and corallite spacing. 

Colonies moved to deeper water showed a significant increase in density at all growth angles 

(Figure 192a). Conversely, colonies transplanted 

to shallower water showed a marked decrease in 

skeletal density for all angles (Figure 192b,c) • 

FIGURE 192.—Growth pattern alterations in x-radiographs 

oi Montastrea annularis colonies after 3 years at transplant site: 

a, colony 3 (Figures 190, 191), transplanted from 1 m to 20 

m off Carrie Bow Cay, shows die-back of live growth surface, 

reduction of annual growth rate at all corallite angles, and 

increase in skeletal density; b and c, colonies 6 and 7 (Figures 

190, 191), transplanted from 45 m to 15 m off Jamaica, show 

increase in maximum growth angle, increase in skeletal 

growth rate, decrease in skeletal density, and decrease in 

intercorallite spacing due to proliferation of interior buds 

(arrowhead = position of maximum growth angle before 

transplant; dot = maximum growth angle 3 years after 

transplant). 

1.0-, 

~ 0.8-

o 0.6-

o 0.4-

0.2-

Vertical Angle (°) 

FIGURE 193.—Change in profile of average annual skeletal growth rate with respect to vertical 

angle for rotated colony 8 (water depth 1 m) 3 years before transplant (dot-plotted curve) and 

3 years after transplant (triangle-plotted curve); after transplant, maximum skeletal growth 

rate remains constant, while maximum growth angle increases about 60° on left side but does 

not change on other side; vertical distance between curves is a measure of increase or decrease 

in growth rate at various locations on colony surface (vertical arrows mark positions of original 

and rotated vertical axes). 
452 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

5 w 

CD <D 

2

 o 

FIGURE 194.—Growth history in x-radiograph of overturned 

Montastrea annularis colony collected by David Meyer in 

Panama, colony oriented with living crest up; corallites 

between original crest and new crest develop sigmoid growth 

paths as they switch from left side of original crest to right 

side of new crest (arrow = maximum corallite growth angle 

before overturning; closed circle = maximum growth angle 

several years after overturning; open circle = intermediate 

die-back position of growth surface at short time after overturning). 

Changes in intercorallite spacing are evident in 

material transplanted from deep to shallow water 

(Figure \92b,c), in that intercorallite spaces become filled with new buds. Because interior budding seldom occurs in the plate morphotype, the 

rapid proliferation of new buds appears to have 

been caused by altered environmental conditions 

at the shallower depth. Despite the decrease in 

spacing at the crest, spacing at the flanks—which 

grow at higher angles—remains greater. Changes 

in intercorallite spacing are difficult to observe in 

colonies moved from shallow to deep water (Figure 192a), but in general the corallites continue 

to diverge without further budding as the growth 

surface expands. Budding would occur eventually, but probably at a set of wider corallite 

spacings. 

These transplant experiments show that the 

morphological parameters that vary with depth 

and angle for in situ colonies are the same ones 

that undergo modification when colonies are 

moved between depths and allowed enough time 

to equilibrate with the new environment. The 

remarkable consistency with which transplanted 

colonies come to resemble in situ colonies at the 

same depth strongly suggests that these morphological parameters and the overall growth form 

are ecophenotypically controlled. Because the 

pattern of morphological variation closely 

matches the pattern of variation of submarine 

light distribution, the most likely factor governing 

this ecophenotypic response is light. 

UNDERWATER RADIANCE DISTRIBUTION.—Data 

on underwater radiance used in the computer 

simulation program were derived from open-water measurements made at depths of 5, 10, and 

20 m by Roos (1967) in Piscadera Baai, Curasao. 

For each depth Roos measured two-dimensional 

light fields at several solar altitudes and azimuths. 

He mathematically corrected the azimuthal data 

to the solar plane, and averaged these values in 

order to obtain an integrated light distribution. 

His final data (Roos, 1967, table 11)—converted 

from microamperes to footcandles using his laboratory to field instrument calibration ratio of 

12.5 lx/juA and the standard International System ratio of 10.8 lx/fc — allowed us to compute 

least squares regression lines of the logarithms of 

his specific angular radiances (0°, 30°, 60°, 90°, 

120°) with increasing depth in order to obtain 

five equations that could be used to calculate 

angle-specific radiance for any depth. Solution of 

the equations at five-meter intervals down to 45 

m gave us a set of gridded data from which the 

contour map in Figure 195 was manually generated. Considering the angular limits of the original data, contours at angles greater than 120° are 

based on extrapolations and should be interpreted with caution. However, because this portion of the contour map is seldom used in our 

simulation program, it has not been problematic. 
NUMBER 12 453 

Vertical Angle (°) 

FIGURE 195.—Logarithm (base 10) of radiance in footcandles contoured with respect to water 

depth and angle from the vertical, radiance data derived from Roos (1967, table 11). The 

dotted area is the range of vertical radiance between 45 m, the reported maximum depth of 

occurrence, and 50 m, the predicted maximum depth of occurrence of Montastrea annularis 

(Figures 186, 187). In this depth range, the maximum growth angle is close to 0°. At shallower 

depths the same radiance exists at greater vertical angles, where it defines the maximum 

corallite growth angle. 

LIGHT-RESPONSE MODELS FOR CALCIFICATION.—Model 1: The light-response model for 

calcification in our previous simulation of skeletal 

growth of Monstastrea annularis (Graus and Macintyre, 1976; Graus, 1977) was based on the light 

and carbon dioxide curve reported by Barnes and 

Taylor (1973). This earlier model (Model 1) assumed that the rates of algal symbiont photosynthesis and skeletal calcification were linearly related and therefore would have similar mathematical expressions with respect to light intensity. 

A subsequent experiment by Chalker and Taylor 

(1975) supported this assumption in observing 

that the rate of calcification in Acropora palmata 

increases proportionately with subsaturating light 

intensity up to 10,500 lx. This result, according 

to the authors, "agrees with studies of photosynthetic carbon fixation in hermatypic corals" (p. 

328). Despite this qualitative agreement, however, the quantitative relationship between the 

rate of photosynthesis and the rate of calcification 

still remains uncertain. 

In our model, the Barnes and Taylor (1973) 

curve was fitted with the exponential function 

r = R[l - e -

A(/

-

/c) ] 

using the reduced major axis method (Imbrie, 

1956). This function states basically that skeletal 

growth rate, r, depends on variation in light 

intensity, /. The relationship is regulated by three 

constants: R, the maximum skeletal growth rate 

at saturation light intensity; Ic, the light intensity 

below which no long-term growth occurs; and k, 

a constant that is equal to the slope of the linearly 

transformed equation. In the equation, R is set at 

1.1 cm/y, which is equivalent to the maximum 

skeletal growth rate of any Belizean colony (pre1975 collection). This rate was measured at the 

crest of a hemispherical colony collected from a 

depth of 1 m. The value of Ic was set at 29 

footcandles (1 fc = 10.8 lx = 1.6 ju,W/cm2) which 

corresponds to the vertical radiance at 45 m 

(extrapolated from light data of Roos, 1967). This 

depth is considered to be the normal maximum 

depth of the species (Goreau and Wells, 1967) 

and is close to the maximum depth for Montastrea 
454 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

annularis in Belize. Because of the pattern of underwater light distribution, Ic not only controls 

the maximum depth of the species, but also the 

maximum angle of corallite growth. Use of this 

equation permitted functional expression of the 

Barnes and Taylor (1973) curve; however, other 

fitting equations such as a polynomial, MichaelisMention equation (Wethey and Porter, 1976) 

could have served as well. 

Model 1 assumed that a single light-response 

curve governs calcification for the entire species. 

Recent reports suggest that the zooxanthella, 

Gymnodimum microadriaticum Freudenthal has highlight and low-light ecotypes that inhabit, respectively, shallow and deep populations of reef corals 

(Barnes and Taylor, 1973; Dustan, 1975; Wethey 

and Porter, 1976). If the photosynthetic response 

mechanism of these ecotypes is indeed distinct, 

there may also be different light-mediated calcification curves; however, any calcification differences between shallow and deep-water populations have yet to be demonstrated experimentally. 

Model 1 also assumed that skeletal growth rate 

in cm/y is proportional to calcification rate in g/ 

cm /y. This assumption is correct if the average 

density of the skeleton is constant. Information 

about systematic density variations was not available before 1975, but since that time various 

workers (Baker and Weber, 1975; Dustan, 1975; 

Highsmith, 1979; and this paper) have shown 

that average skeletal density increases proportionately with depth and probably with corallite 

growth angle and therefore is apparently dependent on light intensity. 

Model 2: Although light-mediated photosynthesis and calcification are known to be related 

(Vandermeulen et al., 1972), the biological mechanism and the mathematics of the interaction are 

still conjectural. Because acceptance of Model 1 

depends on the outcome of this continuing discussion, we decided to propose an independent 

empirical model for calcification in relation to 

light (hereafter referred to as Model 2; see Appendix for summary of computer program) based 

on our skeletal measurements of in situ colonies 

of Montastrea annularis from Carrie Bow Cay. In 

restructuring the original model, we added new 

information on the relationships between light 

and skeletal density and intercorallite spacing. 

Calcification rate in Model 2 is calculated by 

the method of Baker and Weber (1975) as follows: 

Calcification rate (g/cm2/y) = skeletal density 

(g/cm3) X skeletal growth rate (cm/y) (1) 

From the regression equations shown in Figures 

188 and 186, skeletal density and skeletal growth 

rate can be expressed as a function of depth: 

Skeletal density = 0.0184 X depth 

-I- 1.5336 (2) 

Skeletal growth rate = —0.0191 X depth 

+ 0.9739 (3) 

A least squares regression of the logarithm (Ln) 

of mean vertical radiance with depth in Curacao 

(Roos, 1967, table 11) gives the following linear 

equation: 

Depth = 

7.4696 - Ln (Radiance) 

0.08881 (4) 

Sequential substitution of the equations (4) —> 

(3), (2) —» (1) yields calcification rate in terms of 

light intensity. This relationship, plotted in Figure 196, shows how average calcification rate at 

the crest of the colony declines with attenuation 

of vertical radiance. 

Although this model is derived from morphological measurements, use of it does not predetermine the growth form of a simulated colony. It 

merely establishes the skeletal growth rate at the 

crest of a colony at any given depth. Growth rates 

for corallites in other orientations, and hence the 

growth form of the colony, depend upon the 

radiance distribution at the prescribed depth. 

Figure 197 shows that the light-response curves 

for Model 1 and Model 2 are similar in overall 

shape but differ in detail. Both the maximum 

growth rate and the minimum light intensity for 

growth are greater for Model 1 than for Model. 

2. 

SIMULATION EXPERIMENTS.—Morphotypic Variation with Water Depth: Model 2 simulations of 

colony growth oi Montastrea annularis from depths 
NUMBER 12 455 

200 400 6 0 0 800 1000 1200 

Light Intensity (fc) 

FIGURE 196.—Calcification rate for Montastrea annularis as a function of light intensity used in 

Model 2 computer simulations; curve derived from liner equations fit to empirical mesurements 

of maximum skeletal growth rate, skeletal density, and light in relation to depth (Figures 186, 

188). 

1200 

Light Intensity (fc) 

FIGURE 197.—Comparison of Model 1 and Model 2 skeletal growth rate curves in relation to 

light intensity. Minimum light intensity of long-term growth for 1 is 29 footcandles; for 2 it is 

19 fc. Maximum growth rate asymptote for 1 is 1.1 cm/y; for 2 it is 0.9739 cm/y. Curves 

intersect where light intensity is 135 fc and skeletal growth rate is 0.4 cm/y. Vertical distance 

between curves is the difference in skeletal growth rate at that light intensity. 
456 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

of 1 m to 40 m appear in Figure 198. These 

simulations depict the annual growth bands, corallite growth paths, and the development of corallum geometry as seen from x-radiographs of 

vertical skeletal slabs (compare with Figure 185). 

Although these are distinct stages, the program 

can produce any number of transition forms, 

which, taken together, approximate a continuous 

distribution. Some input and output values for 

these simulations are given in Table 35. 

Growth rate values at the crest (Table 35, 

column 3) are derived from the linear equation 

in Figure 186. Growth rates away from the crest, 

as well as the maximum corallite growth angles, 

depend entirely on the angle-specific radiance 

values and their conversion to growth rates via 

the parametric equations of Model 2. The maximum growth angles in the simulations (Table 35, 

column 4) are the angles for which the corallite 

growth rate is 0.1 cm/y. These angles correspond 

closely to values predicted by the regression equation (Figure 187) for real colonies growing at the 

same depths. This outcome lends independent 

support for Model 2. 

The sequence of simulations in Figure 198 

compares favorably with the sequence of real 

colonies in Figure 185. The whole set of morphotypes is represented in correct order, and the 

growth details are also similar. As in the real 

colonies, the maximum growth rate for the simulated colonies occurs at the crest, and the rate 

TABLE 35.—Input and output values for Model 2 simulation 

oi Montastrea annularis growth; in each run, PGRATE is 10% 

(see Appendix); radiance distributions based on open-water 

data (see Figure 195) with no reflection added 

Water 

depth 

(m) 

1 

5 

10 

20 

30 

35 

40 

Growth 

interval 

(y) 

15 

15 

15 

25 

10 

15 

20 

Base 

radius 

(cm) 

2.0 

2.0 

2.0 

2.0 

5.0 

8.0 

8.0 

Growth rate 

at crest 

(cm/y) 

0.959 

0.869 

0.795 

0.587 

0.408 

0.304 

0.204 

Maximum 

growth 

angle (°) 

117 

114 

97 

82 

61 

45 

24 

diminishes gradually with increasing angle to a 

minimum of 0.1 cm/y. In the hemispheric colonies (Figure 198a-c) the corallites near the crest 

follow linear paths. Farther from the crest, where 

differential growth of the surface is greater, the 

corallites curve away from the crest. In the column morphotype (Figure 198a?) corallite curvature begins in the crest region. Colonies a-d begin 

from a circular base of the same radius, but only 

in d does the colony become a column. 

Colonies e-g are developing skirts and plates 

because the annual growth rate of the surface has 

fallen below 10%. As in real colonies, the skirts 

and plates develop a concave upward form because each new peripheral bud grows at a lower 

angle than its predecessor. Plate g has become 

completely horizontal. 

Printout records show that the rate of increase 

in surface area diminishes annually. This means 

that once the rate falls below 10%, it will continue 

to force, at an ever increasing rate, the growth of 

the skirt or plate. Even the hemispheric corals 

will, in time, develop skirts and plates. We estimate that this should happen after about 25-30 

years of growth. This conclusion agrees with the 

observation that large hemispherical colonies of 

Montastrea annularis in shallow water frequently 

develop skirts on their lower flanks. 

The simulated colony of Figure 198g evidently 

compares morphologically with the real colony of 

Figure 185^, but their depths of occurrence do not 

agree. This difference can be accounted for if one 

remembers that the real plate had an in situ 

inclination of 30°. Thus, although the real specimen has a platelike form, it is probably just an 

isolated flank of a flared column. 

Model 1 and Model 2 Morphotypes Compared: Results of Model 1 and Model 2 simulations can be 

compared from output in Figure 198 and output 

in Graus and Macintyre (1976, fig. 1). The light 

vs. skeletal growth rate curves in Figure 197 also 

show the expected outcome. Whenever light intensity exceeds 135 fc (the intersection point of 

the two curves), skeletal growth rate is greater for 

Model 1; conversely, whenever it falls below 135 

fc the rate is greater for Model 2. According to 
NUMBER 12 457 

1 METER 

5 METERS 

10 METERS 

20 METERS 

35 METERS 

40 METERS 

FIGURE 198.—Computer simulation plots of the growth oi Montastrea annularis colonies in relation 

to submarine radiance distribution from depths of 1 m to 40 m, generally reproducing the 

depth-related morphotypic gradient shown in Figure 185; simulations utilize the calcificationlight relationship of Model 2 in Figure 196 and open-water radiance data from Roos (1967, 

table 11) contoured in Figure 195. 
458 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Figure 195, vertical radiance of 135 fc (logio 135 

= 2.13) occurs at a depth of 27 m on the reef. 

Above this depth, then, the growth rate at the 

crest is greater and declines faster with corallite 

growth angle for Model 1 colonies than for Model 

2 colonies. Model 1 colonies are thus relatively 

more peaked than Model 2 colonies. Below 27 m 

the reverse is true. Because the minimum threshold light intensity is lower for Model 2 than for 

Model 1, Model 2 colonies always have a greater 

maximum growth angle at any depth than Model 

1 colonies. Therefore the transition depths between morphotypes as well as the maximum 

growing depth are greater for Model 2 colonies 

than for Model 1 colonies. Despite measurable 

differences in the morphotypes produced by the 

two models, the differences are not readily discernible in the graphical outputs. Hemispheres 

from the two models are similar and can be 

distinguished only by careful observation. 

The simulation results of Model 1 and Model 

2 were compared in order to examine the morphological effects that would result if Montastrea 

annularis had shallow- and deep-water populations with separate light-calcification curves. 

Such a situation exists in the light-productivity 

curves of shallow- and deep-water populations, 

respectively, of Pavona praetorta Dana (Wethey and 

Porter, 1976). These curves closely resemble the 

light-calcification curves of Model 1 and Model 

2. If (1) the light-calcification differences of these 

populations of M. annularis are no greater than 

the differences between the light-productivity 

curves of Wethey and Porter (1976) and (2) these 

populations are separated by depth, except for a 

narrow overlap zone between 15 m and 20 m, as 

suggested by Baker and Weber (1975) and Dustan (1979), then the shallow population would 

have only hemisphere and tapered hemisphere 

morphotypes, whereas the deep population would 

have only the flared column and plate morphotypes. In the overlap zone, the column morphotype could predominate for both populations. 

According to the simulation models, these column 

morphotypes of the two populations would be 

distinguished by the range of maximum corallite 

growth angle: 82° to 92° for the shallow population, 98° to 108° for the deep population. However, because the ranges are close and because 

other factors such as slope, bottom reflection, and 

shading by neighboring organisms also influence 

the maximum angle, recognizing the two populations in the overlap zone would be difficult. 

Therefore shallow and deep populations cannot 

be distinguished with certainty on the basis of 

growth form. Furthermore, the simulation models 

suggest that the sequence of morphotypes is a 

single continuous gradient with depth. 

Morphological Effects of the Bottom Reflection: In 

Model 1 simulations, a 5% light reflection was 

added to all radiance values to render the openwater data of Roos (1967, table 11) more realistic 

in terms of benthic lighting conditions. For Model 

2 simulations, this procedure was eliminated routinely because we could not accurately express 

the distribution of reflection to all radiance angles. However, because corals grow on a highly 

reflective substrate, it is important to speculate 

on the possible morphological effects of reflection. 

Figure 199 shows the effect of including different 

reflection values in Model 2. As can be seen, with 

increasing percentage reflections there is a progressive increase in the growth of the lower flanks 

with respect to growth at the crest. Even with 

2.5% reflection, the maximum angle of corallite 

growth extends to 180°. Thus, if a colony were 

somehow suspended above the substratum it 

could grow vertically downward. Other differences not apparent in the diagrams are the decrease in skeletal density and the increase in 

corallite spacing at all corallite growth angles. 

Reflection from the bottom substrate plays a 

significant role in the morphology of the colonies 

but the amount of reflection is difficult to establish. Brakel (1979) found an average of 18% bottom reflection for sandy substrates and 5% for 

living substrates, but these values can vary considerably from one microhabitat to another. 

The morphological effects of reflection are 

probably greater for shallow-water colonies than 

for deep-water colonies. If we consider two colonies, one shallow and one deep exposed to the 

same percentage reflection (for example, 5% of 

penetrating light), the amount of additional re-
NUMBER 12 459 

0% 2.5 % 10% 

FIGURE 199.—Simulations of morphological effects on colony at 1 m depth resulting from the 

addition of increasing amounts of light reflection (from bottom substrate) to open water 

radiance data: a, no addition; b, plus 2.5%; c, plus 10%. Growth increments that include 

reflection are obtained by adding the percent of vertical radiance equally to radiance values at 

all vertical angles. 

fleeted light in shallow water would be greater 

than that in deep water. Therefore, this factor 

and the postulated relationship of corallite 

growth rate to absolute rather than relative radiance suggest the morphological effects of reflection should diminish toward deep water. Thus, 

any additional reflection would make shallowwater colonies more rounded and increase the 

transition depths between the morphotypes but 

it should have relatively little effect on deeper 

water colonies. 

Transplantation of Colonies: Figure 200 shows 

the simulation of transplanted colonies from shallow water to deep water. The transplant conditions (from 1 to 20 m) in Figure 200a, left, are 

comparable to those of the real colony in Figure 

192a, and the outcome is nearly the same (Table 

36). 

As in the real colony, there is reduction of 

maximum growth angle, which involves a dieback of the live surface, the skeletal density increases at all corallite angles, and the intercorallite spacing becomes wider as the corallites diverge without producing interior buds. Figure 

200a, right, shows an extension of the experiment 

to 9 years at the transplant site. The transplanted 

colony has grown a skirt. Although skirt development on a real transplanted colony has not 

been observed, we would expect it to occur in a 

natural experiment of this duration. Figure 2006 

m 20m 

30m 

FIGURE 200.—Simulations based on Model 2 showing 3 years 

(left) and 9 years (right) of growth in shallow to deep water 

transplant experiments: a, transplanted from 1 m to 20 m; 

b, from 1 m to 30 m. As in actual transplant experiments, 

simulated transplanted colonies experience reductions in 

growth rate and maximum corallite growth angle proportional to transplant depth, and increases in skeletal density 

and intercorallite spacing. After 3 years at transplant depth, 

simulated colonies begin to develop skirts (not observed in 

actual colonies.) 
460 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 36.—Comparison between growth data at different 

depths for real and simulated Montastrea annularis transplanted colonies; both transplants from 1 m to 20 m lasting 

for three years 

Specimen 

Real colony 3 

(Figure 192a) 

Simulated colony 

(Figure 200a left) 

Maximum growth 

rate (cm/y) 

Maximum 

growth 

angle (°) 

1 m 20 m 1 m 20 m 

1.000 

0.959 

0.390 

0.587 

125 

117 

78 

82 

shows the same type of simulation experiment for 

a 1 m to 30 m transplanted colony. Comparison 

of the simulation output (Table 35) with those of 

the real colony 5 (Figures 190, 191) again shows 

a close similarity of results. 

Similar simulated transplant experiments from 

deep water to shallow water were not attempted 

because the computer program would have to be 

modified. We have no doubt they could be done 

successfully. In addition, it should be possible to 

simulate the change in growth pattern produced 

by experimental or natural rotation of the colony 

(see Figure 194). For this, the program would 

have to be altered so that it calculates both sides 

of the colony (at present, because only one half of 

the colony is calculated, it must be bilaterally 

symmetrical). 

Conclusions 

Stained colonies oi Montastrea annularis and their 

x-rays have provided a detailed understanding of 

the major geometric parameters that dicate the 

growth and morphotypic variations of this species. In all colonies studied, for example, maximum skeletal growth rate occurs in vertically 

oriented corallites that are usually situated at the 

crest of the colony and the rate of growth diminishes systematically with corallite growth angle. 

The maximum growth rate and the maximum 

vertical angle of corallite growth also diminish 

linearly with depth. These varying combinations 

of parameters produce a continuous spectrum of 

corallum growth forms from the surface to 50 m, 

ranging from hemispheres, through tapered hemispheres, columns, and flared columns, to plates. 

Similarly, skeletal density and intercorallite spacing gradually increase with increasing corallite 

angle and depth of colony growth. 

Transplanted colonies in both Belize and Jamaica undergo nearly complete modification of 

all the described parameters, and after three years 

the growth form closely matches that of in situ 

colonies at the same depths. These results demonstrate that morphotypic variation oi Montastrea 

annularis with depth is predominantly an ecologically rather than a genetically controlled phenomenon. A similar conclusion was reached by 

Foster (1979), who found, using similar transplant 

experiments, a high degree of phenotypic plasticity in the corallite microarchitecture of this species. 

With these new data we were able to construct 

a computer simulation model to evaluate the 

relationship between submarine light distribution 

and the growth forms of Montastrea annularis. Unlike our earlier model (Model 1 in Graus and 

Macintyre, 1976), which was based on experimental studies of photosynthesis and calcification 

in response to light intensity, this empirical model 

(Model 2) was based on calcification in relation 

to light as established by our measurements of 

skeletal characteristics relative to corallite orientation and water depth. 

Simulation experiments with Model 2 show 

that as in the case of Model 1 a broad range of 

morphotypes produced are comparable to the 

corallum growth form sequence observed with 

increasing depth on the Carrie Bow reef transect. 

In addition, simulations of the transplant experiments replicate the observed changes in growth 

forms and show that the modifications result from 

a combination of altered skeletal growth rate, 

maximum corallite growth angle, skeletal density, 

and intercorallite spacing. Simulation experiments on the effects of bottom reflection indicate 

that variation of the lighting conditions in the 

microhabitat can significantly alter corallum 

morphology. 
NUMBER 12 461 

Our light-response model was developed to test 

quantitatively a working hypothesis, the details 

of which are constantly being revised as new data 

are obtained on the skeletal development of tropical reef corals. In producing morphological variations comparable to those in colonies from habitats of known light fields, this model confirms 

that light is the major factor controlling the skeletal morphogenesis of Montastrea annularis. Many 

other factors not specifically considered by this 

model also influence the skeletal development of 

M. annularis, such as the symbiotic growth interaction of this coral and the sponge Mycale laevis 

(Goreau and Hartman, 1966). 

It should also be noted that ambient light 

conditions can vary considerably within habitats 

at the same depth. For example, shallow-water 

corals growing in shaded microhabitats or on 

sloping substrates exhibit skeletal (morphogenesis) characteristics normally associated with colonies in deeper water. Such variations in light 

field and related variations in skeletal characteristics must be taken into account whenever our 

findings are used to interpret paleoenvironments. 

That is to say, dense platy growths of M. annularis 

might indicate either deep water or restricted 

light conditions in a shallow environment. On the 

other hand, porous hemispherical growths would 

clearly indicate a shallow-water habitat. A relatively porous columnar colony might also be used 

to establish intermediate paleodepths in the range 

of 15-25 m; however, crowding of colonial development or bioerosion on the colonial flanks can 

produce pseudo-columns in very shallow water. 

In restricting our field observations to relatively 

small, isolated, smooth-shaped colonies, we have 

not taken into account variations in shape and 

colonial architecture of other Montastrea annularis 

colonies within the same reef habitat. For example, in the shallow back-reef area off Ambergris 

Cay (Figure 201a), one massive development of 

smooth-surfaced "knobby" (terminology in Dustan, 1975) M. annularis has some areas of distinctly 

"lumpy" surfaces. Although these two surface 

types have a similar yellow-brown color, the 

"lumpy" areas have denser skeletons, wider corallite spacings, and greater corallite relief than 

the "knobby" growths. In the same back-reef 

FIGURE 201.—Variations in colonial growth oi Montastrea annularis in patch reefs in the shallow 

back-reef area off Ambergris Cay: a, large yellow-brown development of M. annularis with areas 

of "lumpy" widely spaced corallites between relatively smooth-surface "knobby" growth; b, 

yellow-brown isolated "knobby" M. annularis in the foreground, which contrasts sharply with 

continuous dark brown surfaces of "lumpy" growth in background. 
462 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

area, dark brown, "lumpy" M. annularis can be 

found growing in juxtaposition with yellowbrown "knobby" colonies (Figure 20lo). In the 

latter case, the colors of the various growth forms 

differ, although the skeletal density and corallite surface characteristics are similar. 

These variations in skeletal morphogenesis— 

which have been observed in the deeper fore-reef 

as well as the shallow back-reef areas off Carrie 

Bow Cay—are still unexplained. The lack of evidence that this skeletal development is a response 

to ecological conditions suggests that it could be 

genetically determined. 

Appendix 

SYNOPSIS OF MODEL 2 COMPUTER PROGRAM 

The computer program for simulation of 

growth oi Montastrea annularis is stored at Museum 

of Natural History, Smithsonian Institution, and 

is available by contacting the authors. The program written in FORTRAN IV is divided into 

two parts: a main program, which performs the 

substantive calculations, and a plotting program, 

which generates plotted output on a Calcomp 

(California Computer Products) plotter. The 

main program consists of seven routines, which 

are described below in program order. 

1. READ INPUT VARIABLES 

External communication with the program is 

carried out by entering values of the following six 

variables, which establish the experimental conditions for each simulation run: 

NYEARS numbers of years of growth of simulated colony 

RADNCE sequence of depth-specific radiance values at 5° intervals from 

0° (up) to 180° (down) 

REFL percent of vertical radiance from bottom substrate to be added to all radiance values 

PGRATE minimum allowable annual increase in colony surface area 

RBASE radius of circular base of simulated 

colony 

TPLANT transplant switch that tells the program to grow the colony using two 

different sets of radiance data, one 

at the origin depth and one at the 

transplant depth 

2. CALCULATE ANGLE-SPECIFIC 

ANNUAL GROWTH INCREMENTS 

The 5° RADNCE values are first linearly interpolated to obtain values at 1° intervals, and 

the entire radiance distribution is then smoothed 

using the three point moving average method. 

The light values for all angles are then converted 

into annual growth increments by the sequence 

of parametric equations of Model 2. The maximum growth angle is set at the angle for which 

the annual growth increment equals 0.1 cm/y. 

This corresponds with the observation that if a 

polyp survives a year, it will grow at least at that 

rate. 

3. ESTABLISH COLONY BASE AND 

INITIAL CORALLITE POSITIONS 

The simulated colony starts growing on a circular base of radius, RBASE. Although a circle 

adequately approximates the shape of an early 

growth stage of a colony, the program can be 

easily modified to accept basal coordinates from 

a real specimen. The base is positioned so that its 

center is on the F-axis at or near origin of the 

rectangular coordinate system. The F-axis is also 

the vertical symmetry axis of the colony. The 

negative F-quadrant is not displayed; thus, the 

^Y-axis is the flat bottom substrate. 
NUMBER 12 463 

The terminus of the first corallite is placed at 

the intersection of the circular base with the Faxis, and is assigned a corallite growth angle of 

0°, thereby establishing its orientation perpendicular to the base. The location of the second 

corallite is then placed on the perimeter in the 

positive X,Y quadrant, 1.5 times the minimum 

budding distance (IPDIST) from the first corallite. Its corallite growth angle, also perpendicular 

to the base, is the angle between vertical and a 

radius drawn from the center of the circle to the 

coordinates of the second corallite. Additional 

corallites are positioned in the same manner, 

relative to the previous corallite, until the maximum growth angle is reached. 

4. CALCULATE NEW GROWTH SURFACE 

AND CORALLITE POSITIONS 

The colony grows beyond the base in yearly or 

other regular events up to the number specified 

by NYEARS. Corallite growth angles are first 

converted into annual growth patterns by adding 

the various calculated annual growth increments 

to the terminal positions of the corallites in the 

directions specified by the growth angles. These 

projected coordinates define the next annual 

growth surface of the colony. 

The corallites cannot be extended merely to 

the new surface, however, because most of them 

would violate the rule that they grow perpendicular to that surface. To solve this problem, the 

correct path of a corallite to the new growth 

surface is determined by calculating a perpendicular line from the previous corallite position to a 

line segment extending between the projected 

coordinates of flanking corallites. The growth 

increment is then adjusted to correspond with the 

new growth angle. 

5. CALCULATE COLONY SURFACE AREA 

In computing the live surface area, the colony 

is assumed to be a radially symmetric solid of 

revolution with a vertical axis of symmetry. Horizontal sections cut through the colony connecting the coordinates of the corallites divide the 

colony into a series of trapezoidal solids. The sum 

of the external surface areas of these trapezoidal 

segments is the total living surface area of the 

colony. The area of the new growth surface is 

calculated and then compared with the area of 

the previous growth surface. If the increase exceeds the minimum growth rate specified by 

PGRATE, program control transfers to the interior budding routine; if not, the program control 

transfers to the peripheral budding routine below. 

6. PERIPHERAL BUDDING 

Peripheral budding occurs only when the annual increase in surface area falls below the minimum value specified by PGRATE. In this routine, a new bud is added at a distance 1.5 X 

IPDIST from, and at a growth angle 3° less than, 

that of the terminal corallite. The growth angle 

of the new bud is prevented from becoming less 

than 0°. After addition of the new bud, the total 

surface area is recomputed and compared with 

the area of the previous growth surface. If it still 

falls below the required minimum, new buds are 

added one at a time until the test is passed 

7. INTERIOR BUDDING 

An interior bud is generated whenever the 

distance between adjacent corallites exceeds the 

maximum intercorallite spacing, IPDIST X 2.0. 

The new bud is positioned midway between 

flanking corallites at a growth angle that bifurcates the angle between those corallites. 

8. PLOTTING PROGRAM 

After the entire colony is calculated by the 

main program, yearly growth events are plotted 

one at a time using the stored coordinates of the 

corallites. The growth surface is drawn first by 

connecting the X, Y coordinates in order from the 

vertical to the terminal corallite, with the plotting 

pen held down on the paper. Next, the axes of 

the corallites are drawn by connecting the coordinates of corallites on the new growth surface to 

the coordinates of those same corallites on the 

previous growth surface. Then, the ^-coordinates 

of all the corallites are reversed, and the mirror 

image of the half-colony is plotted in the — X,Y 

quadrant. 
464 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Literature Cited 

Baker, P. A., and J. A. Weber 

1975. Coral Growth Rate: Variation with Depth. Earth 

and Planetary Science Letters, 27:57-61. 

Barnes, D. J., and D. L. Taylor 

1973. In Situ Studies of Calcification and Photosynthetic 

Carbon Fixation in the Coral Montastrea annularis. 

Helgoldnder Wissenschaftliche Meeresuntersuchungen, 

24:284-291. 

Brakel, W. H. 

1979. Small-Scale Spatial Variation in Light Available 

to Coral Reef Benthos: Quantum Irradiance Measurements from a Jamaican Reef. Bulletin of Marine 

Science, 29(3):406-413. 

Chalker, B. E., and D. L. Taylor 

1975. Light-Enhanced Calcification, and the Role of 

Oxidative Phosphorylation in Calcification of the 

Coral Acropora cervicornis. Proceedings, Royal Society of 

London, series B, 190:323-331. 

Dustan, P. 

1975. Genecological Differentiation in the Reef-Building 

Coral Montastrea annularis. 300 pages. Ph.D. dissertation, State University of New York, Stony Brook. 

1979. Distribution of Zooxanthellae and Photosynthetic 

Chloroplast Pigments of the Reef-building Coral 

Montastrea annularis (Ellis and Solander) in Relation 

to Depth on a West Indian Coral Reef. Bulletin of 

Marine Science, 29(1): 79-95. 

Foster, A. B. 

1979. Phenotypic Plasticity in the Reef Corals Montastrea 

annularis (Ellis and Solander) and Siderastrea siderea 

(Ellis and Solander). Journal of Experimental Marine 

Biology and Ecology, 39:25-54. 

Goreau, T. F., and W. D. Hartman 

1966. Sponge: Effect on the Form of Reef Corals. Science, 

151 (3708) :343-344. 

Goreau, T. F., and J. W. Wells 

1967. The Shallow-Water Scleractinia of Jamaica: Revised List of Species and Their Vertical Distribution Range. Bulletin of Marine Science. 17:442-453. 

Graus, R. R. 

1977. Investigation of Coral Growth Adaptations Using 

Computer Modeling. In D. L. Taylor, editor, 

Proceedings of the Third International Symposium 

on Coral Reefs, 2:463-469. Miami, Florida: 

Rosenstiel School of Marine and Atmospheric 

Science. 

Graus, R. R., and I. G. Macintyre 

1976. Light Control of Growth Form in Colonial Reef 

Corals: Computer Simulation. Science, 193(4256): 

895-897. 

Highsmith, R. C. 

1979. Coral Growth Rates and Environmental Control 

of Density Banding. Journal of Experimental Marine 

Biology and Ecology, 37:105-125. 

Imbrie, J. 

1956. Biometrical Methods in the Study of Invertebrate 

Fossils. Bulletin of American Museum of Natural History, 

108:211-252. 

Macintyre, I. G , and S. V. Smith 

1974. X-Radiographic Studies of Skeletal Development 

in Coral Colonies. In A. M. Cameron et al., editors, 

Proceedings of the Second International Coral Reef Symposium, 2:277-287. Brisbane, Australia: Great Barrier Reef Committee. 

Roos, P. J. 

1967. Growth and Occurrence of the Reef Coral Porites 

astreoides Lamarck in Relation to Submarine Radiance Distribution. Ph.D. dissertation, Drukkerij 

Elink'wijk, Utrecht, 72 pages. 

Vandermeulen, J. H., N. Davis, and L. Muscatine 

1972. The Effect of Inhibitors of Photosynthesis on Zooxanthellae in Corals and Other Marine Invertebrates. Marine Biology, 16:185-191. 

Wethey, D. S., and J. W. Porter 

1976. Sun and Shade Differences in Productivity of Reef 

Corals. Nature, 262(5566):281-282. 
Sponge-Zoanthid Associations: Functional Interactions 

Sara M. Lewis 

ABSTRACT 

Epizoic zoanthids are common in coral reef 

sponge communities in the West Indies. This 

study examines several aspects of the functional 

interactions between zoanthids and their host 

sponges. A predator inclusion experiment demonstrates that a major fish predator on the host 

sponge Callyspongia vaginalis does not discriminate 

between zoanthid-bearing and zoanthid-free 

sponges. In situ measurements of volume-specific 

pumping rates of sponges with and without zoanthids indicate that zoanthid colonization slightly 

depresses pumping rates of host sponges, and may 

increase sponge metabolic expenditure in an effort to overcome increased resistance. The frequencies of zoanthid colonization in two sponge 

species are shown to depend on size distributions 

of the host sponge population. This investigation 

suggests that the functional interactions between 

sponges and zoanthids vary with zoanthid species, 

and that the position of sponge-zoanthid associations along a parasitism-mutualism continuum 

depends on the particular species involved. 

Introduction 

Symbiotic associations between sponges and 

epizoic zoanthids are common throughout shallow tropical marine environments and are particularly abundant in coral reef sponge communities. In tropical western Atlantic sponge communities, six zoanthid species occur obligately on 

approximately 12 common host sponge species, 

with varying degrees of host specificity (West, 

1971; Crocker, 1977). Zoanthids become estabSara M. Lewis, Department of Zoology, Duke University, Durham, 

N.C. 27706. 

lished on a sponge following settlement and subsequent metamorphosis of one or a few planktonic 

larvae. Zoanthid polyps then reproduce asexually 

by budding and eventually colonize extensive 

areas of the host sponge surface. Although the 

systematic and morphological aspects of these 

associations have been described (Duerden, 1900; 

Crocker, 1977; West, 1979), the functional basis 

of the sponge-zoanthid relationship remains 

largely unexplored. 

The present study was designed to examine 

directly some of the interspecific interactions that 

may be involved in the maintenance of the 

sponge-zoanthid symbiosis. Two specific hypotheses were tested: (1) avoidance of zoanthidbearing sponges by sponge-eating fish may result 

in differential predation on zoanthid and nonzoanthid sponges, and (2) extensive zoanthid colonization of the inhalant surfaces of host sponges 

may decrease sponge pumping rates by increasing 

resistance to flow. 

The possibility that zoanthid presence may 

serve to reduce fish predation on sponges was first 

examined by West (1976), using a single species 

of host sponge colonized by a brightly pigmented 

zoanthid species. West demonstrated not only 

reduced predation as a result of zoanthid colonization but also demonstrated ichthyotoxic effects 

of zoanthid extracts. Avoidance of zoanthid-bearing sponges by sponge-eating fish was attributed 

to the contrasting coloration of this sponge-zoanthid pair, with bright zoanthid pigmentation signalling toxicity to potential predators. The 

sponge-zoanthid pair studied by West appears to 

represent a mutually beneficial association 

whereby the sponge benefits through reduced 

465 
466 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

predation while zoanthids are provided with a 

competitor-free substrate (West, 1971) in an 

otherwise highly space-competitive environment 

(Jackson and Buss, 1975). Many of the zoanthid 

species in the Caribbean, however, do not possess 

such bright pigmentation (Crocker, 1977; West, 

1979), and the majority of sponge-zoanthid associations are essentially monochromatic. The lack 

of contrasting coloration in these associations may 

reflect a fundamentally different type of association. In the present study a predator inclusion 

experiment was conducted to determine whether 

zoanthid presence results in reduced predation on 

one of these monochromatic sponge-zoanthid 

pairs. 

Colonization of the sponge surface by zoanthids may directly affect sponge metabolism. 

Zoanthid polyps colonize primarily the inhalant 

surfaces of host sponges, covering up to 75% of 

the total sponge surface (West, 1979). Partial 

occlusion of inhalant apertures due to zoanthid 

colonization may thus substantially increase resistance to water flow through the sponge, and 

reduce the volume of water a sponge can filter 

for a given metabolic expenditure. As suspension 

feeding in tropical water is energetically marginal 

(j0rgensen, 1966; Reiswig, 1974), such a reduction of pumping efficiency may be critical. In the 

present study, water transport rates of sponges 

with and without zoanthids were measured in 

situ to test the hypothesis that zoanthid colonization results in lowered sponge pumping rates. 

The frequency of zoanthid colonization within 

a host sponge species has been studied in several 

locations throughout the Caribbean and has been 

shown to vary widely both among and within 

local areas (West, 1976; Crocker, 1977). Although 

attempts have been made to correlate these variations in colonization frequency with other biological and physical variables, the observed variations have not been satisfactorily explained. The 

present study surveyed the frequency of zoanthid 

colonization as it relates to size distribution in 

populations of host sponge species. 

ACKNOWLEDGMENTS.—I thank A. M. Gibbs and 

A. B. Rath for invaluable assistance in the field, 

E. Richardson for providing the photograph, and 

S. A. Wainwright, P. Klopfer, and T. Michel for 

helpful criticism of the manuscript. I thank K. 

Riitzler for his interest and support of this investigation. I have been supported throughout the 

course of this work by an NSF Graduate Fellowship. 

Materials and Methods 

This study was conducted during November 

1979 at Carrie Bow Cay, Belize, Central America. 

Two sponge-zoanthid species pairs were selected 

for study on the basis of their broad habitat and 

depth ranges, monochromatic coloration, and 

morphological characteristics suitable for measurements of oscular pumping rates. The sponges, 

Callyspongia vaginalis (Lamarck) and Niphates digitalis (Lamarck), host the zoanthid species Parazoanthus parasiticus (Duchassaing and Michelotti) 

(Figure 202) and were abundant on lagoon patch 

reefs and along the fore reef off Belize. For habitat 

descriptions and maps, see Riitzler and Macintyre (herein: 9). 

PREDATOR INCLUSION.—Relative weight loss of 

individuals of Callyspongia vaginalis with and without Parazoanthus parasiticus was measured in the 

presence and absence of fish predation in an 

experiment conducted adjacent to a small lagoon 

patch reef approximately 160 m west of Carrie 

Bow Cay (3 m depth). Callyspongia vaginalis with 

P. parasiticus was chosen on the basis of its abundance and the highly similar coloration of sponge 

and zoanthid tissue. Sixteen specimens of C. vaginalis, eight being densely colonized by zoanthids, were collected from a number of lagoon 

patch reefs and randomly assigned to predation 

and control treatments. Pomacanthus arcuatus (L.), 

the grey angelfish, was selected as a major predator because it is locally abundant in areas of 

high sponge density and on the basis of Randall 

and Hartman's (1968) observation that C. vaginalis constituted the single largest portion of the 

stomach content of this primarily sponge-feeding 

fish (21.9% by volume, totalled over 34 specimens). Adult fish were caught in traps, kept in a 
NUMBER 12 467 

FIGURE 202.—Parazoanthusparasiticus on inhalant surface of Niphates digitalis (scale = 1 mm). 

pen for one week, and starved for two days prior 

to the beginning of the experiment. 

Sponges were weighed individually by an underwater weighing apparatus designed to balance 

submerged sponge weight against the buoyancy 

of a measured quantity of air. Because the percentage of original weight lost during the experiment was the variable of primary interest, underwater weights (expressed in ml of air) were not 

converted to above water weights, and changes 

in tissue density within a sponge during the experiment were assumed to be negligible. The 

precision (±1 SD) of the underwater weighing 

apparatus, as determined by replicate weighings 

of brass weights, was ±0.22 ml air, reflecting less 

than 5% of measured values. 

Weighed sponges were mounted on plexiglass 

plates with vertical supports, and placed in two 

adjacent wire mesh (2.54 cm) enclosures, one 

designated as a control and one with four Pomacanthus arcuatus, each enclosure measuring approximately 1.5 X 1.0 X 0.6 m. An estimate of the 

predation pressure exerted on an individual prey 

in this experiment (number of predators X number of days / included volume per individual 

prey) is 0.013 fish-days/liter. This predation pressure corresponds closely to that from West's 

(1976) predator inclusion experiment (0.010 fishdays/liter) and thus discrepancies in the results 

are unlikely to arise from differences in predation 

intensity. 

The frequency of bites by Pomacanthus arcuatus 

on individual sponges was observed daily for one 

hour periods during the experiment. Small 

wrasses could move freely through the wire mesh 

of both enclosures, but were never observed feed-
468 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

ing on experimental sponges. Sponges were reweighed after 12 days, and the percentage of 

original weight lost was calculated for each individual sponge. Accurate final weights could not 

be obtained for a few specimens due to poor 

weather conditions, and one set of replicates was 

excluded from the analysis. 

FLOW RATES.—Specimens of Niphates digitalis 

having a single distal atrial opening were selected 

for the study of water transport rates. Individual 

sponges were marked and measured in the field, 

and oscular area, atrial volume, and specimen 

volume were estimated by detailed measurements 

and geometric approximations. Axial oscular velocity (cm/sec) of each specimen was monitored 

for three minutes with a tripod-mounted heatedbead thermistor flow meter (LaBarbera and Vogel, 1976), enclosed in an underwater housing 

with a chart recorder. Volume-specific pumping 

rates (cm3 water/cm3 sponge/sec) were calculated 

from oscular velocities (cm/sec), oscular areas 

(cm2), and sponge volumes (cm3). Specimens occurring with Parazqanthus parasiticus were chosen 

for their dense colonization, and zoanthid density 

was estimated by counts in 0.4 cm quadrats {N 

= 16-36 depending on sponge size) located over 

the inhalant surface of the sponge. 

COLONIZATION FREQUENCY.—The frequency of 

zoanthid colonization relative to sponge size was 

determined for two host sponge species, Callyspongia vaginalis and Niphates digitalis. The populations 

of each species were censused in two locations: (1) 

a lagoon patch reef 150 m west of Carrie Bow 

Cay (6 m depth) and (2) a 100 m section along 

the outer ridge of the fringing reef (15 m depth). 

Presence/absence of Parazoanthus parasiticus and 

sponge height from basal attachment to oscular 

rim were noted for all N. digitalis individuals. As 

C. vaginalis generally occurs in clusters of tubular 

elements, both the number of tubes and the 

maximum height from basal attachment were 

measured in order to assess sponge volume more 

accurately. Original measurements in inches were 

later converted to centimeters. 

Results 

PREDATOR INCLUSION.—Mean percentage 

weight change for zoanthid and nonzoanthid Callyspongia vaginalis in control and predation enclosures is shown in Table 37. All zoanthid-bearing 

sponges in the control enclosure gained weight 

during the experiment, while all other sponges 

lost weight. Although both predation and zoanthid presence appear to affect weight loss in 

experimental sponges significantly (Table 38), 

examination of individual contrasts using Scheffe 

(1959) multiple comparison tests reveals that this 

differential weight loss in zoanthid and nonzoanthid sponges occurs only in the control treatment. 

The significant P values for both predation and 

zoanthid main effects, as well as the significant 

interaction term, derive from the marked weight 

gain of control zoanthid sponges. In the control 

enclosure, zoanthid sponges gain significantly 

more weight than nonzoanthid sponges {P < 

0.001); no significant difference {P > 0.05) in 

mean weight loss is found between zoanthid and 

nonzoanthid sponges in the predation enclosure. 

Daily observation of the predator enclosure 

could be conducted during only part of the experimental period because of poor weather conditions. These observations nonetheless indicate 

that Pomacanthus arcuatus preys on C. vaginalis regardless of the presence of zoanthids. 

FLOW RATES.—Sponge height, oscular area, 

volume, zoanthid density, and volume-specific 

TABLE 37.—Mean weight change (% of original weight) of 

Callyspongia vaginalis with and without Parazoanthus parasiticus 

enclosed with and without the predator Pomacanthus arcuatus 

(±SD; three replicates per treatment) 

Zoanthids 

Present 

Absent 

Predation Control 

17.7±12.8 

•10.2±7.3 

+33.0±3.5 

-11.9±12.5 

TABLE 38.—Two-way analysis of variance on percentage 

weight loss in Callyspongia vaginalis with and without Parazoanthus parasiticus enclosed with and without Pomacanthus 

arcuatus 

Source 

Predation 

Zoanthid presence 

Interaction 

Error 

df MS 

1 1793.41 12.43 0.0078 

1 1050.94 7.29 0.0271 

1 2056.70 14.26 0.0054 

8 144.25 
NUMBEJU 12 469 

TABLE 39.—Measurements on Niphates digitalis specimens concerning dimensions, zoanthid 

presence and density, and volume-specific pumping rates 

Sponge 

number 

ON18 

ON10 

ON15 

ON20 

ON19 

ON16 

ONI 

ON12 

L6 

ON14 

ON11 

Height 

(cm) 

27.9 

17.8 

18.4 

22.2 

17.1 

14.0 

14.0 

27.9 

29.2 

29.2 

24.1 

Oscular 

area 

(cm2) 

17.3 

97.9 

10.0 

43.3 

50.5 

50.7 

66.4 

45.7 

94.8 

27.1 

40.9 

Volume 

(cm3) 

338 

549 

184 

472 

355 

173 

309 

504 

639 

616 

764 

Zoanthid 

density (X±SD) 

(no./cm2) 

5.9±.9 

4.3±.7 

4.1±1.0 

5.1±.7 

4.7±1.0 

4.3±.8 

0 

0 

0 

0 

0 

Pumping rate 

(cm 3 water/cm sponge/sec) 

0.16 

0.28 

0.16 

0.02 

0.19 

0.23 

0.60 

0.25 

0.16 

0.13 

0.11 

pumping rates are given in Table 39 for 11 

specimens of Niphates digitalis. Mean pumping 

rates for N. digitalis with and without Parazoanthus 

parasiticus (Table 40) indicate pumping rates for 

both groups of sponges within the range reported 

by Reiswig (1974) for three taxonomically distant 

sponge species. The mean pumping rate of nonzoanthid sponges is higher than that of zoanthidbearing sponges, although not significantly so {P 

> 0.05, Mest for means with unequal variances 

on log-transformed variates). Sponges without P. 

parasiticus show greater among-sponge variance in 

pumping rates than do zoanthid-bearing sponges 

(0.025 < P < 0.05, F-test). Zoanthid densities 

averaged over the entire sponge surface show no 

correlation with pumping rates {P > 0.05, Kendall's rank correlation). 

COLONIZATION FREQUENCY.—For a given location, the frequency of zoanthid colonization appears to be related closely to sponge size. Plots of 

sponge height versus number of tubes for each 

specimen of Callyspongia vaginalis from the lagoon 

patch reef and outer ridge populations (Figure 

203) indicate that the largest individuals in both 

populations generally occur with Parazoanthus parasiticus, while smaller individuals have lower colonization frequencies. 

Size (here represented only by sponge height) 

vs. frequency distributions of C. vaginalis in the 

two locations (Figure 204) indicate that the lagoon population consists of sponges of greater 

height with a lower proportion of small individuals than the outer-ridge population. This difference may reflect lower recruitment rates and an 

older, or possibly a faster-growing, population in 

the lagoon. Colonization frequency is significantly related to C. vaginalis height at both locations (lagoon, P < 0.01; outer ridge, P < 0.005, 

G-tests (log likelihood ratio) used for all frequency 

comparisons, with continuity correction for small 

sample size where appropriate). In the lagoon 

colonization frequencies range from 0% for the 0-

16 cm size classes to 47% for the size classes 

greater than 31 cm. In the outer-ridge population, 

colonization frequencies for the same size classes 

are 33% and 93%, respectively. 

Comparisons of colonization frequencies between the outer-ridge and lagoon populations for 

both sponge species grouped by size class (Table 

41) indicate significantly higher colonization frequencies for C. vaginalis on the outer ridge for 

each size class {P < 0.025 for all size classes). The 

overall (summed over all size classes) colonization 

TABLE 40.—Volume-specific pumping rates (cm water/cm 

sponge/sec) for Niphates digitalis with and without Parazoanthus parasiticus 

Zoanthids 

Present 

Absent 

Af 

Pumping rale 

(X±SD) 

0.173±0.080 

0.250±0.181 
470 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Lagoon Patch Reef 

o Without Zoanthids 

• With Zoanthids 

T r 

2 

i—i—i—r 

4 6 8 IO 12 14 16 

No Tubular Elements 

• • 

Outer Ridge 

® o 

8 ° ° 

o ® • 

o Without Zoanthids 

• With Zoanthids 

6 8 IO 12 14 16 

No Tubular Elements 

-"i—i—i—i—i— 

18 20 22 

FIGURE 203.—Callyspongia vaginalis, sponge height versus 

number of tubes for individuals with and without Parazoanthus parasiticus: a, lagoon patch reef; b, outer ridge. (Multiple 

values indicated.) 

28 

24-

20 

16 

Lagoon 

n = 67 

28] 

24 

20 

16-

s 

p77] With Zoanthids 

| | Without Zoanthids 

Outer Ridge 

n=74 

0 3 9 15 21 27 in. 

0 15 30 45 60 cm 

Max. Height 

0 3 9 15 21 27 in. 

0 15 30 45 60 cm 

Max Height 

FIGURE 204.—Callyspongia vaginalis, size-frequency distributions: a, lagoon patch reef population; b, outer ridge population. (Original measurements in inches). 

frequency of 83% in the outer-ridge population is 

significantly higher than that of 25% in the lagoon 

{P < 0.005). No significant differences were found 

between locations for any size class of N digitalis 

(Figure 205), and there was no significant difference in overall colonization frequencies between 

the two locations (32% lagoon, 38% outer ridge). 

Discussion 

PREDATOR INCLUSION.—The predator inclusion 

experiment clearly demonstrates that the sponge 

predator Pomacanthus arcuatus does not distinguish 

between Callyspongia vaginalis with or without Parazoanthus parasiticus. This finding is contrary to 

results of previous work with Holacanthus tricolor 

(Bloch) preying on the sponge Iotrochota birotulata 

(Higgin) with Parazoanthus swiftii (Duchassaing 

and Michelotti) (West, 1976), where the presence 

of zoanthids appeared to reduce predation on 

experimental sponges. Intraperitoneal injection of 

extracts of two zoanthids, P. swiftii and P. puertoncense West, both highly pigmented species, provided evidence for zoanthid toxicity to fish (West, 

1976). The avoidance of zoanthid-bearing 

sponges by predators was further shown by West 

to be visually mediated, although the effects of 

zoanthid coloration were not distinguished from 

those of zoanthid shape. The bright pigmentation 
NUMBER 12 471 

TABLE 41.—Colonization frequencies of Callyspongia vaginalis and Niphates digitalis comparing 

lagoon patch reef and outer ridge populations (P values are given for G-tests of association 

between colonization frequency and location within size class) 

Size class 

(cm) 

0-23.0 

23.1-31.0 

>31.0 

0-16.0 

16.1-23.0 

>23.0 

Location 

Lagoon 

Outer ridge 

Lagoon 

Outer ridge 

Lagoon 

Outer ridge 

Lagoon 

Outer ridge 

Lagoon 

Outer ridge 

Lagoon 

Outer ridge 

Number of individuals 

With 

zoanthids 

2 

20 

6 

10 

8 

13 

2 

9 

2 

7 

6 

11 

Without 

zoanthids Total 

Callyspongia vaginalis 

26 

27 

14 

3 

9 

1 

Niphates 

8 

29 

6 

8 

7 

8 

dig 

28 

47 

20 

13 

17 

14 

talis 

10 

38 

8 

15 

13 

19 

Colonization 

frequency (%) 

7.1 

42.5 

30.0 

76.9 

47.1 

92.9 

20.0 

23.7 

25.0 

46.7 

46.2 

57.9 

P 

<0.005 

<0.025 

<0.025 

>0.05 

>0.05 

>0.05 

fW] With Zoanthids 

| | Without Zoanthids 

Lagoon 

n=3l 

ia 

24 

20 

I6i 

12-

8' 

4" 

Outer F ^id 

n=76 

— 

—- m 0 

0 

3 9 15 21 in. 

15 30 45 cm 

Max Height 

0 

0 

3 9 J5 21 in 

15 30 45 cm 

Max. Height 

FIGURE 205.—Niphates digitalis, size-frequency distributions: 

a, lagoon patch reef population; b, outer ridge population. 

(Original measurements in inches). 

of P. swiftii was interpreted as warning coloration 

signalling zoanthid toxicity to potential predators. Although toxicity of the remaining zoanthid 

species has not been tested, the results of the 

present experiment with P. parasiticus strongly 

suggest that the lack of zoanthid pigmentation 

and contrasting coloration in many sponge-zoanthid pairs may be related to a lack of protection 

of host sponges from fish predators. 

Three of the zoanthid species known from the 

Caribbean, P. swiftii, P. puertoricense, and Epizoanthus cutressi West, are highly pigmented (West, 

1979). The remaining species, P. parasiticus, P 

catenularis (Duchassaing and Michelotti), and Epizoanthus sp. (see Crocker, 1977:32 for description), 

are greyish white and blend nearly completely 

with the color of their host sponges. The present 

study suggests the existence of two classes of 

sponge-zoanthid associations. The first group, 

comprised of brightly pigmented zoanthids and 

their host sponge species, may represent mutualistic associations in which the presence of toxic 

zoanthids serves to reduce predation on host 

sponges. In the second group dull coloration, 

which may be correlated with a lack of toxicity, 

appears to indicate a lack of protection of host 

sponges from predation. If two such classes of 

sponge-zoanthid associations do exist, the evolutionary development of zoanthid toxicity as a 

means of reducing predation on the sponge-zoanthid complex may represent the crossover point 

from an essentially parasitic association, in which 

no benefit to the sponge can be identified, to a 

mutualistic one. 
472 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

The significant weight gain of zoanthid-bearing sponges relative to nonzoanthid sponges in 

the control treatment of the present experiment 

remains unexplained. West (1976) reports a similar result with Iotrochota birotulata; in his control 

treatment, zoanthid-bearing sponges have a reduced weight loss compared to nonzoanthid 

sponges. The significance of the difference, however, was not tested and sufficient data are not 

given for such tests to be performed now. This 

differential weight change within the control 

treatment is important in that it may bias experimental results in the direction of finding lower 

weight loss in zoanthid-bearing sponges in other 

treatments, thus it should have been considered 

in the analysis of West's results. The weight gain 

of zoanthid-bearing sponges in the present experiment may reflect increased growth rates in zoanthid-bearing sponges relative to nonzoanthid 

sponges, although it is also possible that zoanthid 

growth compensates sponge weight loss. The possibility of differential growth rates of zoanthidbearing and nonzoanthid sponges is currently 

under investigation in natural populations. 

Previous studies of sponge population dynamics have shown that predation by fishes is a 

negligible source of mortality (Reiswig, 1973, in 

Discovery Bay, Jamaica). Field observations of 

Pomacanthus arcuatus at Carrie Bow Cay, however, 

clearly indicate predation on Callyspongia vaginalis 

in fore-reef areas (S. M. Lewis, unpublished observations). In addition, sponges with extensive 

tissue loss, presumably resulting from predation, 

are common in sponge populations on the fore 

reef. The depauperate sponge-eating fish fauna 

in Discovery Bay, the result of intensive trapfishing along the northern coast of Jamaica, may 

be responsible for an underestimate of fish predation in sponge populations in other areas of the 

Caribbean. Further work is needed to quantify 

the effects of fish predation on sponge populations. 

FLOW RATES.—Parazoanthus parasiticus extensively colonizes the inhalant surfaces of Niphates 

digitalis, and polyps may completely or partially 

block inhalant apertures. Zoanthid colonization 

may be expected to cause an increased resistance 

to water flow through the sponge, assuming the 

number of ostia remains constant. The lower 

mean volume-specific pumping rate observed in 

zoanthid sponges may be evidence of this overall 

depression of water transport rates due to zoanthid colonization. If, however, choanocyte pumping activity can be increased to compensate for 

increased resistance, pumping rates of zoanthid 

and nonzoanthid sponges may appear similar. 

The major observed difference in pumping rates 

between zoanthid and nonzoanthid sponges appears in the significantly lower variance among 

zoanthid sponges. This reduced variance may 

indicate that these sponges are uniformly increasing their choanocyte activity to its physiological 

limit in order to compensate for increased resistance due to zoanthid colonization. This would be 

expected to increase metabolic expenditure, without increasing filtration rate, and would thus 

have a significant effect on sponge maintenance 

metabolism, as well as on the energy available for 

growth and reproduction. 

COLONIZATION FREQUENCY.—Previous studies 

of sponge-zoanthid associations in the Caribbean 

have included observations of the frequency of 

zoanthid colonization and have established the 

variability of colonization frequencies among host 

sponge populations in different locations. West 

(1976) noted that the colonization frequency of 

Iotrochota birotulata by Parazoanthus swiftii in shallow 

water (<3 m) near La Parguera, Puerto Rico, 

was low, whereas in deeper water colonization 

frequency was close to 100%. He suggests that 

this difference may be related to fish predator 

abundance, although neither colonization frequency nor predation intensity was quantified. 

Crocker (1977) determined zoanthid colonization 

frequencies for 10 host sponge species in four 

transects along the outer bank reef off the west 

coast of Barbados. Colonization frequencies for /. 

birotulata with P. swiftii in these transects were 

low, ranging from 0% {N = 51) to 8% {N = 422), 

while the colonization frequency for Callyspongia 

vaginalis with P. parasiticus ranged from 59% {N 

= 22) to 76% {N= 42). Within sponge species no 
NUMBER 12 473 

correlation was found between colonization frequency and host sponge density in transect areas; 

also, overall colonization frequency (summed 

over all host sponge species) was not related to 

water depth. 

The present study demonstrates that any attempt to explain variations in colonization frequency among host sponge populations must first 

take into account the size distribution of host 

sponges within the populations. For both Callyspongia vaginalis and Niphates digitalis in two distinct physiographic areas, colonization frequency 

has been shown to be directly related to sponge 

size. Within a given location, size may be assumed 

to be some monotonically increasing function of 

sponge age. The correlation of colonization frequency with sponge size appears to indicate an 

increase in the cumulative probability of zoanthid 

larval settlement with sponge age, although the 

possibility of increased survivorship and/or 

growth rate of zoanthid-bearing sponges cannot 

be excluded. This dependence of zoanthid colonization frequency on sponge size may be responsible for some of the differences in colonization 

frequencies reported in previous studies, as overall 

colonization frequency will clearly depend on the 

size distribution of the population under study. 

The significant difference in colonization frequencies observed in the present study between 

Callyspongia vaginalis on the outer ridge and the 

lagoon cannot be attributed to differences in the 

population size structure between locations, as 

comparisons within all size classes give the same 

result. This difference in colonization frequency 

is not likely to be mediated by differences in 

predator abundance between locations, as suggested by West (1976), because of the demonstrated lack of discrimination between zoanthid 

and nonzoanthid C. vaginalis by fish predators. A 

number of mechanisms might be responsible for 

the observed difference. The outer ridge may 

have higher Parazoanthus parasiticus larval availability or higher rates of successful zoanthid settlement and/or metamorphosis, although the lack 

of significant differences in colonization frequencies between locations for Niphates digitalis suggests uniform P. parasiticus availability. An alternative explanation, assuming equal larval availability and settlement in both locations, would be 

reduced growth rates of C. vaginalis on the outer 

ridge relative to the lagoon population. Individuals of a given size on the outer ridge would then 

have been exposed to zoanthid settlement for 

longer periods of time than similarly sized, but 

younger, individuals in the lagoon. Thus either 

differential rates of successful zoanthid settlement 

and metamorphosis on host sponges or differential sponge growth rates in C vaginalis populations 

may explain the observed difference in colonization frequency between the two locations. 

Conclusions 

Although the present study has considered only 

a subset of the possible interactions between 

sponges and epizoic zoanthids, some aspects of 

the nature of the association have been elucidated. Rates of predation on zoanthid and nonzoanthid sponges by a major fish predator appear 

to be identical for Callyspongia vaginalis and Parazoanthus parasiticus, a common species pair representative of sponge-zoanthid associations lacking 

contrasting coloration. Zoanthid colonization appears to slightly depress sponge pumping rates. 

Reduced variance of pumping rates among zoanthid sponges implies an increased metabolic expenditure to maintain pumping rates against increased resistance. Zoanthid colonization frequency has been shown to increase with sponge 

size within a host sponge species. Comparisons of 

colonization frequencies between sponge populations must take into account potential differences 

in the age structures of the populations. 

The obligate nature of the sponge-zoanthid 

association for the six zoanthid species found in 

the tropical western Atlantic indicates a net benefit to symbiotic individuals relative to hypothetical free-living precursors. Perhaps the most obvious advantage to the zoanthids is the availability of competitor-free substrate for colonization. 

There appear to be two classes of sponge-zoanthid 

associations, distinguished on the basis of whether 

zoanthid presence provides a reciprocal net bene-
474 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

fit to the host sponge through reduced predation. 

Increased resistance to water flow due to zoanthid 

colonization may result in a reduction of energy 

available for sponge maintenance, growth, and 

reproduction. In some sponge-zoanthid associations, zoanthid toxicity appears to reduce predation on zoanthid-bearing sponges, and may thus 

result in increased survivorship and reproductive 

output relative to nonzoanthid sponges. The development of toxicity and bright pigmentation 

by some zoanthid species may be a physiological 

investment serving to increase the survival of the 

host sponge, and may represent a crossover from 

parasitic to a mutualistic association. Additional 

study of growth rates, survivorship, reproductive 

output, and competitive abilities of zoanthidbearing and nonzoanthid sponges is needed in 

order to determine precisely the position of particular sponge-zoanthid associations along a parasitism-mutualism continuum. 

Literature Cited 

Crocker, L. A. 

1977. The Ecology of the Zoanthid-Sponge Symbiosis in 

Barbados. Master's thesis, McGill University, 

Montreal. 

Duerden, J. E. 

1900. Jamaican Actiniaria, Part II: Stichodactyline and 

Zoantheae. Scientific Transactions of the Royal Dublin 

Society, 7:133-206. 

Jackson, J.B.C., and L. Buss 

1975. Allelopathy and Spatial Competition among 

Coral Reef Invertebrates. Proceedings of the National 

Academy of Science, 72:5160-5163. 

J0rgensen, C. B. 

1966. Biology of Suspension Feeding. 357 pages. London: 

Permagon Press. 

LaBarbera, M., and S. A. Vogel 

1976. An Inexpensive Thermistor Flowmeter for Aquatic 

Biology. Limnology and Oceanography, 21:750-756. 

Randall, J., and W. Hartman 

1968. Sponge-feeding Fishes of the West Indies. Marine 

Reiswig 

1973. 

1974. 

Scheffe, 

1959. 

West, D 

1971. 

1976. 

1979. 

Biology, 1:216-225. 

H. M. 

Population Dynamics of Three Jamaican Demospongiae. Bulletin of Marine Science, 23:191-226. 

Water Transport, Respiration and Energetics of 

Three Tropical Marine Sponges. Journal of Experimental Marine Biology and Ecology, 14:231-249. 

H. 

The Analysis of Variance. New York: 477 pages. John 

Wiley and Sons. 

A. 

The Symbiotic Zoanthids of Puerto Rico with 

Observations on Their Ecology. Master's thesis, 

University of Puerto Rico, Mayaguez. 

Aposematic Coloration and Mutualism in Spongedwelling Tropical Zoanthids. In G. O. Mackie, 

editor, Coelenterate Ecology and Behavior, pages 443-

452. New York: Plenum Press. 

Symbiotic Zoanthids (Anthozoa: Cnidaria) of 

Puerto Rico. Bulletin of Marine Science, 29:253-271. 
On the Ecology of Isaurus duchassaingi (Andres) 

(Cnidaria: Zoanthidea) from 

South Water Cay, Belize 

Kathleen S. Larson 

and Ronald J. Larson 

ABSTRACT 

On the shallow, wave-exposed reef crest of 

South Water Cay, Belize, more than 700 Isaurus 

duchassaingi polyps were found aggregated in adjacent clumps of up to 100 or more animals each, 

altogether covering nearly 3 m2 of coral rubble 

and algal substrate. Phenotypic variability of polyps is evident from differences in column and 

capitulum coloration and tubercle size and arrangement. Asexual reproduction by budding 

from anastomotic stolons or coenenchyme predominates, although hermaphroditic and female 

polyps are also present. Behavioral and morphological adaptations enable the zoanthids to exist 

within a harsh, fluctuating environment. The 

zoanthids display nocturnal activity by expanding their columns and tentacles only after dusk. 

Light and food stimuli cause the tentacles to 

retract quickly. Rather than microscopic zooplankton, macroscopic prey (that is, larval fish) 

may form the diet. Nutritional requirements of 

Isaurus spp. probably depend on both zooxanthellae and exogenous food transported by waves. 

Introduction 

Zoanthids of the genus Isaurus have circumtropical distributions and live inconspicuously 

among rocky-coral substrates. Although isolated 

Kathleen S. Larson, Department of Invertebrate Zoology, National 

Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560. Ronald J. Larson, Department of Biology, University 

of Victoria, B. C, Canada V8W 2Y2. 

colonies of 5 to 8 polyps occur at depths to 20 m 

(C. Arneson, pers. comm., 1979), most Isaurus 

species are more abundant in intertidal areas. 

This is the first report on the ecology and behavior 

of the Caribbean zoanthid Isaurus duchassaingi 

(Andres, 1883). This study also extends its known 

range to the continental region of Central America, near Carrie Bow and South Water cays, 

Belize. Hitherto, /. duchassaingi was reported from 

the following West Indies locations; St. Thomas 

and Guadeloupe (Duchassaing and Michelotti, 

1864), Bahamas (McMurrich, 1896), Jamaica 

(Duerden, 1898), and Puerto Rico (Duerden, 

1902). 

Related species are known from similar tropical 

habitats in the Indian, Pacific, and Atlantic 

oceans (Andres, 1877; McMurrich, 1889; Haddon 

and Shackleton, 1891; Verrill, 1928; Carlgren, 

1938; Herberts, 1970, 1972a, 1972b; Day, 1969; 

Walsh and Bowers, 1977). Species oi Isaurus have 

variable morphological features: growth habit, 

color, size, number and shape of tubercles on the 

column, and internal anatomy and cytology. The 

only previous ecological study of these zoanthids 

was done by Herberts (1972a), who examined 

habitat requirements for Isaurus spongiosus 

(Andres) from Madagascar; she found that rubble 

substrate, some degree of dampness, and strong 

currents were principal distributional factors. 

The objectives of this study were to examine 

ecological parameters, morphological variation, 

relative abundance and distribution, reproduc475 
476 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

tive biology, functional morphology, zooxanthellae symbionts, general day and night behavior, 

feeding methods, and prey selection on the basis 

of both field and laboratory observations of Isaurus duchassaingi. Numerous colonies partly concealed among coral rubble were found in an algal 

zone at the northeast reef crest on South Water 

Cay, just north of Carrie Bow Cay. A few colonies 

were also seen on the fore-reef terrace east of 

Carrie Bow Cay. A nocturnal species, /. duchassaingi has been illustrated only in retracted, diurnal postures (Duchassaing and Michelotti, 1860, 

1864; McMurrich, 1896; Duerden, 1898, 1902). 

This paper illustrates the expanded animal at 

night with tentacles exposed. 

ACKNOWLEDGMENTS.—We sincerely thank J. 

Marquardt and his library staff, Smithsonian 

Institution, for making available rare and obscure 

publications; C. Arneson, University of Puerto 

Rico, for valuable comments on the manuscript; 

D. Kapraun, University of North Carolina, for 

algal identifications; I. Jewett, for the figures, 

especially the refined polyp drawings; and L. 

Cullen, Tumor Registry, Smithsonian Institution, 

for meticulous preparation of slides. 

Materials and Methods 

Isaurus duchassaingi occurs in small numbers on 

the reef terrace of Carrie Bow Cay, but is most 

abundant on the northeast reef crest of South 

Water Cay, a 28 km2 island about 1.5 km north 

of Carrie Bow Cay, Belize (Figure 206a). The 

distribution of these colonies was mapped in 

March 1978 with the aid of metal poles as 

markers, compass, and measuring tape. Distances 

between colonies, estimated numbers of animals 

and their color, and habitat were described and 

photographically recorded. Conspicuous associated organisms, if visible without disturbing the 

community, were also recorded. 

Two single animals and more than 30 adult 

polyps from one colony oi Isaurus duchassaingi were 

collected with their substrate attached and were 

kept for over three weeks in shaded aquaria. The 

animals were covered by 15 cm of nonrunning 

sea water, which was changed daily. Morphological features of /. duchassaingi were noted during 

the day. Laboratory observations at night were 

made by direct light and by subdued indirect 

light from a flashlight; specimens were photographed with a prefocused camera and electronic 

flash. Feeding experiments were conducted in 

outside aquaria at night after the animals had 

been expanded for two hours. Live zooplankton, 

lab-cultured brine shrimp, or 1 mm3 pieces of 

fresh fish were fed to the animals by dropping 

them from forceps or pipette onto the tentacles. 

Live crab zoeae and 12 mm long fish were also 

kept overnight with the zoanthids. 

Preserved polyps were dissected and examined 

under low magnification. Transverse sections 

through the column below the actinopharynx, 

including gonads and mesenterial filaments, were 

prepared for microscopic examination. Histological sections (6 jum) of paraffin-embedded tissue 

were stained with Hematoxylin-Eosin. Specimens 

are deposited in the National Museum of Natural 

History, Smithsonian Institution, Washington, 

D.C. 

Results 

STUDY SITE.—South Water Cay is a sparsely 

populated island, about 400 m long by 80 m 

wide, located at 16°49'N, 88°05'W, east-southeast of Belize City, Central America (Figure 

206a). Leeward of the island is a seagrass-covered 

lagoon, 0 to 8 m deep. Windward there is a 

shallow (0 to 0.5 m deep) muddy sand and mixed 

seagrass reef flat extending 25 to 100 m to the 

reef crest. At the northeastern end of the island, 

the storm-rubble reef crest, about 50 m wide, 

meets the shore (Figure 207a). 

Isaurus duchassaingi is found in greatest numbers 

at the northeastern end of South Water Cay. 

Here the entire reef crest is strewn with large 

dead coral boulders 0.5 m or more in height 

(Figure 2066, Zones 1-4). The substrate changes 

from near-shore mud and sand (Zone 1) to a 

compact, diffusely intermeshed cobbled area toward the middle of the crest (Zone 2), to rubble 
NUMBER 12 477 

• I —IO #11-5 0 #51-10 0 ^101-20 0 

Number of polyps per colony 

FIGURE 206.—Maps of study area: a, South Water Cay and Carrie Bow Cay, Belize, showing 

study site; b, study site NE of South Water Cay showing zonation of reef crest and location and 

abundance oi Isaurus duchassaingi colonies (A-J). Zones: (1) Zoanthus sociatus; (2) algal belt with 

Isaurus duchassaingi; (3) Porites astreoides; (4) Diadema antillarum and Palythoa sp. 

and beach rock seaward (Zones 3 and 4). About 

10 to 15 m from shore (Zone 2), various algae 

growing in a yellow-green turf over the rubble 

form an approximate north-south extended algal 

belt, about 33 m long and 4 m wide. The belt is 

composed mainly of the Chlorophyta Cladophoropsis membranacea (C. Agardh), with associated Chlorophyta: Halimeda sp. and Dictyosphaeria sp.; Rhodophyta: Laurencia sp., Hypnea sp., Lophosiphonia 

sp., Gelidiella sp. and Jania sp.; and Phaeophyta: 

Padina sp. 

Average 20 cm tides and predominant northeast trade winds markedly affect the crest biota. 

At extreme low tides the area is exposed for more 

than six hours; at mean high tides near-shore 

substrate is covered by 10 cm or more of water, 

and the low tide wave line is covered by 30 cm. 

Waves paralleling the shoreline break harshly 

over the crest on incoming tides. 

Throughout the region opposite the indented 

shore, invertebrates are distributed in distinct 

areas (Figure 206A, Zones 1-4). Conspicuous sessile animals observed near shore (Zone 1) are 

scattered colonies of the green zoanthid Zoanthus 

sociatus (Ellis) associated with sabellid polychaetes, and the orange or brown burrowing anemone Actinoporus elegans Duchassaing. In the algal 

belt (Zone 2), colonies oi Isaurus duchassaingi are 

clumped in patches or scattered singly among 

coral debris that is channeled by sipunculids and 
478 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

' 

FIGURE 207.—Views of Zone 2: a, rubbled reef crest from shore at South Water Cay, algal belt 

lies in front of wave line; b, close-up view of a small Isaurus duchassaingi colony on coral within 

the algal belt; scale at bottom in mm. 
NUMBER 12 479 

polychaetes. Other colonies are found at the bases 

of coral boulders that are sometimes capped by 

the ribbed barnacle Tetraclita sp. Within 5 m 

seaward of the algal zone, heads of the coral 

Porites astreoides Lamarck are interspersed among 

the rubble (Zone 3). Eastward (in Zone 4), there 

are numerous Diadema antillarum (Philippi) (Echinoidea), clumps oi Palythoa sp. (Zoanthidea) and 

scattered Stoichactis helianthus (Ellis) (Actiniaria). 

Beyond Zone 4, coral growth flourishes and is 

dominated by the coral Acropora palmata (Lamarck). (For coral distribution see Riitzler and 

Macintyre, herein; Cairns, herein). 

FIELD OBSERVATIONS.—At the study site, 10 

colonies or patches oi Isaurus duchassaingi comprising well over 700 polyps covering approximately 

3 m2 were found in the algal belt. Estimated 

number of animals per colony varied from two to 

200; it was impossible to count all the polyps 

without digging colonies from the substrate. A 

clump of 100 or more animals covered approximately 400 cm2

. The colonies were densest at the 

center of the belt, where more than 550 polyps 

were within 7 m of each other (Figure 2066, Zone 

2, E-G). The four northernmost (Figure 2066, 

Zone 2, A-D) and three southernmost colonies 

(Figure 2066, Zone 2, H-J) had less than 200 

polyps combined and less than 10 animals per 

patch at sites farthest from the dense central 

population. 

In the algal belt, Isaurus duchassaingi colonies 

are most easily located when they are emersed at 

low tide. Isolated forms, some solitary, but most 

of them colonial (aggregate, adjacent clumps of 

3 to 8 polyps growing from a common stolon or 

coenenchyme), are abundant on a slightly raised 

area of the reef crest. The only visible part of the 

animal is the gibbose, distal portion of the tuberous column—from 1.5 to 2.5 cm in length and 6 

to 8 mm in diameter—which blends well with the 

green and brown algal growths, encrusting coralline algae, and assorted worm tubes on dead, 

pitted coral pieces (Figure 2076). The lower columns of the animals are hidden, wedged between, 

or half buried beneath, the coral. 

Colonies are united below the substrate by 

holdfasts of anastomotic stolons and coenenchyme bases of adult polyps adhering to one or 

more pieces of coral. Embedded around the coenenchyme and stolon tracts are calcareous sand 

grains, Halimeda chips, sea urchin spines, and 

Homotrema rubra (Lamarck) (Foraminifera) tests. 

Because the lower side of the coral substrate is 

partially covered with bryozoans, serpulid tubes, 

and H. rubra nodules, the zoanthid bases partially 

overgrow these organisms. 

The color of the exposed part of the polyp 

columns varies from grey-green to brown to yellow. In the algal belt, where the colonies are 

densest (Figure 2066, Zone 2, E-G), all animals 

are olive-grey, mottled with metallic turquoise. 

Animals in smaller colonies to the north have 

group colors varying from green-tan to browngold with or without a light turquoise hue (Figure 

2066, Zone 2, A-D). The southernmost colonies in 

the belt are either all dark brown with yellow-tan 

knobs—a few polyps are speckled with turquoise 

(Figure 2066, Zone 2, I)—or are all tan-yellow 

(Figure 2066, Zone 2, H, and Zone 1, J). Only 

the basal 1.5 to 2.0 cm, unexposed lower column 

is partly beige or whitened like the stolon and the 

younger polyp buds below. 

LABORATORY OBSERVATIONS.—At dusk, in an 

aquarium, with the animal straightened and capitulum in view, the external column morphology 

of Isaurus duchassaingi is easily discernible (Figure 

208). The tubercled columns of the animals may 

or may not be transversely wrinkled. In some 

colonies, a thin cuticular sheath sloughs from the 

unincrusted column and minute algal tufts cling 

to the capitular and basal regions. The capitular 

disc has numerous radiating ridges and furrows 

that are the same color as the column, or has 

about 15 or fewer white streaks (Figure 2086). 

The base of each polyp lacks tubercles and is 

firmly adhered to one or more pieces of coral. 

The rest of the polyp, except for the unexposed, 

proximal portion, has variously shaped tubercles, 

which may or may not be contiguous, arranged 

in longitudinal or transverse ridges around the 

column. The most distal tubercles on the column 

(crown tubercles) are arranged in a partial or 
480 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 208.—Isaurus duchassaingi live habit: a, contracted, swelled capitulum; b, contracted 

during the day, with closed capitulum; c, expanded at night, with tentacles exposed; d, extended 

capitulum. (bt = body tubercles, ct = crown tubercles, ec = extended capitulum, sc = swelled 

capitulum.) 
NUMBER 12 481 

complete peripheral circle adjoining the capitular 

disc. The distribution and size of the tubercles on 

both the column and crown vary from polyp to 

polyp. Generally, each polyp has a modified radial symmetry around the oral-aboral axis of the 

column that gives the animal a bilaterally symmetrical appearance. The convex portion of the 

polyp usually contains most of the larger tubercles, whereas the semi-transparent concave part 

is smooth, or has scattered or linear series of 

smaller tubercles. 

Column tubercles on the convex side of a polyp 

may be distributed randomly, or in rows, or both, 

as follows: the tubercles may be arranged in 3 to 

4 main rows of 7 to 12 or more prominent tubercles per row, with variously sized scattered tubercles in between; or there may be merely about 15 

prominent tubercles scattered sparsely on the column. A third pattern of tubercle arrangement 

occurs alone in some polyps or may be incorporated with either of the other tubercle distribution 

patterns on other polyps. In this third pattern 

there may be more than 8 tiny tuberculate or 

larger knoblike rows, each with about 10 to 20 

tubercles per row, resembling kernels of corn on 

a cob. 

Crown tubercles, 3 to 15 or more in number, 

may or may not be prominent. Similar-sized tubercles may extend completely around the capitular disc or only partially around the disc on the 

convex part. Or the tubercles may extend all 

around the disc, but decrease in size on the 

concave portion. Crown and column tubercles 

vary in size from 0.5 to 2.0 mm projections, and 

in shape from tiny verrucae to more alated protuberances. 

At night, with polyps fully expanded and capitula unfolded to reveal the tentacles, the column wall extends 3 mm or more above the crown 

tubercles and increases the oral diameter of the 

peripheral capitular edge to more than 1 cm 

(Figures 208d, 2096). This edge supports a double 

circle of alternating simple, short, acuminate, 

white marginal tentacles that expand to more 

than 5 mm in length in a flabellate array. The 

exocoelic tentacles drape slightly downward over 

the edge of the capitulum or they hang vertically 

downward near the column wall, whereas the 

endocoelic tentacles extend obliquely outward, 

about 90° from the exocoelic ones or droop next 

to the exocoelic tentacles (Figures 208c,a', 2096). 

All tentacles are about the same length. From 

photographs and observations of an expanded 

polyp, a diagrammatic sagittal view has been 

sketched to show the outstretched tentacles above 

an elongate preoral chamber, which probably 

serves as an enlarged basin to contain prey before 

it enters the actinopharynx (Figure 2096). 

In the laboratory, Isaurus duchassaingi expelled 

zooxanthellae in mucus-covered packets, about 1 

mm in diameter, which appear as brown rings 

surrounding a bright green center; the mucus was 

teaming with ciliates. Photographs show the endoderm of the tentacles with numerous brownish 

green zooxanthellae. 

The behavior of colonies in aquaria during the 

day is similar to that of colonies observed in the 

field at low tide. During the day, polyps are 

arched toward the substrate and the closed capitulum is oriented downward or perpendicular to 

the ground (Figures 2076, 2086). Polyp orientation is random with regard to waves. By dusk 

(1830 h) polyps expand their hook-like columns 

somewhat and inflate the capitulum (Figure 

208a), but this movement is too gradual to be 

seen. By 2000 h, many of the polyps are vertically 

expanded 4 to 5 cm or more above the substrate, 

having increased their column lengths by about 

1 cm; their columns may be straight or slightly 

curved, and may be sustained in an erect or 

angled posture; their capitula are open and tentacles exposed (Figure 208c). 

At night during the first two weeks in aquaria, 

the polyps reacted to direct light by retracting all 

tentacles and closing the preoral cavity within 

two seconds. No noticeable contraction of the 

column occurred. The tentacles reexpanded 

within three hours after the direct light was extinguished. In the third week, when subjected to 

direct light, the zoanthids did not close until two 

to three minutes later, for unknown reasons. 

During feeding experiments observed with 
482 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 209.—Isaurus duchassaingi, median sagittal sections through anterior portion of polyp: a, 

preserved, retracted; b, live posture, expanded (diagrammatic), (a = actinopharynx, b = body 

wall, ca = capitulum, ct = crown tubercles, m = mouth, p = preoral chamber, me = mesentery, 

mf = mesenterial filaments, sm = sphincter muscle, t = tentacle.) 

minimal light, Isaurus duchassaingi closed its 

preoral cavity as soon as food touched the tentacles. A piece of fresh fish less than 1 mm3

 dropped 

onto the tentacles caused the tentacles and capitulum to infold completely with the food within 

three seconds. When live microzooplankton was 

gently pipetted onto the tentacles or preoral cavity, some of it was caught. Live isopod crustaceans 

dropped into the cavity usually escaped, except 

for some of the less active ones, which became 

enclosed in the chamber when the capitulum 

folded. Brine shrimp, Artemia salina (Linnaeus), 

were quickly caught on the tentacles, which then 

retracted into the chamber. Whether these prey 

were digested was not observed. When a 10 mm 

long cardinal fish touched the tentacles of a polyp 

that had previously been starved for a week, the 

fish adhered and was immediately transferred 

into the chamber, which closed as the tentacles 

simultaneously folded in. The following morning, 

the zoanthid had ejested a bolus, indicating that 

the fish had been digested, the process taking less 

than 12 hours. 

Because it took several hours for Isaurus duchassaingi to reexpand after a disturbance by feeding 

or photographing, only one feeding experiment 

could be done each night. Potential food organisms, however, were allowed to swim freely overnight in the same aquarium. For several days, 

three trigger fish about 12 mm in length swam 

near the zoanthids but were not eaten. Likewise, 

crab zoeae left in aquaria overnight with these 
NUMBER 12 483 

zoanthids were not captured, even though, owing 

to their erratic movements, they probably swam 

into the tentacles. A small gastropod shell and 

some crustacean appendages were found inside 

one of the dissected polyps, at its base. Presumably these animals were eaten, but the relatively 

heavy shells probably could not be expelled by 

the weak musculature of the zoanthid's column. 

STUDY OF PRESERVED SPECIMENS.—Asexual reproduction seems to be predominant in Isaurus 

duchassaingi colonies because most adult individuals had pale tan gemmae sprouting from their 

stolons and growing obliquely from their bases. 

Also present among the colonies were solitary, 

smooth polyps, about 4 mm or more in length, 

which probably formed as buds and then detached from the parent. The mesenteries of many 

of the colonial polyps from the algal belt contained numerous large ova, about 200 jum in 

diameter, and oval spermaries, about 50 jum by 

100 /xm (Figure 210). Some polyps were sterile, 

and others had only developing ova. 

All preserved polyps are in retracted postures 

even though they had been narcotized for several 

days. Polyps are yellowish white, and adult forms 

average 4 to 5 cm in length and 7 to 8 mm in 

width. A sagittal cut through a preserved retracted polyp reveals a preoral chamber below 

the closed capitulum, wherein lay the infolded 

tentacles, about 1 to 3 mm in length, usually 

varying in number from 38 to 42 (Figure 209a). 

The thick mesodermal sphincter muscle above 

the tentacles surrounds the top of the contracted 

cup-like, preoral chamber. At the base of the 

chamber is a slightly elevated, slit-like mouth 

opening, 2 to 3 mm in length, which leads to a 

longitudinally plicated actinopharynx, about 6 

mm in length. The body wall is about 0.5 mm or 

more in thickness, and of greater thickness at the 

level of the tubercles. Most gastric filaments are 

brownish in color. 

Discussion 

Numerous colonies of Isaurus duchassaingi live 

on the reef crest northeast of South Water Cay, 

Belize, and survive under austere environmental 

conditions. Endean (1976) described the influence 

of such factors on the biota on coral reefs: mechanical damage from force of water movement, 

and movement of coral sand and rubble by 

waves; salinity changes and exposure to fresh 

water from rain; temperature changes; emersion 

of animals at low tide; changes in light intensity 

and degree of turbidity; predation by animals; 

and competitive interaction for space. Nonetheless, /. duchassaingi has successfully adapted to the 

intertidal, shallow, reef-crest zone of South Water 

Cay, where temperatures, salinity, sunlight, wind 

and wave action are extreme and variable. 

Isaurus duchassaingi also lives in deeper water. A 

few colonies were seen at 10 m depth on the reef 

terrace, east of Carrie Bow Cay. C. Arneson (pers. 

comm., 1979) has seen them attached to dead 

coral at depths of 15 to 20 m between coral ridges 

in sandy areas off Mona Island, west of Puerto 

Rico; off Puerto Rico; and off St. Croix, Virgin 

Islands. The polyps, however, were not as numerous at any of these locations (only isolated groups 

of five to eight polyps) as those found on the reef 

crest of South Water Cay. 

Previous observers have noted that all Isaurus 

species prefer a coral rubble or rocky substrate 

(McMurrich, 1889, 1896; Haddon and Shackleton, 1891; Duerden, 1898, 1902; Carlgren, 1938; 

Day, 1969; Walsh and Bowers, 1977). Herberts 

(1970, 1972a, 1972b) found that /. spongiosus from 

Madagascar occurred only in areas where variously sized coral rubble was present. Mona Island specimens have been seen nestled in rocks in 

1 m of water with strong currents (R. Larson, 

unpublished data, 1975). Isaurus sp. has also been 

seen adhered to pieces of rock in the intertidal 

zone of northwest Tobago (R. Larson and K. 

Larson, unpublished data, 1977). The bases of 

Belize specimens were likewise attached to coral; 

the cementation of stolon and coenenchyme to 

parts of adjacent coral rubble enhanced stability 

of the animals and of substrates, and helped to 

retain additional sediments. Whether they adhere 

to shells, rocks, or pieces of coral, all known Isaurus 

species do attach to a hard substrate. 

Degree of emersion is apparently a major factor 

determining vertical zonation of intertidal zoanthids (Herberts, 1972a). Herberts found Isaurus 
484 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 210.—Isaurus duchassaingi, transverse section across mesentery below the actinopharynx 

of hermaphroditic polyp: a, ova and mesenterial filament; b, ova and spermaries. (o = ovum, 

mf = mesenterial filament, s = spermary.) 

spongiosus near midtide level, intermediate between species of Zoanthus and Palythoa. Similarly, 

in Belize / duchassaingi was located between very 

shallow Zoanthus sociatus and deeper Palythoa 

sp. There we noticed that Z. sociatus tolerates a 

high degree of desiccation by contracting the 

column and closing the oral disc around the 

tentacles, whereas Palythoa sp. probably can survive only brief periods of exposure to air. Isaurus 

species, however, withstand moderate periods of 

exposure, during which they are dampened by 

wave swash and stray water droplets from pounding waves nearby. 

On the reef crest, Isaurus duchassaingi has 
NUMBER 12 485 

adapted to its habitat by protecting itself from 

desiccation and from abrasion by coral debris 

and potential predators. During daytime low 

tides, water loss from the polyps is minimized 

because the adults are partially buried and contracted, the capitulum is closed, and the body is 

near the moist substrate; the pale, smooth, thinner young polyps are never exposed, being sheltered by the adults above them. The adult polyps 

are also protected by a thick, tubercular column 

wall covered with a cuticle, sometimes with adhering unicellular algae (Duerden, 1898, 1902). 

Duchassaing and Michelotti (1860, 1864) noted 

the coriaceous tegument of I. duchassaingi. Herberts (1970, 1972b) remarked that her specimens 

had an ectodermal column thickness of 60-100 

/am and a 300-600 jum mesogleal layer that was 

even thicker at the level of the tubercles; Belize 

specimens are similar. Furthermore, abundant 

nematocysts in the column ectoderm (Andres, 

1877; Duerden, 1902; Carlgren, 1938; Herberts, 

1970, 1972b) may deter some predators from 

grazing on the polyps. Thus, behavioral adaptations as well as the animal's own morphological 

adaptations protect the colony from desiccation 

and abrasion. 

Although not an obvious limiting factor for 

distribution oi Isaurus, light is essential for growth 

of symbiotic zooxanthellae. Isaurus species are 

known to have zooxanthellae in the ectoderm and 

endoderm of the body wall, in the endoderm of 

the mesenteries, as well as in the endoderm of the 

tentacles (McMurrich, 1889; Haddon and Shackleton, 1891; Duerden, 1898, 1902; Carlgren, 1938; 

Herberts, 1970, 1972b; Trench 1974), although 

the significance of these dinoflagellates in Isaurus 

is unknown. Sebens and DeRiemer (1977) postulated that anthozoan structures adapted for 

photosynthesis—that is, those containing dense 

populations of zooxanthellae—respond positively 

to light (expansion, positive orientation). This is 

true for the exposed, sunlit—though contracted— 

column oi Isaurus. Trench (1974) showed that in 

the zoanthid Zoanthus sociatus, zooxanthellae contribute substances that are utilized. Sebens and 

DeRiemer (1977) noted the high oxygen production of zooxanthellae. Perhaps conservation of 

nutrients and energy is part of the reason for 

expansion and contraction cycles (Sebens and 

DeRiemer, 1977) in Isaurus species, but endogenous rhythms, individual physiological states of 

the polyps, and other external factors must also 

account for diurnal and nocturnal behavior patterns. For instance, after I. duchassaingi polyps are 

disturbed by light at night, why do they require 

many hours in the dark before reexpanding their 

tentacles? Obviously, light is important for zooxanthellae, which presumably benefit the zoanthids; however, the reason for tentacle retraction 

of Isaurus species during the day or in response to 

light at night remains undetermined. 

Located in a high energy area where waves are 

nearly always present, Isaurus duchassaingi from 

South Water Cay is ensured of a food supply of 

zooplankton and larger macroscopic prey. Sebens 

(1976) found that Stoichactis helianthus, abundant 

on reefs in Panama wherever there was heavy 

wave action, ate gastropods and echinoids 

brought in by the waves. Likewise, Isaurus spp. 

may capture prey tossed nearby; it has tentacle 

spirocysts (Carlgren, 1938; Herberts, 1970, 

1972b) capable of adhering to prey and has muscular tentacles capable of quickly transferring the 

food into its preoral chamber, which is simultaneously capped by the contracting capitulum. 

Competition with other organisms for space or 

food may be minimal for Isaurus duchassaingi colonies inhabiting the reef crest of South Water 

Cay. Being the largest sessile organism in the 

algal belt, /. duchassaingi can overgrow the smaller 

organisms in its niche and can rise above them 

when expanding to feed at night. Motile animals 

(such as mollusks, echinoids, crustaceans, and 

fish) could enter its habitat during high tides, but 

this infringement would be only temporary. Although /. duchassaingi may be rasped by fish or 

other invertebrates, or jabbed by shore birds, we 

saw no apparent damage to the zoanthids. 

Colony size and growth form of Isaurus duchassaingi vary. At South Water Cay, there were 

greater numbers of this zoanthid than have ever 

been reported elsewhere. The largest colony of/. 
486 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

duchassaingi had more than 100 animals in adjacent groups of five to six polyps, joined either by 

the coenenchyme or stolon; some polyps were 

separate, and the smallest group had two solitary 

forms. Duchassaing and Michelotti (1860, pl. 8: 

fig. 5) originally showed four adjacent polyps on 

one coenenchymal base, but in their later figures, 

they drew two polyps united by a stolon (Duchassaing and Michelotti, 1864, pl. 6: figs. 2, 3). 

McMurrich (1896, pl. 17: fig. 6) reported specimens from the Bahamas as solitary, but aggregated in groups of five to six, with one specimen 

having a bud connected to it. Duerden reported 

Jamaican and Puerto Rican polyps in small clusters, or more often, solitary (Duerden, 1898, pl. 

17: fig. 4; 1902, pl. 2: fig. 5). Other related species 

of Isaurus also vary in colony size and mode of 

growth, that is, solitary or colonial; and growing 

from stolons or coenenchyme (Gray, 1828; 

Andres, 1877; McMurrich, 1889; Haddon and 

Shackleton, 1891; Verrill, 1928; Carlgren, 1938; 

Herberts, 1970, 1972b; Day, 1969; Walsh and 

Bowers, 1977). 

Isaurus duchassaingi shows a remarkable variability in dimensions of the retracted column, in 

color, and in size, shape and distribution of tubercles. Our adult retracted specimens have average column lengths between 4 cm and 5 cm. 

Bermuda, Bahama, and some Caribbean specimens have shorter columns, such as reported by 

Duchassaing and Michelotti (1860, 1864), 

McMurrich (1889, 1896), and those examined by 

us (British Museum (Natural History): B.M. 

1901.3.8.58-60, B.M. 1927.9.9.1; American Museum of Natural History No. 691, No. 1290); 

many of these specimens do not exceed 3.5 cm in 

length. On the other hand, C. Arneson (pers. 

comm., 1979) has noted /. duchassaingi polyps as 

long as 12 cm. Therefore, column length oi Isaurus 

may vary according to age of the polyps, habitat, 

and individual states of contraction when alive 

and during preservation. 

Color of the exposed columns of Isaurus varies 

with species and location but generally blends 

with the surroundings. Hawaiian specimens, like 

the Caribbean ones, are brownish, with upper 

areas of the column bright green (Verrill, 1928; 

Walsh and Bowers, 1977). Animals from the Indian and Pacific oceans have some additional red 

color on the column, particularly at the level of 

the tubercles (Haddon and Shackleton, 1891; 

Carlgren, 1938; Herberts, 1970, 1972b). Color of 

the exposed portion of the polyps may depend on 

the genetic composition of the animal, and, to a 

greater extent, on zooxanthellae and epizoic algae, such as the algal tufts noted by Duerden 

(1898, 1902). The buried lower portion of the 

columns of all Isaurus species is whitish. 

Arrangement of column and crown tubercles 

appears to be random in all Isaurus species; distribution of tubercles on the concave and convex 

aspects of the columns varies from polyp to polyp. 

External morphology differs even among polyps 

that have grown from the same stolon. Column 

tubercles vary in shape from tiny verrucae to 

more alated protuberances, and may or may not 

be contiguous. For South Water Cay animals, 

tubercles are usually more prominent on the convex side, being situated randomly or in rows, or 

both, whereas the concave part has fewer, smaller 

tubercles or none at all; however, a preserved 

specimen from nearby Water Cay (B.M. 

1927.9.9.1) has prominent tubercles (in rows on 

the convex side, scattered on the concave) completely around the column. Other Caribbean 

specimens may have prominent tubercles scattered all around the polyp. Crown tubercles are 

prominent knobs or slight bumps, or both; these 

tubercles are in partial or complete circular arrangement. External features oi Isaurus may vary 

because of differential, endogenous rates of development, varying states of nutrition from prey 

or symbiotic zooxanthellae, or environmental influences. 

Asexual reproduction by budding from the 

gastrodermal solenia of the stolon and at the 

coenenchymal base of adults may be the most 

successful means of reproduction for Isaurus duchassaingi on South Water Cay. Colonial forms 

are numerous, most of them are adjacent to each 

other in groups of three to eight polyps that are 

joined by coenenchyme or anastomotic stolons, or 
NUMBER 12 487 

both. As their basal tissue atrophies from the 

stolon or coenenchyme, polyps become solitary. 

Little is known about sexual reproduction in 

zoanthids, probably because of small samples, 

inability to culture larval forms, or lack of seasonal data owing to short observation times. C. 

Arneson (pers. comm., 1979), who has observed 

Isaurus duchassaingi in aquaria for many years, has 

never seen them spawn. Previous authors did not 

mention gonads in species of Isaurus, except for 

noting female gametes in a few specimens (Duchassaing and Michelotti, 1860; Haddon and 

Shackleton, 1891; Herberts, 1970, 1972b). 

McMurrich (1889) was the only worker to find 

both ova and spermatozoa together in preserved 

Isaurus sp. polyps from Bermuda. Many polyps of 

/. duchassaingi from South Water Cay, Belize, are 

also hermaphroditic. As in McMurrich's descriptions (1889), the male and female gonads in 

Belizean polyps are irregularly arranged without 

specific positions along the mesenteries. Many 

Belizean polyps had only ova of various sizes, 

some that were possibly ready to be released; 

other polyps were sterile. Hence, it appears that 

/. duchassaingi from the Caribbean has monoecious 

and possibly also dioecious reproduction. Seasonal reproductive cycles have been reported for 

Zoanthidea from Okinawa and Hawaii (Yamazato et al., 1973; Cooke, 1976) and perhaps this 

is also the case for species oi Isaurus. Yamazato et 

al. (1973) have determined that some colonies of 

Palythoa tuberculosa Esper from Okinawa undergo 

sexual changes with the passage of time: female 

polyps change after six months to hermaphroditic 

polyps; after the ova are released, the testes remain and the polyps became functional males. 

Whether sexual changes occur in Isaurus remains 

undetermined until reproductive states of the 

polyps can be studied year-round. 

The natural prey oi Isaurus is unknown. Since 

/. duchassaingi expands its tentacles to feed only at 

night, it is a nocturnal carnivore. In the laboratory, /. duchassaingi preferred larval fish, and did 

not capture small crustaceans readily, except for 

Artemia salina nauplii. On the reef crest, however, 

/. duchassaingi may select larger organisms in the 

zooplankton as food because the zoanthid has a 

widened preoral chamber, which is rimmed by 

nematocyst-laden tentacles. Furthermore, the 

zoanthid has strong sphincter muscles that close 

the capitulum simultaneously with the introversion of the tentacles, thus facilitating swift containment of the prey. C. Arneson (pers. comm., 

1979) has kept /. duchassaingi from Puerto Rico in 

good condition in aquaria for over three years 

without feeding them. The importance of prey 

and zooxanthellae for the nutritional requirements of these zoanthids is unstudied. 

Literature Cited 

Andres, A. 

1877. On a New Genus and Species of Zoanthina malacodermata (Panceria spongiosa, sp.n.). Quarterly Journal of 

Microscopical Science, 17:221 -226. 

1883. Le Attinie [of the Gulf of Naples]. Atti della Reale 

Accademia dei Lincei, series 3a, 14:211-673. 

Carlgren, O. 

1938. South African Actiniaria and Zoantharia. Kungliga 

Svenska Vetenskapsakademiens Handlingar, 3rd series, 

17(3):1-148. 

Cooke, W. J. 

1976. Reproduction, Growth and Some Tolerances of 

Zoanthus pacificus and Palythoa vestitus in Kaneohe 

Bay, Hawaii. In G. O. Mackie, editor, Coelenterate 

Ecology and Behavior, pages 281-288. New York: 

Plenum Press. 

Day.J.H. 

1969. A Guide to Marine Life on South African Shores. 300 

pages. Cape Town: A. A. Balkema, for the University of Cape Town. 

Duchassaing, P., and J. Michelotti 

1860. Memoire sur les coralliaires des Antilles. Memoires 

de l'Academie des Sciences de Turin, 19(2):279-365, 10 

plates. 

1864. Supplement au memoire sur les coralliaires des 

Antilles. Memoires de l'Academie des Sciences de Turin, 
488 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

23(2):97-206, 11 plates. 

Duerden, J. E. 

1898. Jamaican Actiniaria, XIV, Part I: Zoantheae. The 

Scientific Transactions of the Royal Dublin Society, 6(2): 

329-376, plates 17-20. 

1902. Report on the Actinians of Porto Rico. United 

States Fish Commission Bulletin for 1900, 2:321-374, 

12 plates. 

Endean, R. 

1976. Destruction and Recovery of Coral Reef Communities. In O. A. Jones and R. Endean, editors, 

Biology and Geology of Coral Reefs, 3:215-254. New 

York: Academic Press. 

Gray, J. E. 

1828. Radiata. In Spicilegia Zoologica, page 8. London. 

Haddon, A. C , and A. M. Shackleton 

1891. Reports on the Zoological Collections Made in 

Torres Straits by Professor A. C. Haddon, 1888-

1889, Actiniae, I: Zoantheae. The Scientific Transactions of the Royal Dublin Society, 4(2):673-701. 

Herberts, C. 

1970. Note sur un zoanthaire de la cote sud-ouest de 

Madagascar. Recueil des Travaux de la Station Marine 

d'Endoume, Marseille, Supplement, 10:305-315. 

1972a. Contribution a l'etude ecologique de quelques 

zoanthaires temperes et tropicaux. Marine Biology, 

13(2): 127-136. 

1972b. Contribution a l'etude systematique de quelques 

zoanthaires temperes et tropicaux. Tethys Supplement, 3:69-156. 

McMurrich, J. P. 

1889. A Contribution to the Actinology of the Bermudas. 

Proceedings of the Academy of Natural Sciences of Philadelphia, 1889:102-126. 

1896. Notes on Some Actinians from the Bahama Islands, Collected by the Late Dr. J. I. Northrop. 

Annals of the New York Academy of Sciences, 9:181-

194, plates 17-20. 

Sebens, K. P. 

1976. The Ecology of Caribbean Sea Anemones in Panama: Utilization of Space on a Coral Reef. In G. 

O. Mackie, editor, Coelenterate Ecology and Behavior, 

pages 67-77. New York: Plenum Press. 

Sebens, K. P., and K. DeRiemer 

1977. Diel Cycles of Expansion and Contraction in Coral 

Reef Anthozoans. Marine Biology, 43:247-256. 

Trench, R. K. 

1974. Nutritional Potentials in Zoanthus sociatus (Coelenterata, Anthozoa). Helgoldnder Wissenschaftliche 

Meeresuntersuchungen, 26:174-216. 

Verrill, A. E. 

1928. Hawaiian Shallow Water Anthozoa. Bernice P. 

Bishop Museum, Bulletin, 49:29-30. 

Walsh, G. E., and R. L. Bowers 

1977. Order Zoanthiniaria. In D. M. Devaney and L. G. 

Eldredge, editors, Reef and Shore Fauna of Hawaii. Bernice P. Bishop Museum Special Publication, 

64(1): 148-157. 

Yamazato, K., F. Yoshimoto, and N. Yoshihara 

1973. Reproductive Cycle in a Zoanthid Palythoa tuberculosa Esper. In T. Tokioka and S. Nishimura, 

editors, The Proceedings of the Second International 

Symposium on Cnidaria, pages 275-283. Japan: Nippon Printing and Publishing Company Limited. 
Larval Settlement Behavior and Shell Morphology 

of Malleus candeanus (d'Orbigny) 

(Mollusca: Bivalvia) 

Thomas R. Waller 

and Ian G. Macintyre 

ABSTRACT 

Data on larval settlement behavior and shell 

morphology of the little known bivalve, Malleus 

{Malvufundus) candeanus (d'Orbigny, 1842), were 

obtained from an experiment designed for obtaining data on the distribution of boring microorganisms. Plastic (PVC) tubes with screened ends 

enclosing crushed coral fragments were set out on 

a transect line crossing the reef at Carrie Bow 

Cay, Belize. Epifaunal bivalves, half of which 

proved to be members of this species, attached to 

the containers during the 10-momh sampling 

period. The distribution of specimens of M. candeanus among opaque versus transparent tubes 

and the positions of attachment sites on the walls 

of containers suggest that their planktonic larvae 

are both photonegative and geonegative at time 

of settlement. In nature the larvae probably swim 

to a dark area beneath an overhang and then 

move laterally to the intersection of two surfaces, 

such as the bottom of a crevice, where they attach 

with a byssus and then metamorphose. 

Malleus candeanus trapped in the tubular containers approached full adult size in less than 10 

months. Unlike naturally occurring specimens 

they have a more regular shell outline and better 

developed posterior auricles and lack a central 

constriction and distal flare in the posteroventral 

prismatic extension of the shell. Therefore, the 

lack of a posterior auricle and presence of an 

irregular shell outline are probably phenotypic 

Thomas R. Waller and Ian G. Macintyre, Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560. 

characteristics that cannot be relied on in taxonomic studies. 

Introduction 

Malleus {Malvufundus) candeanus has received little attention in the literature on western Atlantic 

Mollusca since its original description by 

d'Orbigny (1842) as Avicula candeana (type-locality, Santiago de Cuba, in a large sponge). Johnson 

(1918) reported the presence of the species in 

Bermuda and briefly discussed its taxonomy. The 

only other detailed treatment is that of Boss and 

Moore (1967), who established that the species is 

broadly distributed, though "rarely encountered," in Bermuda, the tropical western Atlantic, 

and the tropical eastern Pacific. More recently, 

however, Waller (1973 and unpublished data) 

and Abbott (1974) found that the species is common among corals in Bermuda, the east coast of 

Florida, and in Belize, and Rosewater (1975 and 

unpublished data) found it to be common among 

rocks at Ascension Island in the South Atlantic. 

Specimens are seldom encountered not because 

they are rare, but because their cryptic living 

habit and irregular, fragile shell make them difficult to find and collect. 

The present study is concerned with specimens 

of this species found byssally attached in cylindrical plastic containers with screened ends that 

were placed on a transect line crossing the reef at 

489 
490 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Carrie Bow Cay, Belize. The distribution of these 

specimens among opaque and transparent containers and their preferred living positions yield 

information on the settling behavior of larvae; 

the size of the shells gives an estimate of minimum 

growth rate; and the shapes of the post-larval 

shells provide insight into the relationship between shell form and living habit. Comparison of 

specimens from the containers with those from 

natural habitats in the same area suggests that 

some features previously considered important in 

the taxonomy oi Malleus are phenotypic responses 

to living conditions. 

MORPHOLOGY AND SYSTEMATICS.—Like other 

members of the Order Pterioida (emend. Waller, 

1978), Malleus {Malvufundus) candeanus has a shell 

composed of an inner layer of nacreous aragonite 

and an outer layer of simple prismatic calcite 

(Taylor et al., 1969). As in the closely related 

species Malleus {Malvufundus) regulus (Forskal), the 

anatomy and habits of which have been described 

by Yonge (1968), nacreous aragonite is limited to 

the dorsal one-fourth of the shell and prismatic 

calcite forms a long extension of the shell margin 

in a posteroventral direction. The visceral-pedal 

mass occupies the nacreous dorsal region, whereas 

only the mantle and ctenidia lie in the prismatic 

extension. 

Several subgeneric names based primarily on 

the presence or absence of anterior and posterior 

extensions of the hinge have been introduced for 

groups within the genus Malleus. Boss and Moore 

(1967) questioned the taxonomic reliability of 

hinge extensions except in two groups: one having 

both anterior and posterior hinge extensions 

[Malleus {Malleus) Lamarck, the "hammer oyster"]; the other having "hardly any lateral extension of hinge line" and designated Malleus {Parimalleus) Iredale. Hertlein and Cox (1969) and 

Keen (1971) used the prior subgeneric name Malvufundus De Gregorio for the second group, 

thereby suggesting that minor differences in 

poorly developed hinge extensions originally used 

to separate Parimalleus and Malvufundus are insignificant for the distinction of subgenera. Variations in the development of a posterior auricle in 

M. candeanus are well documented: d'Orbigny 

(1842) showed a distinct posterior auricle; Johnson (1918) showed only a slight irregular posterior 

extension of the dorsal region; and Boss and 

Moore (1967) showed none. 

Many species names have been introduced for 

populations of the subgenus Malvufundus that occur in the tropical and warm-temperate regions 

of the world oceans. All exhibit a highly irregular 

shell form and tend to settle on hard substrates in 

crevices or beneath overhanging ledges. A taxonomic review of these nominal taxa has not yet 

appeared, and there is no agreement on the number of valid species. Boss and Moore (1967:87) 

have summed up the problem of relationships 

concisely: "The major problem with Malvufundus 

is whether or not there is only a single extremely 

variable species in the Mediterranean-Indo-Pacific region [Malleus regulus], and if so, whether 

that species can be truly differentiated from M. 

candeanus." 

In distinguishing the New World Malleus candeanus from the Mediterranean-Indo-Pacific species complex, Boss and Moore (1967) considered 

three regions of the shell: (1) a dorsal part frilled 

on the exterior by broad, overlapping sheets of 

prismatic calcite, some of which may fuse to form 

large, enclosed voids; (2) a central, narrow region—the "shank"—which is not strongly frilled 

on the exterior and is buttressed by a single rib 

on the inner shell surface; and (3) a flared, "fanshaped" ventral area. They suggested that this 

tripartite division of the shell is more distinct in 

M. candeanus than in other species of the subgenus 

Malvufundus. 

ECOLOGY.—In the vicinity of Carrie Bow Cay, 

Belize, specimens oi Malleus candeanus are common 

from a depth of 5 m in the high-relief spur and 

groove zone seaward to the deepest area explored 

by SCUBA diving, 46 m on the steeply inclined 

fore-reef slope (Figure 211). Specimens are less 

common in shallower water. In the wave-washed 

Millepora complanata Lamarck of the reef-crest zone 

and in the lagoon landward of Carrie Bow Cay, 

Malleus is uncommon and is replaced by Isognomon 

radiatus (Anton), which is very similar morpholog-
NUMBER 12 491 

Lagoon 

Thalassia 

Zone 

0 

Sand& 

Rubble 

Zone 

Back Reef 

Potch Reef Zone 

1100 

Rubble & Pavement Zone 

faoo 

Reef 

Crest 

Inner Fore Reef 

High Spur & Groove Zone tow Spur & Groove Zone 

1300 I400 I500 

Outer Fore Reef 

Inner 

Reef 

Slope 

Sand 

Trough 

Outer 

Ridge 

I600 

Fore Reef 

Slope 

Meters 

M5 M7 

1.2 

* * 

M6 M8 

1.5 

* * 

Station Number 

Station Depth (m) 

M3 M4 Mi l M12 Ml M2 M9M10 

24.4 12.2 27.4 

tt mt 

FIGURE 211.—Profile of reef at Carrie Bow Cay, Belize, showing location of stations, each 

station consisting of two containers, one opaque and the other transparent (asterisks indicate 

algal-covered and damaged containers that yielded no specimens). 

ically and ecologically, living in holes and crevices 

and beneath overhanging ledges. Although morphologically similar species of Isognomon and Malleus {Malvufundus) may co-occur in abundance in 

some regions of the Pacific Ocean (Yonge, 1968), 

this is not the case in Belize. 

Individuals of Malleus candeanus commonly live 

on the downward facing surfaces of overhanging 

ledges. The shells are attached by a strong byssus, 

with the dorsal or right anterodorsal region of the 

shell against the attachment surface and the posteroventral extension hanging downward. Specimens were common at the inner limits of overhangs, less common on vertical walls. In only one 

case were young specimens found on an upward 

facing surface; one individual was hanging downward inside a dead, articulated, nearly closed 

shell of Spondylus. As noted by Boss and Moore 

(1967), Waller (1973), and in the present study, 

specimens do not ordinarily live in sponges, the 

type description notwithstanding. Quantitative 

data on population density are lacking, but our 

observations during diving suggest that Malleus 

candeanus is not unusually abundant in the area of 

the reef transect. 

ACKNOWLEDGMENTS.—We are grateful to D. S. 

Putnam and J. Cooper for help in the field, W. 

Blow for asssistance in photography, and M.-J. 

Mann for operation of the scanning electron microscope. E. G. Kaufmann, Department of Paleobiology, and J. Rosewater, Department of Invertebrate Zoology, Smithsonian Institution, kindly 

read the manuscript and offered suggestions for 

its improvement. 

Materials and Methods 

Twelve pairs of sediment containers holding 

crushed coral fragments were placed at intervals 

along a transect line crossing the reef at Carrie 

Bow Cay, Belize, by Macintyre for the purpose of 

gathering information on distribution of boring 

microorganisms (unpublished). Each container 

was a cylinder of either opaque gray or transparent colorless plastic PVC (polyvinyl chloride) 

pipe 65 mm in length and 50 mm in diameter 

and each had gray, plastic-coated fiberglass 

screening (mesh size about 1 mm) fastened with 

epoxy resin over each end. The containers were 

positioned in pairs, one opaque and one transparent, at each station in the manner shown in 

Figure 212. Distance from the containers to the 

bottom was approximately 50 cm. Distribution of 

stations along the transect line is shown in a 

profile (Figure 211), and the location of the transect is shown by Riitzler and Macintyre (herein, 

Figure 4). The containers were set out between 7 

June and 13 June 1972 and were collected between 5 April and 11 April 1973, approximately 

10 months later. 
492 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 212.—Substrate station M6 in the patch reef zone. 

Browsing fish have destroyed the screened ends of the cylindrical sediment containers. 

The containers were placed in neutral buffered 

formalin at the time of collection. Because it was 

not apparent until they were emptied at a later 

time that they contained Malleus, no on-site data 

on the final orientation of the cylinders or of the 

exact living positions of bivalves in them were 

taken. The top and bottom of the containers, 

however, could be inferred later from the distribution of encrusting organisms, the top surface 

generally being covered with algae and the bottom having sparsely distributed worm tubes. The 

attachment sites of Malleus on the interiors of 

several of the cylinders could be determined by 

examination with a binocular microscope, because the byssus was left attached to the wall of 

the container when specimens were removed. 

The dorsal region of the shell of one small 

specimen from station Mil (USNM 782900, Figure 213) was washed in neutral distilled water, 

sputter coated with carbon and then gold palladium to a thickness not exceeding 7.5 nm, and 

examined in a Coates and Welter 106B scanning 

electron microscope. 

Results and Discussion 

Containers shoreward of the low-relief spur and 

groove zone on the transect (stations M5-M8, 

Figure 211) were covered with algae and their 

screens were destroyed by browsing animals. Because they contained no specimens of Malleus or 

of any other epifaunal bivalves on either their 

outer or inner surfaces, they will not be considered 

further. 

In contrast, the eight pairs of containers seaward of the high-relief spur and groove zone 

remained intact (with the exception noted below) 

and contained an assemblage of five species of 

living epifaunal bivalves at the time of collection. 

The distribution of these specimens with respect 

to the inside and outside of opaque and transparent containers at each station is given in Table 

42. Thirteen of the 27 individuals tabulated are 

Malleus candeanus, a surprising result in view of the 

fact that there does not seem to be an unusually 

dense population of the species in this area. 

A striking difference in settlement behavior 

among the five species of bivalves can be seen in 

Table 42. Of the 13 specimens of Malleus candeanus, nine attached to the inner walls of the 

cylinders. Only one individual of the 14 specimens of other species was found inside a cylinder; 

all of the other specimens were on outer walls or 

on outer sides of screens. Furthermore, all of the 

specimens of M. candeanus found on inner walls 

were in opaque containers. In contrast, individuals of all species attached to outer walls were 

about evenly divided between opaque and transparent containers. 

The veliger larvae of Pinctada, Spondylus, and 

Chama are known to be planktonic (MartinezEscarbassiere, 1970; Kennedy et al., 1970; LaBarbera and Chanley, 1971; Rosewater, 1975; 

Waller, unpublished data), lifting themselves off 

the bottom by the action of a ciliated velum. 

Larvae of Lopha, after an initial brooded stage 

(Stenzel, 1971), are also planktonic. Although 

living larvae of Malleus candeanus have never been 

observed, their small prodissoconch-I stage (now 

known to represent the initial phase of larval 
NUMBER 12 493 

TABLE 42.—Epifaunal bivalves in or on cylindrical containers at Carrie Bow Cay, Belize 

(stations listed landward to seaward for each species) 

Species 

Malleus candeanus (d'Orbigny) 

Pinctada imbricata Roding 

Spondylus sp. 

Lopha frons (Linnaeus) 

Chama sp. 

Station 

M3 

M4 

M i l 

M12 

Ml 

M2 

MIO 

M3 

M9 

M3 

M4 

M12 

MIO 

M i l 

M9 

Number of 

Opaque 

containers 

Inside 

0 

1 

3 

1 

1 

2 

1 

0 

0 

0 

1 

0 

0 

0 

0 

Outside 

0 

2 

1 

0 

0 

0 

0 

0 

1 

2 

0 

2 

0 

0 

0 

specimens 

Transparent 

containers 

Inside 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

Outside 

1 

0 

0 

0 

0 

0 

0 

2 

0 

1 

0 

0 

2 

1 

2 

Maximum 

dimension 

of measured 

(mm) 

12 

37 

24 

41 

(broken) 

40 

(broken) 

14 

5.5 

16 

13 

5.6 

28 

5 

5 

growth before the valves of the larval shell can 

enclose the body) and relatively large prodissoconch-II stage (Figure 213) indicate a planktonic, planktotrophic mode of life (Ockelmann, 

1965; Waller, 1981). A planktonic mode of life is 

also indicated by the fact that the larvae entered 

containers 50 cm off the bottom. 

Th~ fact that all of the specimens of M. candeanus occurring on inner walls were in opaque 

containers cannot readily be explained except in 

terms of larval behavior during the swimming 

and crawling stage just prior to and at the time 

of settlement. The sizes of prodissoconchs (larval 

shells) of all species found on and in containers 

(Table 43) are well below the 1 mm mesh size of 

the screens. Despite the reduction in size of mesh 

openings caused by algal encrustations, these 

openings were still adequate to allow penetration 

of bivalve larvae. It thus appears that larvae of 

M. candeanus are distinctly photonegative at the 

time of settlement and that they have a greater 

ability to select and enter dark, obstructed spaces 

than other larvae present during the sampling 

period. 

Additional information on the settling behavior of Malleus candeanus can be obtained from the 

attachment sites on container walls. Three byssal 

attachments found on the inside walls were at or 

near the top adjacent to a plug of epoxy resin, 

which was holding the rope that fixed the container to the pole (Figure 212). Attachment sites 

on the outside walls were either at the bottom, 

between the tube wall and the folded-back edge 

of the screen, or (in one case) in a downward 

opening crevice formed between an attached oyster and the wall of the container. The larvae of 

M. candeanus thus appear to be geonegative as well 

as photonegative at the time of settlement. They 

probably swim upward until they contact a suitable overhang and then move laterally until they 

perceive a crevice or an intersection of two surfaces. 

The sizes of the shells of trapped specimens of 

Malleus candeanus give a rough indication of 

growth rate. With the exception of one individual 

(station Ml), all specimens in the cylinders were 

living and presumably actively growing at the 

time of collection. The only dead specimen, which 
494 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 213.—Prodissoconch oi Malleus (Malvufundus) candeanus from station Mil , USNM 

782900: a, dorsal view of dorsal region, anterior toward top, X 43, scale bar = 200 jum; b, detail 

of right valve of prodissoconch showing small prodissoconch-I stage (top) and relatively large, 

commarginally ridged prodissoconch-II stage, X 280, scale bar = 50 ftra. 

TABLE 43.—Prodissoconch size of specimens on or in containers at Carrie Bow Cay, Belize 

Species 

Malleus candeanus 

(d'Orbigny) 

Pinctada imbricata 

Roding 

Spondylus sp. 

Lopha frons 

(Linnaeus) 

Chama sp. 

Number of 

specimens -

measured 

13 

1 

1 

4 

1 

Length of 

prodissoconch (/im) 

Min. 

265 

-

-

398 

-

Max. 

326 

-

428 

Mean 

288 

306 

194 

416 

184 

had a circular hole with beveled edge on its left 

valve, was killed by a predatory gastropod that 

entered the container through a large hole in one 

of the screens. Measuring from beak to the margin 

of the posteroventral prismatic extension of the 

shell, we observed a maximum size of 56 mm in 

shells of this species in the Division of Mollusks, 

National Museum of Natural History (USNM 

457011, Southwest Channel, Tortugas, Florida, 

depth 18 m). The largest trapped individual (40 

mm, station Ml 2) had attained 73 percent of the 

size of the largest museum specimen and must 

have been approaching maturity. If it be assumed 

that the larva of the 40 mm specimen entered the 

container at the beginning of the 10-month sampling period, its growth rate would be 4 mm per 

month. This represents a minimum estimate; if 

the larva entered later, the calculated growth rate 

would be higher. 

The shells of specimens removed from the containers differ from those of specimens living under 

natural conditions in that they have a posterior 

auricle and lack a central "shank" and distal flare 

(Figure 214). Both differences appear to result 

from the unusual environment in which larvae 

settled. Compared to natural settlement surfaces, 

the walls of the plastic tubes were relatively free 

of epifauna and presented a smooth surface for 

attachment. Subsequent growth away from the 
NUMBER 12 495 

10 mm 

FIGURE 214.—Malleus (Malvufundus) candeanus: a, right side of specimen from inside container at 

Station M4, USNM 782901; b, right side of specimen from natural habitat on Carrie Bow Cay 

reef at depth of 9 m, USNM 782902; c, left side of specimen from inside container at Station 

Ml2, USNM 782903; all X 2, lightly coated with ammonium chloride. 

surface was into a protected and, in view of the 

small maximum size of this species, unconfined 

space. 

The development of a posterior auricle is apparently hindered in naturally occurring specimens because dorsal shell expansion is confined 

to a narrow or irregular space. Among museum 

specimens, a posterior auricle is absent from specimens of Malleus candeanus and closely related or 

conspecific Pacific taxa collected from reefs, 

where they presumably lived in holes and crevices. In contrast, a posterior auricle is present in 

many specimens of M. {Malvufundus) regulus, 

which appears to occur on bare rock surfaces 

more commonly than in holes and crevices 

(Yonge, 1968). Extreme development of a posterior auricle and an anterior auricle-like extension 

of the dorsal margin occurs in M. {Malleus), the 

"hammer oyster." Yonge (1968:387), in referring 

to M. malleus, said that the shell "is always buried 
496 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

with only the distal quarter or less exposed" and 

that "the extent of exposure is usually revealed 

by the presence of encrusting organisms absent 

from buried regions of the shell." In specimens of 

this species in the collections of the National 

Museum of Natural History both valves are entirely encrusted with organisms that were alive at 

the time of collection, and thus we infer that these 

specimens cannot have been buried in the manner 

assumed by Yonge to be typical. They more likely 

were attached to rocky surfaces or to rubble on 

the bottom, as reported by Chuang (1961) and 

R. T. Abbott (pers. comm.). 

These differences between specimens living under natural conditions and those living in artificial, protected, unconfined spaces support suppositions that minor differences in hinge extension are unimportant for the distinction of subgenera. They further suggest that minor differences 

in shell outline, such as the central constriction 

and distal flare of the prismatic region thought 

by Boss and Moore (1967) to be useful in distinguishing M. candeanus from congeneric species, 

may also be unreliable. 

Conclusions 

The planktonic planktotrophic veliger larvae 

of Malleus {Malvufundus) candeanus respond negatively to light and gravity at the time of settlement. These responses allow them to seek out 

dark crevices and holes beneath overhanging 

ledges for byssal attachment and subsequent 

growth. M. candeanus can attain a maximum dimension of about 40 mm, which is about threefourths of its maximum known size, within a 

period of time not exceeding 10 months and 

possibly considerably less. The degree of development of a posterior auricle, a central constriction, and a distal flared portion of the shell are 

all phenotypic responses to the nature of the 

attachment surface and are therefore not reliable 

subgeneric and specific differentia. 

Literature Cited 

Abbott, R. T. 

1974. American Seashells. Second edition, 663 pages, illustrated. New York: Van Nostrand Reinhold Co. 

Boss, K. J., and D. R. Moore 

1967. Notes on Malleus (Parimalleus) candeanus (d'Orbigny) (Mollusca: Bivalvia). Bulletin of Marine Science, 7(l):85-94, 2 figures. 

Chuang, S. H. 

1961. On Malayan Shores, xvi + 225 pages, 28 figures, 112 

plates. P.O. Box 1813, Singapore: Muwu Shosa. 

d'Orbigny. See Orbigny, A. d' 

Hertlein, L. C , and L. R. Cox 

1969. Family Malleidae Lamarck, 1819. In R. C. Moore, 

editor, Treatise on Invertebrate Paleontology, Part N: 

Mollusca, 1:326-332. Lawrence, Kansas: Geological Society of America and University of Kansas 

Press. 

Johnson, C. W. 

1918. The Avicula candeana of d'Orbigny, from Bermuda. 

The Nautilus, 32(2):37-39, plate 3. 

Keen, A. M. 

1971. Sea Shells of Tropical West America. Second edition, 

xiv + 1064 pages, illustrated. Stanford, California: 

Stanford University Press. 

Kennedy, W. J., N. J. Morris, and J. D. Taylor 

1970. The Shell Structure, Mineralogy, and Relationships of the Chamacea (Bivalvia). Palaeontology 

13(3):379-413, 7 figures, plates 72-77, 2 tables. 

LaBarbera, M., and P. Chanley 

1971. Larval and Postlarval Development of the Corrugated Jewel Box Clam Chama congregata Conrad 

(Bivalvia: Chamidae). Bulletin of Marine Science, 

21(3): 733-744, 8 figures. 

Martinez-Escarbassiere, R. 

1970. Descripcion de la larva veliconcha y postlarva 

(disoconcha) de la ostra perlifera Pinctada imbricala 

Rdding del oriente de Venezuela. Memoria, Sociedad 

de Ciencias Naturales La Salle, 87:183-195, 5 figures, 

1 table. 

Ockelmann, K. W. 

1965. Development Types in Marine Bivalves and Their 

Distribution along the Atlantic Coast of Europe. 

In L. R. Cox and J. F. Peake, editors, Proceedings of 

the First European Malacological Congress (London, Sep-
NUMBER 12 497 

tember 17-21, 1962), pages 25-35, 5 figures. London: Conchological Society of Great Britain and 

Ireland and the Malacological Society of London. 

Orbigny, A. d' 

1842. Atlas in Mollusques. In R. de la Sagra, Histoire 

physique, politique et naturelle de Vile de Cuba, [volumes 6, 7], atlas, 28 plates. Paris: Libraire de la 

Societe de Geographie et de la Societe Royale des 

Antiquaires du Nord. 

Rosewater, J. 

1975. An Annotated List of the Marine Mollusks of 

Ascension Island, South Atlantic Ocean. Smithsonian Contributions to Zoology, 189: iv + 41 pages, 24 

figures, 3 tables. 

Stenzel, H. B. 

1971. Oysters. In R. C. Moore, editor, Treatise on Invertebrate Paleontology, Part N: Mollusca, 3:953-1224. 

Lawrence, Kansas: Geological Society of America 

and University of Kansas Press. 

Taylor, J. D., W. J. Kennedy, and A. Hall 

1969. The Shell Structure and Mineralogy of the Bivalvia, Introduction: Nuclacea-Trigonacea. Bulletin of the British Museum (Natural History), Zoology, 

supplement, 3: 125 pages, 77 figures, 29 plates. 

Waller, T. R. 

1973. The Habits and Habitats of some Bermudian 

Marine Mollusks. The Nautilus, 87(2):31-52, 33 

figures. 

1978. Morphology, Morphoclines and a New Classification of the Pteriomorphia (Mollusca: Bivalvia). 

In C. M. Yonge and T. E. Thompson, editors, 

Evolutionary Systematics of Bivalve Molluscs. 

Philosophical Transactions of the Royal Society of London, 

series B, 284:345-365, 2 figures, 2 tables. 

1981. Functional Morphology and Development of Veliger Larvae of the European Oyster, Ostrea edulis 

Linne. Smithsonian Contributions to Zoology, 328: iv 

+ 70 pages, 152 figures, 1 table. 

Yonge, C. M. 

1968. Form and Habit in Species oi Malleus (Including 

the "Hammer Oysters") with Comparative Observations on Isognomon isognomon. The Biological 

Bulletin, 135(2):378-405, 13 figures, 1 table. 
Habitat and Resource Partitioning 

between Two Species of Acanthemblemaria 

(Pisces: Chaenopsidae), with Comments 

on the Chaos Hypothesis 

David W. Greenfield 

and Teresa A. Greenfield 

ABSTRACT 

Four of the eight western Atlantic species of 

the chaenopsid fish genus Acanthemblemaria occur 

in Belize, Central America. At Carrie Bow Cay 

on the barrier reef two inquiline species, A. spinosa 

and A. greenfieldi, exhibit subtle habitat separation: A. spinosa occurs only in holes left by invertebrates on vertical surfaces of dead coral, and A. 

greenfieldi occurs only in holes left by invertebrates 

on horizontal surfaces of dead coral. Consequently, A. spinosa individuals are found primarily 

just behind the reef crest, whereas A. greenfieldi 

individuals are most abundant in the back reef, 

although both species can be found on the same 

coral head. A total of 168 specimens of A. greenfieldi and 90 specimens of A. spinosa was collected 

and analyzed for length-frequency, stomach contents, and fecundity. 

Length-frequency analysis showed that Acanthemblemaria greenfieldi is larger than A. spinosa and 

that in both species males are larger and more 

numerous than females. Stomach content analysis 

revealed different preferential feeding of each 

species among the same groups of invertebrates. 

Fecundity analysis confirmed that females of A. 

spinosa mature at a smaller size than those of A. 

greenfieldi. Smaller females (including most A. spinosa females) have fewer and smaller eggs than 

David W. Greenfield and Teresa A. Greenfield, Department of 

Biology, Northern Illinois University, DeKalb, III. 60115; and 

Division of Fishes, Field Museum of Natural History, Chicago, 

III. 60605. 

larger females (including many A. greenfieldi females). The fine habitat partitioning exhibited 

by these and other coral-reef fishes suggests that 

the chaos hypothesis of habitat selection may only 

apply to larger species. 

Introduction 

The chaenopsid fish genus Acanthemblemaria is 

represented by eight species in the western Atlantic Ocean (Smith-Vaniz and Palacio, 1974). Four 

of these eight species occur in Belize, Central 

America: A. aspera (Longley), A. greenfieldi SmithVaniz and Palacio, A. maria Bbhlke, and A. spinosa 

Metzelaar; all except A. maria have been taken on 

the barrier reef at Carrie Bow Cay, (Greenfield 

and Johnson, 1981). 

At Carrie Bow Cay, Acanthemblemaria aspera has 

been taken only at a depth of 9 m in the lowrelief spur and groove zone of the inner fore reef; 

however, both A. spinosa and A. greenfieldi occur in 

shallow areas on the reef crest and across the back 

reef. Both of these species are inquiline, dwelling 

in holes left by worms or other boring invertebrates (Figure 215). The present study was 

prompted by our observation of subtle differences 

in habitats during a faunal survey at Carrie Bow 

Cay. 

ACKNOWLEDGMENTS.—We are indebted to the 

499 
500 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 215.—Acanthemblemaria greenfieldi in natural habitat. 

government of Belize, and especially to G. W. 

Miller, Fisheries Administrator, for granting permission to collect fish specimens. P. Illg, University of Washington, kindly helped to identify food 

items and C. F. Rakocinski, Northern Illinois 

University, assisted in the field. J. Glaser, Northern Illinois University, prepared the graphs. H. 

Bowman, Jr., was of considerable assistance in 

arranging field logistics. This research was supported in part by National Science Foundation 

Grant BMS75-08684 to D. W. Greenfield. 

Materials and Methods 

Specimens were collected individually by 

squirting into their holes a mixture of quinaldine 

and isopropanol, which caused them to leave 

their holes. After the anesthetic effect of the quinaldine took effect, individuals were picked up by 

hand and placed into plastic bags labeled with 

detailed information on habitats. Six collections 

were made during the period 10-17 May 1977 

and three collections during 1-4 January 1978. A 

total of 168 specimens of Acanthemblemaria greenfieldi and 90 specimens of A. spinosa was taken. 

Lengths, which are given as the standard 

length (SL) in mm, were made to the nearest 0.1 

mm using dial calipers. Twenty specimens each 

of A. greenfieldi (24.3-33.8 mm SL) and A. spinosa 

(19.8-26.4 mm SL) were analyzed for stomach 

contents: eight individuals of each species were 

taken at 0930 h on 16 May 1977 and 12 of each 

species at 1500 h on 15 May 1977, and the data 

combined. Food items were sorted by taxa, 

counted, and their volume determined using a 

squash technique modified from Hellawell and 

Abel (1971) by Ross (1974:8, 1977:561). Relative 

importance of each food category was evaluated 

using the percentage of occurrence of food items 

in stomachs, the number of food items in each 

category, and the volumetric contribution of each 

prey category to total stomach volume. The relative importance of each food category was determined by the index of relative importance 

(IRI) (Pinkas 1971:9): F (N+V) = IRI, where N 

= numerical percentage, V = volumetric percentage, and F = frequency of occurrence percentage. The degree of food overlap between the 

two species of Acanthemblemaria was measured using the overlap coefficient of Horn (1966:420): 

CA = 

2 J XiYi 

i=l 

i x\ + i n 

i=l i=l 

CA varies from 0 (when two species have no food 

category in common) to 1 (when the proportions 

of each food category are identical); s = total 

number of food categories; Xi is the proportion of 

the total diet of species X for food category i; Fi 

is the similar proportion of the total diet of species 

Y. 

Eggs were counted and measured from 36 Acanthemblemaria greenfieldi females (18 in January; 18 

in May) and from 22 A. spinosa females (12 in 

January; 10 in May) with the aid of a dissecting 

microscope fitted with an ocular micrometer. 

Eggs were categorized by size class (ovum diameter) into separate broods: <0.10 mm, 0.11-0.20 

mm, 0.21-0.30 mm, 0.31-0.50 mm, 0.51-0.75 
NUMBER 12 501 

A. greenfieldi 

-A. spinosa 

FIGURE 216.—Relative positions oi Acanthemblemaria greenfieldi and A. spinosa on a single brain 

coral. 

mm, 0.76-1.00 mm. Individual broods contained 

from two to 28 eggs, and individual females 

contained from two to three broods. Average 

brood size was determined by averaging the number of eggs in all broods for all females of each 

individual standard-length measurement. 

Results 

Acanthemblemaria spinosa occurs only in holes left 

by invertebrates on vertical surfaces of dead coral. 

This vertical habitat is present in three distinct 

situations: dead basal portions of living trees of 

Acropora palmata (Lamarck), vertical surfaces of 

fallen dead trees of A. palmata, and less commonly 

on vertical surfaces of other corals. The restriction 

of this species to vertical surfaces results in an 

uneven horizontal distribution across the reef, 

with most individuals being found just behind 

the reef crest and in reduced numbers in the back 

reef. 

Acanthemblemaria greenfieldi occurs only in holes 

left by invertebrates on the horizontal surfaces of 

dead coral. As a result, this species is most abundant in the back reef, which is composed of coral 

rubble and pavement. Less commonly, this species occurs closer to the reef crest on the horizontal 

surfaces of fallen dead trees oi Acropora palmata. 

On 15 May 1977, 44 specimens oi Acanthemblemaria were collected without regard to species 

identification. They were placed into plastic bags 

labeled "horizontal" or "vertical" depending on 

the location of capture. When the collection was 

subsequently examined it was apparent that 20 

A. greenfieldi had been taken from the horizontal 

surfaces and 24 specimens of A. spinosa from the 

vertical surfaces. A single brain coral, Diploria sp., 

65 cm high and 60 cm in diameter, located on 

the rubble and pavement zone of the back reef, 

provided a striking example of the sharpness of 

the habitat separation between these two species. 

The top of the brain coral, which was dead and 

covered by sediment, provided a horizontal surface. A small dead portion on the side of the coral 

head also provided suitable vertical habitat. Specimens of A. greenfieldi were living on the top of the 
502 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

coral head, whereas A. spinosa was living on the 

side (Figure 216). Additional collections made for 

the feeding and life history studies have also 

demonstrated this separation. 

On 4 January 1978, a collection was made in 

the coral rubble and pavement zone of the back 

reef just south of Carrie Bow Cay to obtain 

specimens of Acanthemblemaria greenfieldi for life 

history information. This location was chosen 

because it is a typical A. greenfieldi habitat and 

samples taken in May 1977 had contained only 

members of this species. Examination of the new 

series of 53 specimens, however, showed that it 

included not only 36 A. greenfieldi (7.6-30.3 mm 

SL; x = 24.5 mm SL), but also 17 young A. 

spinosa (13.4-18.4 mm SL; x = 15.7 mm SL). 

Length-frequency analysis (Figure 217) shows 

that Acanthemblemaria greenfieldi is a larger species 

than A. spinosa and that males of both species are 

larger than females: A. greenfieldi males attain a 

length of at least 32.5 mm SL (x = 25.5 mm) and 

females reach 30.0 mm SL (x = 23.5 mm), 

whereas A. spinosa males attain a length of 26.4 

mm SL (x = 19.9 mm) and females reach 23.2 

mm SL (x = 18.4 mm). Males of both species also 

outnumbered females in our collections; males 

constituted 61% (January) and 64% (May) of the 

total A. greenfieldi sample, and 60% (January) and 

65% (May) of the total A. spinosa sample. Within 

each species, the proportions of individuals within 

each size range were basically similar for both 

January and May, with the majority of A. greenfieldi individuals falling between 20 and 30 mm 

SL and the majority of A. spinosa individuals 

falling between 15 and 25 mm SL for both 

months. 

A fecundity analysis (Figure 218) revealed that 

Acanthemblemaria spinosa females mature at a 

smaller size (12.4 mm SL) than do A. greenfieldi 

females (19.0 mm SL), as might be expected on 

the basis of the smaller overall size range of A. 

spinosa. Females of both species contained from 

two to three broods, or size classes of eggs. Predictably, the largest females {A. greenfieldi) had 

more eggs than the smallest females (A. spinosa). 

In addition, the largest eggs of the largest A. 

greenfieldi females averaged 0.76-1.00 mm in diameter, whereas the largest ones of the smaller 

females, including A. spinosa, averaged 0.51-0.75 

mm in diameter. 

An analysis of the food habits of these two 

species during the month of May has shown that 

although they may feed on the same groups of 

invertebrates, the dependence on, or preference 

for, specific taxa differs considerably. The overlap 

coefficient yields a value of 0.4, indicating a major 

separation. The index of relative importance 

shows that both species depend heavily on copepods; however, the next item of importance for 

Acanthemblemaria spinosa is isopods, whereas isopods are only the fourth most important food 

20 

10 

0 

20 h 

10 

A. spinosa 

DO. 

MAY 1977 

am vm 

JAN 1978 

5-9 10-14 15-19 20-24 25-29 30-34 

Standard Length, mm 

FIGURE 217.—Length frequencies oi Acanthemblemaria spinosa 

and A. greenfieldi males and females from Carrie Bow Cay, 

May 1977 and January 1978. 
NUMBER 12 503 

10 -

tn 30 - A 

20 -

A. greenfieldi 

• JAN 

• MAY 

A. spinosa 

O JAN 

• MAY 

0 

- o a D 

I,I , 

0 

• 

i • 

O 

• 

a 

1 

o 

• 

• 

• 

1 

0 

• 

a 

• 

i 

• 

• 

1 

• 

• 

• 

• 

I 

• 

• 

• 

1 

• 

• 

12 14 16 18 20 22 24 

Standard Length, mm 

26 28 30 

FIGURE 218.—Average brood size vs. SL of females oi Acanthemblemaria greenfieldi and A. spinosa 

from Carrie Bow Cay, May 1977 and January 1978. 

category for A. greenfieldi (Figure 219), which 

depends more on ostracods and tanaids (Figure 

220). 

Discussion and Conclusions 

Acanthemblemaria spinosa and A. greenfieldi exhibit 

extremely fine habita t partitioning; they achieve 

100% separation as adults by using holes mad e 

by invertebrates on vertical or horizontal surfaces. 

The relatively low level of food overlap between 

these two species (CA = 0.4) is also probably the 

result of their habita t separation. Both species 

feed by two methods, either by extending partially out of their holes and taking food items 

from the substrate near the hole, or by darting 

out and taking plankters. Th e plankters taken by 

our specimens were mainly harpactacoid copepods, a group usually associated with the substratum. Food organisms of second importanc e were 

isopods for A. spinosa and ostracods for A. greenfieldi. Th e vertical dead coral surfaces where A. 

spinosa is found tend to be covered with filamentous algae that provide a typical substrate for 

isopods. Th e horizontal surfaces where A. greenfieldi occurs are generally exposed to sedimentation so that sand surrounds the holes in the coral, 

providing suitable ostracod habitat. 

T h e 4 Januar y 1978 collection from a typical 

Acanthemblemaria greenfieldi habitat, which also 

yielded young of A. spinosa, provides valuable 

information concerning the interactions of these 

two species. Although the adults exhibit complete 

habitat separation, apparently planktonic larval 

Acanthemblemaria species settle and occupy any 

available hole or crevice. Because individuals of 

A. spinosa found here were only 18.4 mm or less, 

it may be assumed that when they outgrow their 

holes and search for a larger hole, they are excluded by A. greenfieldi from this habitat. Stephens 

et al. (1970:228) have described similar shifts in 

holes with growth for the California blenny Hypsob lennius Jenkins i. Because A. greenfieldi is a larger 

species, it might have an advantage in holding a 

hole. 

T h e horizontal habitat of Acanthemblemaria 

greenfieldi compared to that of A. spinosa is generally in areas of lower wave and surge action. 

Collecting A. spinosa just behind the reef crest is 

difficult because of the strong surge that tends to 

sweep the diver back and forth, whereas collecting 

A. greenfieldi does not present this problem. If the 

force of the water behind the reef crest poses a 

problem for a diver, it is logical to assume that it 

would be an equal or greater problem for A. 

spinosa when it darts out to obtain a food item. It 
504 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

may be that the horizontal habitat available in 

the back reef is more advantageous for Acanthemblemaria, but that A. spinosa is excluded from the 

superior habitat by A. greenfieldi. 

The fine partitioning of habitat by these two 

species of Acanthemblemaria is particularly interesting in light of Sale's (1977) views concerning 

habitat partitioning by coral-reef fishes. Sale concluded (1977:341): "We do not find, as a general 

rule, a high degree of specialization on microhabitats and a resulting fine partitioning of living 

space." He also noted that "there are a minority 

of species on reefs with highly specialized habitat 

needs, especially among fish with inquiline 

habits;" however, "the habitat requirements of 

coexisting habitat specialists often overlap 

greatly." Sale listed four examples from the literature of sponge-dwelling gobies, coral-dwelling 

100 -

% Frequency of Occurrence 

gobies, and anemone fishes where two or more 

species were collected in the same sponge, coral 

or anemone. Until more detailed studies have 

been conducted, however, a single organism, such 

as a sponge, should not necessarily be considered 

to provide a single habitat. Position of fishes 

within the sponges must first be investigated, for 

it is certainly possible that one species may be 

restricted to the opening and another to the inner 

part of the sponge atrium. 

The chaenopsid blennies found in Belize provide an example of a group of fishes that have 

finely partitioned the habitat. The two species of 

Acanthemblemaria investigated in this study demonstrate that a single, small brain coral cannot 

be considered a single habitat because habitat 

partitioning was effected by distinguishing between horizontal and vertical surfaces. Other 

chaenopsid species provide additional examples. 

100 

80 -

.S 60 

E 

4 0 -

20 -

20 -

4 0 = 

— Acanthemblemaria greenfieldi 

A 

. 

— 

_ 

-

-

-

I R I 

A. COPEPODA - 11770 

B. OSTRACODA -1834 

C. TANAIDACEA -49 8 

D. ISOPODA -22 5 

E. AMPHIPODA -114 

F ANNELIDA -101 

G. DECAPODA - 48 

H. GASTROPODA -14 

1. CIRRIPEDIA - 4 

J. INSECTA - 0 

B 

7 0 % 

GHI 

C 

3 0% 

U 

30% 

£ F\W 

^ 

|207 

15% 5% 

9 5% 

% Frequency of Occurrence 

FICURE 219.—Percent composition of food by taxonomic group in number, volume, and 

frequency of occurrence in two species of Acanthemblemaria: a, A. spinosa; b, A. greenfieldi. (IRI = 

index of relative importance.) 
NUMBER 12 505 

Emblemariopsis pricei Greenfield may occur in the 

same tree of Acropora palmata as A. spinosa, but A. 

spinosa is restricted to the basal dead portions and 

E. pricei is found only in holes in the living portion 

of the coral (Greenfield, 1975:714). Acanthemblemaria aspera occurs more commonly on the dead 

bases of Montastrea heads and Emblemaria pandionis 

Evermann and Marsh is found only in a very 

specific habitat, for instance empty worm or clam 

holes in small pieces of coral rubble occurring on 

open sand substrate in areas of moderate current 

(Greenfield and Johnson, 1981). Lucayablennius 

zingaro (Bohlke), which occurs in deep water, is 

usually found free-living when feeding (Greenfield, 1972:590). 

Examples of fine habitat partitioning among 

coral-reef fishes in Belize are not restricted to 

inquiline species. The damselfishes Eupomacentrus 

dorsopunicans (Poey) and E. diencaeus Jordan and 

Rutter both occur at Glover's Reef but are ecologically well separated. The first of this pair 

occurs only in shallow, wave-swept areas on the 

windward reef, whereas the second occurs only in 

deeper water or within the lagoon where wave 

action is absent. Eupomacentrus diencaeus also occurs 

on the reef crest at the leeward side of Glover's 

Reef, another area where wave action is absent. 

Nowhere at Glover's Reef have the two species 

been found occurring in the same area (Greenfield and Woods, 1974:19). 

The basslets in the genus Gramma provide another striking example of habitat partitioning at 

the dropoff (fore-reef slope) of the outer fore reef 

at Glover's Reef. Gramma loreto occurs in shallow 

waters and over the edge of the dropoff to a depth 

of 24.5 m. A second species, G. melacara, occurs 

from a depth of 21 m on the vertical face of the 

dropoff down to 110 m. Thus only a 3.5 m zone 

of overlap occurs at the dropoff at Glover's Reef. 

Three ichthyocide collections from various depths 

A. spinosa 

A. COPEPODA 

B. OSTRACODA 

C. TANAIDACEA 

D. ISOPODA 

E. AMPHIPODA 

F ANNELIDA 

G. DECAPODA 

H. GASTROPODA 

I. CIRRIPEDIA 

J . INSECTA 

E F Gr ^ • _ J 

_J

 v Y//A IH7777I 

M = 10% 

% Frequency of Occurrence 

FIGURE 220.—Comparison of percent composition of food by taxonomic group in number, 

volume, and frequency of occurrence for Acanthemblemaria spinosa and Acanthemblemaria greenfieldi. 
506 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

demonstrate the partitioning of habitat by depth: 

(a) at 18 m, 64 G. loreto, 0 G. melacara; (b) 21-24 

m, 18 G loreto, 91 G. melacara; and (c) 34 m, 0 G. 

loreto, 32 G. melacara. Colin (1974:37) has observed 

that the bathymetric distribution of G. melacara 

varies geographically depending upon local conditions, particularly the presence of vertical reef 

faces. Thus, the sharp separation of G. melacara 

and G. loreto is in dropoff areas where strong depth 

changes occur along relatively short horizontal 

distances, such as at Glover's Reef. Another species of Gramma, G. linki, observed at Belize only at 

the lower depth range of G. melacara (Colin, 1974: 

34), provides an additional example of habitat 

partitioning by depth. Starck and Colin (1978: 

146) stated that in localities where G melacara is 

absent, G. linki occurs in considerably shallower 

water. 

Since the publication of Sale's work in 1977, 

several papers have appeared relative to the question of the importance of stochastic larval recruitment of coral-reef fishes. Molles (1978), working 

in the Gulf of California, found the fish faunas of 

his small replicate reefs showed a high degree of 

similarity and a correlation between species diversity and habitat complexity. Gladfelter and 

Gladfelter (1978) found both a high degree of 

similarity between fish assemblages and also correlations between assemblages and structural features of a series of relatively large coral patch 

reefs in the tropical western Atlantic. Gladfelter 

et al. (1980:1167), recognizing the problems associated with studying colonization in very small 

artificial structures and the fact that various studies had been conducted by different workers in 

different oceans, attempted to solve these problems by determining the degree of similarity between fish assemblages of groups of replicate 

patch reefs in the tropical western Atlantic (Virgin Islands) and in the tropical central Pacific 

(Marshall Islands). They concluded, ".. . the 

high diversity of coral reef fish communities is 

permitted by a variety of factors other than random recruitment of larvae alone; of these, habitat 

structure is the most evident." 

Anderson et al. (1981:494) presented data on 

the distribution of butterfly fishes of the northern 

Great Barrier Reefs of Australia in relation to 

various theories of coral reef fish community 

structure. They found there were conspicuous 

niche differences between locally coexisting species, geographical replacement occurred between 

species from the same niche, and similarities existed between species present at many different 

sites of the same habitat type. Based on these 

data they concluded, "Our data do not support 

the need for alternative hypotheses centered on 

larval habitat preferences and stochastic recruitment." 

Also, additional field work (since the 1977-

1978 Carrie Bow Cay observations) has 

strengthened the findings reported in Greenfield 

and Johnson (1981) relative to habitat specificity 

of blennioid fishes. 

The examples just cited emphasize the importance of viewing habitat associations of fish species on a sufficiently small scale. The co-occurrence of species from single collections recorded 

in the literature may often be misleading in suggesting that habitat partitioning is not a reality. 

Considering that detailed studies have been conducted for only a few species in the western 

Atlantic Ocean and in the central Pacific, it is 

premature to conclude that habitat partitioning 

does not occur among coral-reef fishes. General 

studies of the blennioid fishes of Belize and Honduras have shown significant differences in species 

distribution across various major habitat types 

(Greenfield and Johnson, 1981). Detailed studies 

of these small species should yield valuable information on the importance of habitat partitioning. 

Many examples of the absence of habitat partitioning provided by Sale (1977) concerned larger, 

often herbivorous species; however, the high species diversity found in coral-reef fish communities 

is largely due to the small sedentary species, such 

as the blennies and gobies. To date, we have 

collected over 300 species of reef-associated fishes 

in Belize, of which 84 are blennies and gobies. 

Thus, the lottery or chaos hypothesis proposed by 

Sale may apply to some of the larger species he 

studied, but it does not appear to apply broadly 

to all coral-reef fish species in the western Atlantic. 
NUMBER 12 507 

Literature Cited 

Anderson, G.R.V., A. H. Ehrlich, P. R. Ehrlich, J. D. 

Roughgarden, B. C. Russell, and F. H. Talbot 

1981. The Community Structure of Coral Reef Fishes. 

American Naturalist, 117:476-495. 

Colin, P. L. 

1974. Observations and Collection of Deep-Reef Fishes 

off the Coasts of Jamaica and British Honduras 

(Belize). Marine Biology, 24:29-38. 

Gladfelter, W. B., and E. H. Gladfelter 

1978. Fish Community Structure as a Function of Habitat Structure on West Indian Patch Reefs. Revista 

de Biologia Tropical, 26 (supplement l):65-84. 

Gladfelter, W. B., J. C. Ogden, and E. H. Gladfelter 

1980. Similarity and Diversity among Coral Reef Fish 

Communities: A Comparison between Tropical 

Western Atlantic (Virgin Islands) and Tropical 

Central Pacific (Marshall Islands) Patch Reefs. 

Ecology, 61:1156-1168. 

Greenfield, D. W. 

1972. Notes on the Biology of the Arrow Blenny, Lucayablennius zingaro (Bdlhke) from British Honduras. 

Copeia, 1972:590-592. 

1975. Emblemariopsis pricei, a New Species of Chaenopsid 

Blenny from Belize. Copeia, 1975:713-715. 

Greenfield, D. W., and R. K. Johnson 

1981. The Blennioid Fishes from Belize and Honduras, 

Central America, with Comments on their Systematics, Ecology, and Distribution (Blenniidae, 

Chaenopsidae, Labrisomidae, Tripterygiidae). 

Fieldiana, Zoology, new series, 8:1-106. 

Greenfield, D. W., and L. P. Woods 

1974. Eupomacentrus diencaeus Jordan and Rutter a Valid 

Species of Damselfish from the Western Tropical 

Atlantic. Fieldiana, Zoology, 65:9-20. 

Hellawell, J. M., and R. Abel 

i971. A Rapid Volumetric Method for the Analysis of 

the Food of Fishes. Journal of Fish Biology, 3:29-37. 

Horn, J. S. 

1966. Measurement of "Overlap" in Comparative Ecological Studies. American Naturalist, 100:419-424. 

Molles, M. C , Jr. 

1978. Fish Species Diversity on Model and Artificial and 

Natural Patch Reefs: Experimental Insular Biogeography. Ecological Monographs, 48:289-305. 

Pinkas, L. 

1971. In L. Pinkas, M. S. Oliphant, and I.L.K. Iverson, 

Food Habits of Albacore, Bluefin Tuna, and Bonito in California Waters. California Division of Fish 

and Game, Fish Bulletin, 152:1-105. 

Ross, S. T. 

1974. Resource Partitioning in Searobins (Pisces, Triglidae) on the West Florida Shelf. Ph.D. Dissertation, University of South Florida, Tampa. 

1977. Patterns of Resource Partitioning in Searobins 

(Pisces: Triglidae). Copeia, 1977:561-571. 

Sale, P. F. 

1977. Maintenance of High Diversity in Coral Reef Fish 

Communities. American Naturalist, 111:337-359. 

Smith-Vaniz, William F., and Francisco J. Palacio 

1974. Atlantic Fishes of the Genus Acanthemblemaria, with 

Description of Three New Species and Comments 

on Pacific Species (Clinidae: Chaenopsinae). Proceedings of the Academy of Natural Sciences of Philadelphia, 125:197-224. 

Starck, W. A., II, and P. L. Colin 

1978. Gramma linki: A New Species of Grammid Fish 

from the Tropical Western Atlantic. Bulletin of 

Marine Science, 28:146-152. 

Stephens, John S., Jr., Robert K. Johnson, Gerald S. Key, 

and John E. McCosker 

1970. The Comparative Ecology of Three Sympatric 

Species of California Blennies of the Genus Hypsoblennius Gill (Teleostomi, Blenniidae). Ecological 

Monographs, 40:213-233. 
Reconnaissance Study of the 

Geomorphology and Benthic Communities 

of the Outer Barrier Reef Platform, Belize 

Randolph B. Burke 

ABSTRACT 

The first general survey of the entire 257 km 

outer platform of the Belizean barrier reef has 

disclosed three provinces having distinct community distribution and geomorphic characteristics: (1) the northern province, which contains 46 

km of shallow-water reefs; (2) the central province, with 91 km of shallow-water reefs; and (3) 

the southern province, which includes only 10 km 

of shallow-water reefs. Central province reefs are 

the best developed. They are wide and continuous 

shallow reefs (<—0.2 m MSL) with flourishing 

communities of branching corals. The fore reef is 

characterized by high relief and a pronounced 

ridge, capped with Acropora cervicornis at depths of 

less than 20 m, extending along the shelf edge. 

The reef in the northern and southern provinces 

is discontinuous except along Ambergris Cay in 

the north. Wide passages of uniform depth occur 

in the northern province, in contrast to the narrow, deep channels within passes typical of the 

southern province. In both the north and the 

south, the reef occurs adjacent to islands and 

generally is more distant from the shelf edge than 

is the reef along the central province. In the south, 

however, some sections of the reef occur as reentrants and are closer to the shelf edge. 

On the seaward side, framework-building communities have the same composition along the 

barrier reef, but relative species abundance and 

density vary significantly. The greatest abundance occurs in the central province in association 

with the coral ridge and high-relief spur and 

groove structures. The distribution of these structures coincides with the greatest extent of sea 

Randolph B. Burke, North Dakota Geological Survey, University 

Station, Grand Forks, N D. 58202. 

level reef development. Their distribution also 

corresponds to the region of the barrier reef most 

protected from open ocean waves. The shelter 

afforded by seaward platforms has favored 

growth of rapidly accumulating communities of 

branching corals, and thereby has substantially 

contributed to the central province's development 

of the most extensive and luxuriant shallow portions of the barrier reef. 

Introduction 

Although the longest continuous barrier reef in 

the West Indies occurs along the continental shelf 

edge of Belize (Figures 221, 222; Smith, 1971:29) 

little work has been done on the seaward part of 

the reef. Previous studies have focused on geological and biological zonations in the lagoon (Wantland and Pusey, 1975). In addition, during the 

Cambridge Expedition to British Honduras 

(Thorpe and Stoddart, 1962), Thorpe and Bregazzi (1960) made preliminary observations on 

corals at Rendezvous Cay. Using their results for 

baseline data, Stoddart (1962, 1963, 1969, 1974) 

studied hurricane damage to the reef and subsequent recovery. York (1971) compiled a photolog of common Belizean corals. Recent geologic 

studies on the deep seaward margin (James and 

Ginsburg, 1978) and studies of shallow-water submarine cementation (James et al., 1976) have 

described parts of the windward section of the 

barrier reef. 

This paper presents a comprehensive survey of 

the Belizean barrier reef based largely on aerial 

reconnaissance and underwater observations. It 

509 
510 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

FIGURE 221.—Belizean barrier reef looking south towards Tobacco Cay entrance, an example 

of the extensive shallow-water reef development characteristic of the central province. 

thus places the Carrie Bow Cay transect data (see 

Riitzler and Macintyre, herein: 9) into regional 

perspective. The study considers shelf bathymetry, regional patterns of wave energy, and the 

distribution and development of reef communities and their architecture. 

ACKNOWLEDGMENTS.—Field work was carried 

out with W. Adey, P. Adey, C. Shipp, and pilot 

T. Metts. A grant from the Smithsonian Institution Fluid Research Fund awarded to K. Riitzler 

provided financial assistance. J. Suhayda, Institute of Coastal Studies, Louisiana State University, provided initial calculations of wave energy 

and a wave diagram that was modified with data 

from Belize. An early version of the manuscript 

was reviewed by L. C. Gerhard, H. J. Humm, I. 

G. Macintyre, and F. T. Manheim; I am especially grateful for the critical comments of W. H. 

Adey, M. S. Gould, and S. B. Upchurch. The 

aerial photograph was provided by J. A. Miller, 

Union Oil Company. I. Jewett drafted the final 

version of the figures. 

Methods 

Low altitude aerial photography was used to 

map major geomorphic features and reef communities along the margin of the Belizean barrier 

reef platform from Ambergris Cay in the north 

to Zapotilla Cays in the south. Surveys by 

SCUBA diving were also made along seven transects perpendicular to the reef to record the bottom profile and zonation of substrates and organisms (Figure 222). 

Photographs were taken through the open window of a high-wing plane at an altitude of 150 m 

and air speed of 160-170 km/h, using 35 mm and 

60 mm cameras equipped with polarizing filters. 

Pictures were taken between 0900 h and 1500 h 

when glare was at a minimum. Two photographic 
1180

00' 

89°O0 88°30 

FIGURE 222.—Map of the coastal areas of Belize showing distribution of sea level reefs, as 

determined by this study, and location of transects (a-g). 
512 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

surveys were flown, the initial one concerned 

primarily with establishing transect sites. After 

ground truth transects were completed, the second photography flight documented their location. Calm water after a norther improved visibility through the water, and thus provided reliable resolution to depths of 20 m for mapping the 

geomorphic features and the principal bottom 

communities. 

The underwater transects, measured with 

depth gauges and from charts, extended from the 

lagoon, across the reef crest, and down the shelf 

edge to depths of 40 m. Dominant biota and 

visual estimates of percentage of cover were recorded on mylar sheets attached to a PVC wrist 

slate. In addition, characteristic zones of the transects were photographed. Wave energy was calculated from mean annual wind direction and 

wave heights resolved from SSMO (Snyoptic 

Summary of Meteorological Observations) data 

(United States Naval Weather Service Command, 1974). See Burke (1979) for details on 

wave energy calculations. 

Composition and Distribution 

of Major Reef Structures 

PHYSIOGRAPHIC PROVINCES 

The barrier reef platform (<5.5 m depth) seaward of Belize is 257 km long, of which 147 km 

consists of shallow-water reefs (<0.2 m depth; 

Figure 222). The remainder of the shelf-edge 

platform is cut by channels or is composed of 

carbonate shoals, fleshy-algae-covered pavements, or deeper reef development. The reef complex can be divided into three provinces on the 

basis of community distribution and geomorphic 

characteristics: the northern, central, and southern provinces. The biological and topographic 

characteristics of these provinces are defined as 

follows. 

NORTHERN PROVINCE.—Extending approximately 78 km southward from Reef Point to the 

northern tip of Gallows Point Reef, this province 

includes approximately 46 km of reef and the 

muddy, shallow Chetumal Bay-lagoon complex 

with average water depth less than 6 m. The 

barrier platform in this province is distinguished 

by a line of mangrove cays set well inside the 

shelf edge. Passes between the reef and these 

islands are wide and shallow (<5 m deep). Except 

for areas along Ambergris Cay and Cay Corker 

where the reef crest is formed by Millepora-coxaV 

line mounds or coralline-algae-encrusted corals, 

the area of the reef crest and inner fore reef in the 

northern province has few living corals and is 

rounded and gently sloping. Coral is abundant 

behind the reef crest along Ambergris Cay (Plate 

1: top left). The shoals between this cay and 

Gallows Point are primarily a pavement covered 

by rubble, sand, and only sparse corals. The biota 

along the shelf-edge break are luxuriant except 

where numerous wide sand channels cross the 

shelf and are probably similar in composition to 

biota along the shelf break at Gallows Point Reef 

(see Figure 223a). 

CENTRAL PROVINCE.—Extending south approximately 120 km from Gallows Point Reef to Gladden Spit, this province has more than 91 km of 

shallow-water reef (Figure 221; Plate 1: top right, 

center left); and includes the central and southern 

lagoons, which have depths of less than 27 m. 

Numerous mangrove cays occur near the center 

and landward side of the barrier platform, and 

along the boundary of the southern lagoon. 

Unique to the central province are a few sand 

cays built on the inner edge of the shallow-water 

reef, and three carbonate platforms 8 to 40 km 

seaward: Turneffe Islands, Lighthouse Reef, and 

Glover's Reef (Figure 222). In contrast to the 

others, the central province of the barrier platform supports long and wide sections of uninterrupted reef, which are cut by only two major 

channels and scattered narrow passes. The central 

province is characterized by three distinct structural features: high-relief spur and groove zones, 

double high spur and groove zones, and a shelfedge coral ridge (Figure 224). 

The high-relief spur and groove system occurs 

over 52 km of the reef. The relief of spurs ranges 

from 2 m to more than 10 m; spurs are dominated 

by coral communities in less turbulent water and 

by Millepora-coraWine algae in higher wave en-
NUMBER 12 513 

ergy zones. A double high-relief spur and groove 

system occurs over 16 km of the reef an d is 

separated from the reef crest by a channel tha t 

runs parallel to the reef trend an d contains boulder rubble. Th e communitie s on these spurs are 

rich in coralline algae and Millepora complanata 

Lamarck. 

Coral ridges occur along 48 km of the outer 

reef at the shelf edge and are capped by Acropora 

cervicornis (Lamarck) at depths less than 20 m. At 

depths greater tha n 20 m, the ridge is less continuous and less wide; A. cervicornis becomes less 

abundant and platy corals, particularly Montastrea 

annularis (Ellis an d Solander), prevail. Ridges having high relief occur only leeward of the seaward 

platforms. 

SOUTHERN PROVINCE.—Extendin g approximately 59 km from Gladden Spit to Zapotilla 

Cays, this province has about 10 km of shallowwater reef an d a deep lagoon (>27 m deep), 

which is known as Victoria Channel. Th e barrier 

platform supports a few small cays in this province, most of which are well inside the shelf edge. 

Passes are wide and generally have narrow, deep 

channels (6-10 m deep). In passes, the platform 

crest is broad an d rounded and supports a dense 

crop of algal macrophytes. 

In the southern province, shallow-water reefs 

occur only as fringe around islands and reentrants 

to the lagoon. Th e reef crest is composed mainly 

of weakly coalesced Millepora-coraWine mounds 

and rubble. Coral density is greatest in the back 

reef and along the shelf edge. 

TRANSECTS ACROSS THE BARRIER REE F 

Seven underwate r transects are described from 

north to south: five in the central province, and 

two in the southern (Figures 222, 223). Descriptions proceed from the lagoon to the shelf edge. 

Gallows Point Reef is described in detail as a 

reference for the remainde r of the transects. Constituents change little between transects, but population densities and areal distributions vary 

markedly. 

GALLOWS POINT REEF , CENTRAL PROVINCE (Figure 223a).—Almost due east of Belize City, Gallows Point Reef is the northernmost transect. A 

77ia/a.5\na-dominated lagoon grades into reef-derived rubble that surrounds patch reefs which 

fringe the moderately gentle back-reef slope. 

Commo n constituents of the patch reefs are the 

corals Siderastrea radians (Pallas), S. siderea (Ellis 

and Solander), Porites astreoides (Lamarck), Favia 

fragum (Esper), Agaricia sp., Diploria clivosa (Ellis 

and Solander), D. strigosa (Dana), an d the hydrozoans Millepora alcicornis Linnaeus and M. complanata. Th e rubbl e is interrupted locally by pavement covered with algal turf and by sand patches. 

Hydrolithon boergesenii (Foslie) Foslie, and a few 

species of Neogoniolithon are common encrusting 

corallines on back-reef rubble. O n the turf-covered pavement, dominated by microscopic algae 

(<1 cm)—primarily Phaeophyt a and Rhodophyt a that stabilize a sediment veneer—there is 

an increasing abundanc e of Porites astreoides and 

coralline algal mounds toward the reef crest. 

T h e somewhat elongate, steep-sided (0.5 m 

high) mounds of the reef crest, dominated by 

Millepora complanata and crustose corallines, support diverse biota, including the zoanthid Palythoa 

sp., Porites astreoides, Favia fragum, Siderastrea siderea, 

Caulerpa sp., Halimeda sp., and a few Diploria clivosa, D. strigosa, Agaricia sp., and Acropora palmata 

(Lamarck). A patchwork of pavement with some 

rubble, coated by both corallines and algal turfs, 

surrounds the mounds. Coralline algae encrusting 

the rubble include Porolithon pachydermum (Webervan Bosse and Foslie in Foslie) Foslie and Neogoniolithon sp. 

Scattered thickets oi Acropora palmata commonly 

interfinger with the reef crest and the spur and 

groove zone of the fore reef. Rubbl e is a major 

component in the sand-floored grooves and on 

the turf-covered pavement. Like the reef crest, the 

surface of the spurs supports a variety of corals, 

predominantly A. palmata, Montastrea annularis, 

Agaricia agaricites (Linnaeus), and Porites astreoides. 

Seaward, on the low-relief spur and groove pavement, rubble is commonly covered by the encrusting sponge Cliona sp. an d other encrusting organisms such as the coralline algae Paragoniolithon 

sp., Neogoniolithon sp., and some Tenarea prototype 

(Foslie) Adey. Along with the pavement, the rub-
514 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Gallows Point Reef 

Lagoon 

Sand 

a 

Sea Grass 

Zone 

Back Reef 

Rubble 8 Pavement 

Zone 

Reef 

Crest 

MilleporaCorallinacea 

Zone 

Inner Fore Reef 

Moderate 

Spur 8 Groove 

Zone 

Low Spur 8 Groove 

Zone 

High 

Spur 8 Groove 

Zone 

1 0-

2 0 -

3 0 -

4 0 -

1 

meters 

—Mi . com., Ac. pal., Si. sid. 

Di. str. 

Ac. cer. 

Po. ast 

Paly 

Caul. 

Hali. 

Ag. aga., Mo. ann 

Por, Octo. 

Diet. 

Mo. cav. 

Ma. mir. 

Colp.... 

Lobo. 

Myce.... 

I 

200 4 0 0 

Rendezvous Cay 

Lagoon Back Reef Reef 

Crest Inner Fore Reef 

Sand 

8 

Sea Grass 

Zone 

Coral 

8 

Rubble 

Zone 

Rubble 

8 

Pavement 

Zone 

MilleporaCora. 

Zone 

Moderate 

Spur 

8 Groove 

Zone 

Low Spur 8 Groove 

Zone 

High 

Spur 

8 Groove 

Zone 

Inner 

Reef 

Slope 

Ma. mir, Ac. cer., 

Mi. ale, Hali. 

Sarg., Lobo., Colp. 

200 

I 

4 0 0 
NUMBER 12 515 

Outer Fore Reef 

Fore Reef Slope 

Mean Sea Level 

Di. sto. 

600 

&L 

^ \ 

X? 

p 

r\ 

^ 

7 ^ ^ 

V 

* 

Sand 

Rubble 

Mounds or Spurs 

Coral Heads 

Platy Corals 

Fleshy Algae 

Seagrass 

Sponges 

Millepora complanata 

Acropora cervicornis 

Acropora palmata 

Octocorals 

Gorgonia sp. 

Alternate Contour 

Trace or Absent 

Common 

FIGURE 223.—Idealized sketches of barrier reef transects 

(located in Figure 222), showing reef zonation and major 

components of surface coverage: a, Gallows Point Reef; b, 

Rendezvous Cay (configuration of coral ridges determined 

from hydrographic charts and aerial rconnaissance); c, Colson Cay (the inner coral ridge is a branch of the outer ridge, 

not a parallel structure); d, Carrie Bow Cay (after Riitzler 

and Macintyre, herein, Table 1, Figures 4, 5); e, South Cut; 

/, Gladden Spit (shelf edge not observed, dashed profile 

taken from hydrographic charts); g, Queen Cay. (Coverage 

indicated by location of abbreviation and solid or dotted 

lines; arrowheads mark extended distribution; abbreviations: Ac. cer. = Acropora cervicornis, Ac. pal. = A. palmata, Ag. 

aga. = Agaricia agaricites, Caul. = Caulerpa sp., Colp. = 

Colpophyllia sp., Cora. = Corallinacea, Di. sto. = Dichocoenia 

stokesi, Diet. = Dictyota sp., Dip. cl. = Diploria clivosa, Dip. la. 

= D. labyrinthiformis, Dip. st. = D. strigosa, Dipl. = Diploria sp. 

Fa. fra. = Faviafragrum, Gorg. = Gorgonia sp., Hali. = Halimeda 

sp., Lobo. = Lobophora sp., Ma. mir. = Madracis mirabilis, Me. 

rnea. = Meandrina meandrites, Mi. ale. = Millepora alcicornis, 

Mi. com. = M. complanata, Mille. = M. sp., Mo. ann. = 

Montastrea annularis, Mo. cav. = M. cavernosa, Myce. = Mycetophyllia sp., Octo. = Octocorallia, Padi. = Padina sp., Paly. 

= Palythoa sp., Para. = Paragoniolithon sp., Por. = Porifera, Po. 

ast. = Porites astreoides, Po. por. = P. porites, Sarg. = Sargassum 

sp., Scol. = Scolymia sp., Si. rad. = Siderastrea radians, Si. sid. 

= S. siderea, Sid. = Siderastrea sp., Styp. = Stypodium sp., Turb. 

= Turbinaria sp., Thai. = Thalassia sp., Udot. = Udotea sp.) 

Outer Fore Reef 

Sand Trough Inner Coral Ridge Sand Trough Outer Coral Ridge 

Fore Reef 

Slope 

Mean Sea Level 

Unexplored 

600 800 1000 
516 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Colson Cay 

Lagoon 

Sand 

a 

Sea Grass 

Zone 

Back Reef 

Rubble 

a 

Coral 

Zone 

Rubble 8 Pavement 

Zone 

Reef 

Crest 

MilleporaCora. 

Zone 

Inner Fore Reef 

Moderate 

Spur 8 Groove 

Zone 

Low Spur 8 Groove 

10-

2 0 -

3 0 -

40 -

1 

meters 

Mi. com 

Po. ast 

Ac. pal.... 

.Si. sid 

Ag. aga 

.Dip. cl. 

Dip. st 

.Fa. fra. 

Paly 

Turb 

Sarg Lobo 

Gorg., Hali., Para. 

Mo. ann. ..Ac. cer._ 

Di. sto. 

Octo 

I 

200 400 

Carrie Bow Cay 

Lagoon Back Reef Reef 

Crest 

Sand 8 

Sea 

Grass 

Zone 

Sand 8 

Rubble 

Zone 

Patch Reef 

Zone 

Rubble 

a 

Pavement 

Zone 

MilleporaCora. 

Zone 

High Spur a 

10-

2 0 -

3 0 -

4 0

 J 

meters 

•-Thai. 

Diet. 

Dip. la... 

Mo. ann. 

-@"-v-s3pac^ 

Si. sid. _ 

.Ac. cer. 

Po. ast.. 

Ac. pal.. 

Mi. com. 

Ag. aga. 

Paly 

Hali 

Po. por. 

200 
NUMBER 12 517 

Outer Fore Reef 

Zone 

Inner 

Reef 

Slope 

Sand Trough 

a 

Inner Coral Ridge 

Outer 

Coral 

Ridge 

Fore Reef 

Slope 

Mean Sea Level 

Por 

Po. por., Mo. cav.,..Colp 

Mi. ale 

600 800 

Inner Fore Reef 

Groove Zone Low Spur 8 Groove Zone 

Outer Fore Reef 

Inner 

Reef 

Slope 

Sand 

Trough 

Outer 

Coral 

Ridge 

Fore Reef 

Slope 

Mean Sea Level 

Octo, Por. 

Mo. cav 

Myce. 

Ma. mir. 

4 0 0 600 
518 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

South Cut 

Lagoon 

Sand 8 Sea Grass Zone 

Back Reef 

Rubble 

a 

Coral 

Zone 

Rubble 

a 

Pavement 

Zone 

Reef 

Crest 

MilleporaCorallinacea 

Zone 

Low 

Spur 8 Groove 

Zone 

10-

2 0 -

3 0 -

40 -

1 

meters 

Udot. 

Sid 

Po. por. 

Dipl 

Por. ast.. 

Ac. pal._ 

Mi. com.. 

Ag. aga.. 

Mo. ann. 

Diet. 

Paly._ 

Fa. fra. 

Hali 

Turb. 

~~r~ 

400 

200 

Gladden 

Lagoon 

Sand 

8 

Sea Grass 

Zone 

Spit 

Back Reef 

Rubble 8 Coral Zone 

Reef 

Crest 

MilleporaCorallinacea 

Zone 

Moderate 

Spur 8 Groove 

Zone 

io2 0 -

3 0 " 

4 0

 J 

meters 

370 m 

M/ M/.-^ 

Mi. com.. 

Ac. palm. 

Fa. fra._ 

Mo. ann.. 

Dip. cl 

Dip. st 

.Po. ast., Si. sid., Si. rad. 

Paly. 

Ag. aga. 

Sarg.,_ 

200 400 
NUMBER 12 519 

Inner Fore Reef 

Pavement 

Zone 

Mod. 

Spur a 

Groove 

Zone 

Pavement 

Zone 

Outer Fore Reef 

High 

Spur 8 Groove 

Zone 

Fore Reef 

Slope 

Octo., Por. 

Styp 

Sarg., Lobo.. 

Mean Sea Level 

Mo. cav. 

Ac. cer, Ma. asp., Me mea 

Myce., Colp 

600 800 

Inner Fore Reef 

Pavement 8 Sand Channel Zone 

Outer Fore Reef 

Fore Reef Slope 

Mean Sea Level 

W_<2 

.Diet., Lobo., Styp.. 

Octo 

Por., Mo. cav., 

Di. sto., Colp.. 

1800 2000 2200 
520 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

QUEEN CAY 

Lagoon 

Sand 

a 

Sea Grass 

Zone 

Back Reef 

Rubble 

a 

Coral 

Zone 

Rubble 8 Pavement Zone 

Reef 

Crest 

MilleCora. 

Zone 

Moderate 

Spur 8 Groove 

Zone 

10-

2 0 -

3 0 -

Thal. 

Udot. 

Mi. com . 

Po. ast... 

Sid 

Ag. aga. 

Dip. cl. 

Dip. St.. 

40

 J 

meters 

Hali. 

Ac. pal. 

Paly. 

Sarg 

Turb. 

Mo. ann. 

Gorg 

Por, Octo. 

Diet 

200 400 

ble provides the primary substrate for the ubiquitous alga Dictyota sp., the common Gorgonia sp., 

other octocorals, sponges, and diverse but scattered coral heads, mostly Diploria strigosa and 

Montastrea annularis. 

Nea r the shelf edge, a weakly developed spur 

and groove system with about 1.5 m relief marks 

the transition to the fore-reef slope. Spurs along 

the seaward face are characterized by a rather 

dense (approaching 50 percent cover) and diverse 

coral assemblage in which Montastrea cavernosa 

(Linnaeus), M. annularis, and Agaricia spp. are 

most abundant. Corals common to deeper water, 

such as Mussa angulosa (Pallas), Mycetophyllia sp., 

Madracis mirabilis (Duchassaing and Michelotti), 

and Colpophyllia sp. are also conspicuous, together 

with sparse populations of Acropora cervicornis, Diploria labyrinthiformis (Linnaeus), D. strigosa, and 

Siderastrea siderea. Turf and the alga Lobophora sp. 

are most abundant, but large vase sponges and 

octocorals are also common. 

At the base of the spur faces a pavement slope 

is coated with a thin veneer of fine sand and 

abundan t Lobophora sp., but as the slope steepens, 

corals again dominat e and in places form local 

overhangs along the shelf-edge wall. 

RENDEZVOUS CAY, CENTRAL PROVINCE (Figure 

22?>b).—This transect lies 2.2 km north of Rendezvous Cay on the outer barrier reef. Th e shallow reef is narrow here and the back-reef slope is 

steep, rising abruptly 10 m from a sand bottom 

having seagrass and algae {Udotea sp., Penicillus 

sp., and Padina sp.). A distinct rubbl e zone lies 

immediately below the patchy but dense colonies 

of Acropora palmata and mounds of Millepora complanata, Montastrea annularis, and Siderastrea siderea 

tha t span the sharp slope of the back reef. Turfcovered rubble dominates the back-reef flat, 

which also has a sparse coral fauna that includes 

Porites astreoides, Agaricia agaricites, Diploria sp., and 

P. porites (Pallas). Palythoa sp. coats the rubble 

crest where Millepora-coraWine mounds constitute 

less than 40 percent of the crest's irregular trend. 

Rubbl e continues to predominat e through the 

moderately high-relief (1.3 m) spur zone in which 

Montastrea annularis is the principal coral. Less 
NUMBER 12 521 

Inner Fore Reef 

Low Spur a Groove Zone 

Outer 

Fore Reef 

Fore Reef 

Slope 

Mean Sea Level 

Styp. 

Dip. la. 

Mo. cav.. 

Ac. cer. Lobo. 

600 800 

abundan t species include Isophyllastrea rigida 

(Dana), Dendrogyra cylindrus Ehrenberg, and Acropora palmata. Turf, Dictyota sp., and fine sediment 

cover muc h of the low-relief spur and groove 

pavement as the amoun t of rubble decreases seaward. Th e rubbl e consists primarily of branches 

oi Acropora cervicornis. Pseudopterogorgia sp., Gorgonia 

sp., other octocorals and sponges are the most 

conspicuous components of the pavement, which 

also includes the corals Dichocoenia stokesi (Milne 

Edwards and Haime ) and Dendrogyra cylindrus. 

Massive spurs of u p to 4.8 m relief mark the 

slope break, which drops to 44 m into a sand 

trough. Uppe r spurs are characterized by scattered Acropora cervicornis, Madracis sp., Porites porites, 

Montastrea annularis, and M. cavernosa and aboun d 

with octocorals an d sponges. Coral diversity is 

highest on the seaward face, which also supports 

P. astreoides, Mycetophyllia spp., Colpophyllia sp., 

Isophyllia sp., Diploria labyrinthiformis, Eusmilia fastigiata (Pallas), an d encrusting Millepora alcicornis 

on octocorals. Lobophora sp. is mor e commo n on 

horizontal surfaces an d Halimeda spp. on spur 

margins. Depths between 20 and 35 m are characterized by a sandy slope having patches of 

raised pavement that support the alga Sargassum 

sp., octocorals, and large vase sponges. 

Aerial reconnaissance revealed two large ridge 

systems bordering the shelf edge which were not 

seen during the dive because of turbid water. Th e 

ridges could be comparabl e to the Acropora cervicornis'-capped coral ridges at Colson and Carrie 

Bow cays (see below). These ridges, however, are 

more segmented and, according to hydrographic 

charts, have u p to 35 m of relief. 

COLSON CAY , CENTRAL PROVINCE (Figure 

222>c).—Lagoon and back-reef areas are similar to 

Gallows Point although sand patches are mor e 

common within the rubble of the back reef. A 

discontinuous series of Millepora-coraWine mounds 

mark the reef crest, which is composed of over 60 

percent rubble. Rubbl e coverage increases to over 

70 percent in an irregular zone that enters the 

spur and groove system. Algal biomass on the 

crest approaches 450 g/m2 and is composed 

mainly of Sargassum sp. and Turbinaria turbinata 
522 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

(Linnaeus) Kuntze . Lobophora sp. and Halimeda sp. 

become the primary algal macrophytes on the 

turf-dominated rubble zone that separates the 

reef crest from the adjacent spur and groove zone. 

Living corals in this spur and groove zone cover 

less tha n 10 percent of the surface. Rubble , prevalent in grooves, is covered mainly by turf, the 

coralline alga Paragoniolithon sp., and Lobophora sp. 

T h e most obvious coral is Acropora cervicornis with 

some Montastrea annularis, Agaricia agaricites, Siderastrea siderea, and only a few patches of Acropora 

palmata. These spurs of moderate relief grade 

seaward into a low-relief spur and groove system 

covered primarily by fine sediment stabilized by 

turf an d a scattering of Sargassum sp. Th e few 

corals present are scattered widely. Coral diversity 

and numbers increase on steeper spur faces at the 

slope break. At the base of the spur face, the 

bottom is about 50 percent low-relief spurs an d 

the remainde r is sand. Whip-like octocorals are 

abundant, as are several types of large sponges. 

An anastomosing coral ridge, capped by Acropora cervicornis and separated from the inner fore 

reef by a sand trough, characterizes the outer fore 

reef and shelf edge. This ridge, which does not 

appea r on hydrographic charts, has a maximum 

relief of 16 m. Lobophora sp. is common in the 

areas of pavement, which, along with dead 

branches of A. cervicornis, serve as attachment sites 

for the abundan t octocorals, a variety of vase 

sponges, and less conspicuous corals such as Madracis asperula Milne Edwards and Haime , Colpophyllia sp., Montastrea cavernosa, Mycetophyllia sp., 

Isophyllia sinuosa (Ellis and Solander), and Scolymia 

sp. 

CARRIE Bow CAY, CENTRAL PROVINCE (Figure 

223</).—This is the IMSW E Program reference 

transect, which is described in detail in Riitzler 

and Macintyr e (herein: 9). 

SOUTH CUT , CENTRAL PROVINCE (Figure 

223f).—The transect traverses the barrier reef on 

the south side of the South Cut reentrant (Figure 

222). Rubbl e dominates the immediat e back reef, 

which grades into a crest of sparse mounds of 

Millepora and coralline algae. Th e crest is composed of about 50 percent rubble. Fine sediment 

stabilized by turf covers most hard surfaces, with 

Neogoniolithon sp., some Porolithon pachydermum, and 

Millepora complanata present on higher areas. 

Fleshy algal macrophyt e coverage is relatively 

low (<100 g/m2) and consists mainly of Turbinaria 

sp. Palythoa sp. forms mats over less tha n 3 

percent of the area. A series of low-relief (<1 m) 

spurs surrounded by rubble extends below the 

crest. Biota on the spurs are similar to tha t found 

on the crest, but some Agaricia agaricites, Acropora 

palmata, and Halimeda spp. also occur. 

T h e numbe r an d size of the spurs an d mounds 

decrease seaward of the crest. Th e mounds 

broaden into a pavement covered primarily by 

turf, rubble, and sand. Gorgonia sp., other octocorals, and the alga Stypopodium sp. are the most 

conspicuous components, but are quantitatively 

unimportant in terms of surface coverage. Another set of mounds occurs at a depth of about 6 

m and in places they coalesce to form a moderately high-relief (~4.5 m) spur and groove system. 

Acropora palmata, Agaricia agaricites, Millepora complanata, and an encrusting Millepora sp. are the 

most obvious components, although turf with 

sediment, Sargassum sp., and Lobophora sp. are 

actually more abundan t (~200 g/m2). 

Nea r the slope break, coral and octocoral densities increase as mounds of Montastrea annularis 

( s i m high) mark the transition into the highrelief (~4.5 m) spur and groove zone. Commo n 

corals are shown in Figure 223.?. Porites astreoides, 

P. porites, and Acropora cervicornis are most abundant along groove margins. Madracis asperula occurs principally along the spur faces. 

Turf-covered pavements with abundan t Lobophora sp. broaden from the base of the spurs. 

These hardgrounds constitute almost 30 percent 

of the fore-reef slope, the remainde r being steep 

sandy slopes that include some coral rubble. Th e 

hard bottom supports numerous large vase 

sponges and scattered coral, mainly of the species 

Montastrea cavernosa, Colpophyllia sp., Meandrina 

meandrites (Linnaeus), Mycetophyllia sp., Agaricia 

sp., and Scolymia sp. 

GLADDEN SPIT, SOUTHERN PROVINCE (Figure 

223/).—The barrier reef bends aroun d Gladden 
NUMBER 12 523 

Spit and consists of large patch reefs that rise 

from a turf and fleshy-algae-covered pavement 

with sand channels. From here southward, the 

shallow-water reef is absent except as island 

fringes and reentrants. The transect crosses a 

large patch reef on the south side of Gladden Spit 

that has a segmented, locally emergent reef crest 

approximately 400 m long. 

Lagoon and back-reef areas are similar to those 

at Rendezvous Cay. Rubble constitutes 40 to 60 

percent of the reef surface across the crest and 

continues through the moderate-relief spur and 

groove zone. The rubble is composed mainly of 

branches oi Acropora palmata (~0.5 m long). Turf 

is the main cover along with the encrusting alga 

Neogoniolithon sp.; however, Porolithon pachydermum 

is also common. Millepora complanata and Palythoa 

sp. dominate the mounds on the reef crest. As the 

mounds grade into spurs, Acropora palmata and 

Agaricia agaricites become the most obvious components on spur margins. Knobby pillars of Montastrea annularis (~2 m high) increase in abundance where the spurs begin to break up and 

decrease in size as they grade into a pavement 

with sand channels. 

Fleshy algal macrophytes are dominant on the 

pavement and increase in biomass seaward. Sargassum sp. is the principal constituent; Dictyota sp., 

Lobophora sp., and Stypopodium sp. are abundant; 

and Galaxaura sp. and Neomeris sp. are common. 

Veneers of turf-stabilized sediment combine with 

red boring algae to form the predominant cover. 

Sand channels that are common in the shallow, 

inner fore-reef pavement decrease along the outer 

part of the transect. Mounds of coral are present 

on the pavement adjacent to the edges of channels 

and holes. Corals associated with the mounds are 

Diploria strigosa, Siderastrea radians, S. siderea, Montastrea annularis, and Agaricia agaricites. Corals, 

large octocorals, and vase-shaped and encrusting 

sponges are found on the outer pavement, but 

they are sparse. 

QUEEN CAY, SOUTHERN PROVINCE (Figure 

223^).—This transect crosses a 1.2 km section of 

reef, the longest segment of shallow-water reef 

south of Gladden Spit (Figure 222). Typical of 

southern province reefs that form reentrants, the 

back-reef flats are generally much narrower and 

the lagoons deeper than in the north. From the 

lagoon, a moderate slope ascends into coral 

mounds surrounded by scattered rubble. Patches 

of seagrasses occur in some of the slightly deeper 

water on the back-reef flat. Mounds of corals and 

coralline algae increase in size and frequency as 

they grade into the reef crest. Surface cover on 

mounds is primarily turf and encrusting coralline 

algae, but coral cover can reach 15 percent. Rubble is not prominent except in the crest, where it 

covers the moderately deep (~1 m) grooves between the broad massive spurs. These spurs coalesce locally to form the irregular reef crest. Turf, 

encrusting coralline algae, and Sargassum sp. are 

predominant, with lesser amounts of Palythoa sp. 

and Millepora complanata also present. Turbinaria 

turbinata occurs on the shallowest mound surfaces. 

Thickets of Acropora palmata, covering 50-60 percent of the bottom, occur immediately seaward 

of the reef crest and locally interfinger with the 

crest where mounds are spaced more widely. The 

low-relief spur and groove zone is mainly a turfcovered pavement with coral coverage approaching 25 percent. In places, the bottom is as much 

as 60 percent coarse sand; the remainder consists 

of low, turf-covered spurs. Gorgonia sp. and other 

octocorals mark the deeper parts of the spurs, 

spanning the transition from spurs to fleshy algal 

pavement. Spurs grade abruptly into a broad 

pavement where the most conspicuous components are Sargassum sp. and Dictyota sp. Encrusting 

and vase-shaped sponges highlight the surface. 

Near the shelf edge, a spur and groove zone 

with moderate relief is formed, and coral diversity 

and population densities increase. Dominant corals are listed in Figure 223g. Some of the less 

conspicuous forms include Madracis asperula, Eusmila fastigiata, Stephanocoenia michelinii Milne Edwards and Haime, Meandrina meandrites, Dichocoenia stokesi, Mycetophyllia spp., and Porites porites. 

Acropora cervicornis is found only along margins in 

the upper parts of grooves along the shelf edge. 

Lobophora sp. becomes increasingly abundant 

across the shelf-edge spur and groove zone. The 
88° 

«W Chinchorro 

" Bank 

A~zz&*k >$§£-

Lighthouse 

Reef 

WIND 

January July 

Sea Level Reef 

High Spur and Groove Features 

Coral Framework Ridge 

74.5° Mean Wave Direction 

Unimpeded Wave Force 

Modified Wave Force 

Maximum Impedence of Wave Force 

FIGURE 224.—Simplified diagram of the relationship between the magnitude of wave energy 

and sea level reefs, spur and groove features, and shelf-edge ridges along the Belizean barrier 

reef. Wave force impact on the reefs is controlled by wind speed and direction and by the 

impedence and modifying effect of the off-shore carbonate platforms. Wave patterns indicate 

mean wave direction (74.5°) and plus and minus 22.5° (97° and 52°), the sector containing 

87% of the frequencies. Note the extensive development of fore-reef structures in the shaded 

(wave impeded) zones. The question mark indicates lack of data for the shelf edge behind 

Chinchorro Bank. 
NUMBER 12 525 

fore-reef slope along the shelf edge drops abruptly 

from 27 m to below 1400 m. 

Discussion and Conclusions 

The Belizean barrier platform has a reef built 

within 0.2 m of sea level along 57 percent of its 

length. Of this sea level reef development, approximately 31 percent occurs in the northern 

province, 62 percent in the central province, and 

6 percent in the southern province. The distribution of fore-reef structures, their associated 

communities, and the distribution of the shallowwater reef system correlates with the distribution 

of wave energy reaching the barrier reef (Figure 

224). Mean wind direction, and thus mean wave 

direction, is equal to 74.5° (east-northeast) as 

determined by vector addition of directional frequencies of the wind. More than 95 percent of 

the waves along the fore reef come from easterly 

directions between 0° (north) and 180° (south). 

The remaining 5 percent are from westerly directions, particularly the northwest, and have little 

mechanical effect on the fore-reef communities. 

Eighty-seven percent of the easterly waves come 

from between 52° and 97°, or 45° about the 

mean wind. Wave energy impinging on the barrier reef is differentially attenuated and modified 

by the three carbonate platforms lying seaward 

of the barrier reef (Turneffe Islands, Lighthouse 

Reef, and Glover's Reef). The amount of wave 

modification and decrease in wave force depends 

on the direction of the incoming wave train relative to these carbonate platforms. 

The shelf-edge coral ridge and the shallow, 

high-relief spur and groove systems occur only in 

the zone of modified wave force. Preliminary 

calculations of mean annual wave energy indicate 

that the coral ridge occurs where wave energy is 

reduced to less than 40 percent of the total unimpeded wave energy. Shallow fore-reef spur and 

groove structures of high relief occur where wave 

energy is between 25 to 50 percent of the total. 

Double high spur and groove formations occur 

where wave energy is between 40 and 50 percent 

of the total. The positive correlation between 

zones of impeded and modified wave forces and 

extensive shallow-water reefs with abundant fragile corals in the fore reef suggests that seaward 

platforms have affected favorably the development of a continuous barrier reef in the central 

province by interfering with regional wave energy. Where open ocean waves have full access to 

the barrier reef, durable, slow-growing communities prevail and shallow-water reefs are narrow 

and discontinuous. 

High-relief spurs and coral ridges built by fragile branching corals are uncommon in Caribbean 

reefs. From the literature and personal observation of over 80 percent of the windward Caribbean coastlines, similar community development 

and structures are known only from Jamaica, 

north Haiti, and southeast Alacran. All of these 

areas are exposed to modified wave settings and 

are protected from long-period storm waves generated by prolonged and severe mid-latitude winter storms. Exposure to such severe wave energy 

conditions was postulated to account for the absence of emergent shelf-edge reefs in the eastern 

Caribbean (Adey, 1978). Conversely, protection 

from such extreme conditions has affected significantly the shelf-edge development along the central province of Belize by encouraging high rates 

of carbonate production and accumulation by 

rapid-growing, branching corals (Adey, 1978; 

Burke, 1979). 

Obviously, other factors have substantially influenced present reef distribution on the outer 

Belizean platform. A model proposed by Adey 

and Burke (1977) incorporating wave energy, 

turbidity, and pre-Holocene shelf depth was applied by Burke (1979) to explain Holocene reef 

development and distribution along the platform. 

This subject will be presented in greater detail in 

a forthcoming paper by Burke. 
526 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

Literature Cited 

Adey, W. H. 

1978. Coral Reef Morphogenesis: Multidimensional 

Model—New Data from Coring and C14 Dating 

Provide Data for Unravelling Some Classical Enigmas. Science, 202:831-847. 

Adey, W. H., and R. B. Burke 

1977. Holocene Bioherms of Lesser Antilles—Geological 

Control of Development. In S. H. Frost, M. P. 

Weiss, and J. B. Saunders, editors, Reefs and 

Related Carbonates—Ecology and Sedimentology, The American Association of Petroleum Geologists, 

Studies in Geology, 4:67-81. Tulsa, Oklahoma. 

Burke, R. B. 

1979. Morphology, Benthic Communities, and Structure 

of the Belize Barrier Reef. 78 pages. Master's 

thesis, University of South Florida, Tampa. 

James, N. P., and R. N. Ginsburg 

1978. The Deep Seaward Margin of Belize Barrier and Atoll 

Reefs. 201 pages. Miami Beach, Florida: University 

of Miami. 

James, N. P., R. N. Ginsburg, D. S. Marszalek, and P. W. 

Choquette 

1976. Facies and Fabric Specificity of Early Subsea Cements in Shallow Belize (British Honduras) Reefs. 

Journal of Sedimentary Petrology, 46(3):523-544. 

Smith, F.G.W. 

1971. A Handbook of the Common Atlantic Reef and ShallowWater Corals of Bermuda, the Bahamas, Florida, the 

West Indies, and Brazil. 164 pages. Coral Gables, 

Florida: University of Miami Press. 

Stoddart, D. R. 

1962. Three Caribbean Atolls: Turneffe Islands, Lighthouse Reef and Glover's Reef, British Honduras. 

Atoll Research Bulletin, 87:1-151. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays, October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 

1969. Post-Hurricane Changes on the British Honduras 

Reefs and Cays: Re-survey, 1965. Atoll Research 

Bulletin, 131:1-31. 

1974. Post-Hurricane Changes on the British Honduras 

Reefs: Re-survey, 1972. In A. M. Cameron, B. M. 

Campbell, A. B. Cribb, R. Endean, J. S. Jell, O. 

A. Jones, P. Mather and F. H. Talbot, editors, 

Proceedings of the Second International Coral Reef Symposium, 473-483. Brisbane, Australia: The Great 

Barrier Reef Committee. 

Thorpe, J. E., and P. K. Bregazzi 

1960. Experiments and Observations on the Corals at 

Rendezvous Cay. In Cambridge Expedition to British 

Honduras, 1959-1960, General Report, pages 22-28. 

Cambridge, England. 

Thorpe, J. E., and D. R. Stoddart 

1962. Cambridge Expedition to British Honduras. Geography Journal, 128:158-171. 

United States Naval Weather Service Command 

1974. Summary of Synoptic Observations—Caribbean and 

Nearby Island Coastal Marine Areas. Volume 1, 474 

pages. Asheville, North Carolina. 

Wantland, K. F., and W. C. Pusey III, editors 

1975. Belize Shelf—Carbonate Sediments, Clastic Sediments, and Ecology, The American Association of 

Petroleum Geologists, Studies in Geology, 2: 599 pages. 

York, M. E. 

1971. Patch Reef Coral Communities of Southern British Honduras and Illustrated Catalogue of Common British Honduras Corals. In K. F. Wantland 

and W. C. Pusey III, A Guidebook for the Field Trip 

to the Southern Shelf of British Honduras, appendix I. 

New Orelans, Louisiana: New Orleans Geological 

Society. 
Species-Area Relationships on Small Islands: 

Floristic Data from Belizean Sand Cays 

David R. Stoddart 

and F. Raymond Fosberg 

ABSTRACT 

Analysis of species records of plants from 34 

sand cays on the barrier reef and atolls of Belize, 

Central Amerca, shows a linear relationship between species number and the logarithm of island 

area over the range 0.13 ha to 16.50 ha. Shrubs 

and trees show only a slight increase in numbers 

with area, because of the small number of shrubs 

available to colonize available habitats and the 

clearing of native woodland for coconut plantations. Herb numbers increase more strongly with 

island area, probably as a result of disturbance of 

habitats by man. Distance from source areas 

forms a much weaker control of species numbers, 

because of the proximity of the cays to Central 

America and their equal distance from eastern 

Caribbean source areas. Comparison with other 

island groups, however, shows that the Belize cays 

have much larger floras than more remote oceanic 

islands of similar sizes. Limited inferences on 

colonization and extinction can be made from 

the species records. 

Introduction 

This paper presents the results of floristic studies on sand cays of the barrier reef and lagoon 

and the three offshore atolls (Turneffe Islands, 

Glover's Reef, Lighthouse Reef) of the coast of 

Belize (formerly British Honduras), Central 

America (Figure 225). 

David R. Stoddart, Department of Geography, University of Cambridge, Downing Place, Cambridge CB2 3EN, England. F. Raymond Fosberg, Department of Botany, Smithsonian Institution, Washington, D.C. 20560. 

Niering's (1963) study of the distribution of 

vascular plants on islets of Kapingamarangi Atoll, eastern Caroline Islands, showed constant 

species numbers independent of island size below 

a threshold of 1.4 ha, and a linear relationship 

between species number and logarithm of island 

area above this threshold (Figure 226). Wiens 

(1962:369) suggested that the threshold marked 

the size of the smallest island supporting a freshwater lens under Kapingamarangi rainfall conditions (mean annual rainfall 2000 mm). These 

data, based on 31 islands ranging in size from 

0.16 to 32.00 ha, were used by MacArthur and 

Wilson (1967) both to illustrate the fundamental 

control of species diversity by area, and also to 

suggest that on very small islands this control is 

less effective. Instead of Wiens' explanation, they 

proposed that very small islets are regularly subject to inundation and wave damage during 

storms, and that below a threshold of island 

stability, periodic extinction of species could be 

complete. Supporting data used by MacArthur 

and Wilson came from earlier surveys of the Dry 

Tortugas and the Florida Keys (Millspaugh, 

1907; Bowman, 1918), though recent work has 

shown that the Dry Tortugas data are unreliable 

(Stoddart and Fosberg, in prep.). 

The Kapingamarangi data provided an apparently simple illustration of MacArthur and Wilson's equilibrium theory of island biogeography: 

on the local (within-atoll) scale, area is the main 

determinant of species number, whereas the species pool available for colonization of the islands 

is filtered on a regional scale by distance from 

527 
TURNEFFE ISLANDS 

SANDBORE CAY 

IGHTHOUSE 

REEF 

,'fiU^.HALF MOON CAY 

GLOVER'S 

REEF 

NORTHEAST CAY 

i /j> LONG CAY 

--^MIDDLE CAY 

SOUTHWEST CAYS 

BUTTONWOOD CAY 

S'^'p \ /^NORT H , 

^F-MIDDLE^SILK CAYS 

'-^rtf sour 

HATCHET CAY 

, ROUND CAY 

/"—TPOMPION CAY 

A-/RANGUANA CAY 

/•J/NORTH SPOT 

/-/TO M OWEN'S CAYS 

/ / 

/^NORTHEAST SAPODILLA CAY 

'••^FRANK'S CAY 

-NICOLAS CAY 

-HUNTING CAY 

^LIME CAY 

RAGGED CAY 

km 50 

Three fathom line 

FIGURE 225.—Reef islands of the Belizean coast. 
NUMBER 12 529 

o 30— 

0-1 

> T T 

05 1 

"i I I | l I I I I 

5 10 

Area (ha) 

50 

FIGURE 226.—Relationship of number of species to logarithm of island area for plants on 

Kapingamarangi islands (source: Niering 1963:137). 

continental sources. Niering's analysis was further 

refined by Whitehead and Jones (1969), who 

classified the species on each of the Kapingamarangi islands into ecological groups: strand (salttolerant) species; non-strand species; and introduced species. They showed that the smallest 

islands on Kapingamarangi supported only 

strand species, and that introduced species were 

most numerous on larger, inhabited islands. 

Numbers of strand species per island were only 

weakly influenced by area, presumably because 

the pool of such species is itself limited in size, 

whereas the number of non-strand species increased sharply with increasing area. Whitehead 

and Jones (1969:176) concluded that "species 

diversity on both the large and small islets is 

controlled primarily by the ecology of the island." 

Unfortunately they did not indicate which species 

in the total flora of 99 species they assigned to 

each ecological category; the status of many must 

necessarily be conjectural. 

A rather different analysis of the same data 

was made by Stoddart (1975), who grouped the 

species into categories of herbs, shrubs, and trees. 

It was shown that (a) numbers of shrub species 

did not vary greatly with area because of the 

limited number of such species present in the 

Western Pacific and capable of surviving on such 

islands; (b) numbers of tree species did not increase on larger islands, largely because indigenous forests had been replaced by coconut plantations; and (c) herb species numbers (which 

include many weedy species) increased dramatically on larger, inhabited islands as a result of 

human interference with natural vegetation. Similar results came from an analysis of data from 

Aitutaki, Cook Islands, for 15 islands ranging in 

size from 1.0 to 71.3 ha. The Aitutaki case is 

particularly interesting, for the total flora is small 

(53 species) even though the islands are immediately adjacent to an inhabited volcanic island 

which forms a local reservoir of possible colonizers. The fact that many species have failed to 

colonize across a water gap of less than 10 km 

suggested that the restricted flora on the islets 

reflected ecological control rather than the remoteness of Aitutaki itself from source areas. 

There remains, however, a dearth of floristic data 
530 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

on similar small islets with which the interpretations of MacArthur and Wilson (1967) and 

Whitehead and Jones (1969) can be tested and 

refined. 

ACKNOWLEDGMENTS.—This paper results from 

collections made by Stoddart during 1959-1965, 

supported by the Department of Scientific and 

Industrial Research (London), the Office of Naval Research (Washington), the Coastal Studies 

Institute (Louisiana State University), and the 

Royal Society (London); by Fosberg, M.-H. 

Sachet (Smithsonian Institution), and Stoddart 

on Glover's Reef in 1971, as part of the Smithsonian Institution's CITRE (Comparative Investigations of Tropical Reef Ecosystems) Program; 

and by Fosberg, D. L. Spellman (Missouri Botanical Garden), and Stoddart along the barrier reef 

in 1972, as part of the Smithsonian Institution's 

IMSWE (Investigations of Marine Shallow-Water Ecosystems) Program. 

Methods and Data 

Sand cays have been mapped on the coast of 

Belize in 1960, 1961, 1962, 1965, 1971, and 1972, 

all more than once and many on several occasions, and vascular plants have been both collected and recorded. The islands are accumulations of coral cobbles and reef-derived sand, 

mostly 50 to 200 m in greatest dimension, exceptionally reaching 1500 m, usually rising 1 to 3 m 

above mean sea level and occasionally reaching 

5 m. They support a varied vegetation of pioneer 

herbs, vines and grasses, scrub, scrub woodland, 

broadleaf forest, and coconut woodland, with, in 

protected situations, mangrove woodland. All the 

islands are periodically subject to severe hurricane 

damage. Island areas reported here derive from 

planimetric measurements of maps made during 

the surveys. Where the areas have changed over 

the period of study, the figure quoted is that for 

the most recent survey, usually 1971 or 1972 

(Stoddart, 1962, 1963, 1965, 1974). 

The total cay flora comprises approximately 

171 species, of which 68 are common to islets of 

the coastal barrier-lagoon system and to one or 

more of the atolls, 91 are found only on the 

barrier and or lagoon cays, and 12 are restricted 

to the atolls. One hundred and fifty eight species 

are found on the barrier and lagoon islets, of 

which 31 are introduced. The three offshore atolls 

have a combined flora of 87 species (of which 

three are introduced): 36 are recorded from Turneffe, 55 from Lighthouse Reef, and 63 from 

Glover's Reef. The atoll records, which reflect 

collecting intensity, compare with other Caribbean atolls as follows: Dry Tortugas, 45 species, 

plus more than 50 introductions (Millspaugh, 

1907; Bowman, 1918; Davis, 1942); Alacran, 25 

species, including 5 introductions (Fosberg, 1962), 

Morant Cays, 16 species, including one introduction (Chapman, 1944; Asprey and Robbins, 

1953), and Pedro Bank, 10 species, including one 

(possibly two) introductions (Zans, 1958; Fosberg, 

unpublished records in 1962; Chapman, 1944). 

Plants have been recorded from a total of 97 

Belizean reef islands. The records suggest that the 

floristic data are sufficiently comprehensive for 

analysis for 40 islands, but other considerations 

(such as presence of extensive mangroves) reduced 

this to 34, located as indicated in Tables 44 and 

45. The most extensive plant collecting was carried out by Fosberg, M.-H. Sachet, and Stoddart 

on Glover's Reef in 1971, and by Fosberg, D. L. 

Spellman, and Stoddart on the barrier reef and 

lagoon cays in 1972, supplementing earlier collections made by Stoddart in 1960, 1961, 1962, and 

1965. The vegetated islands studied range in size 

from 0.13 ha (North Spot) to 16.5 ha (Half Moon 

Cay); islands smaller than 0.10 ha did not possess 

vascular plants (for instance, Curlew Cay, 0.03 

ha; Paunch Cay, 0.06 ha). Most of the collections 

made are represented in the U.S. National HerTABLE 44.—Number and location of Belizean reef islands 

for which data on vegetation are available 

Location 

Barrier reef 

Lagoon 

Turneffe Islands 

Lighthouse Reef 

Glover's Reef 

Data 

available 

Data 

acceptable 

Data 

used 

29 

34 

22 

6 

6 

25 

5 

0 

4 

6 

24 

2 

0 

2 

6 
NUMBER 12 531 

barium, with a fairly good set at the Missouri 

Botanical Garden. 

Results 

SPECIES-AREA-DISTANCE RELATIONSHIP.— 

Numbers of species of vascular plants, excluding 

seagrasses, for the 34 selected islands are plotted 

against logarithmic island area in Figure 227, and 

the data are listed in Table 45. As in Amerson's 

(1975) study of the Leeward Hawaiian Islands, 

species number refers to the total number of 

species ever recorded from the island rather than 

the number recorded during a single survey. Under conditions of rapid turnover, therefore, it will 

generally exceed the number of species present at 

any one time. Twenty of the islands are smaller 

than Wiens' Kapingamarangi threshold of 1.4 ha, 

but no inflection is apparent in the Belize distribution; there is a linear relationship between 

species number and logarithm of island area over 

the range 0.13 ha to 16.5 ha. 

The species recorded for each island have also 

been classified into trees, shrubs, and herbs, and 

each category plotted against logarithmic area in 

Figure 228. Assigning species to these categories 

raises considerable problems. Decisions have been 

made on a taxonomic rather than a physiognomic 

basis, so that a tree species is recognized even if 

only a seedling is present. Some distinctions are 

arbitrary; thus, all mangroves are classified as 

trees except for Conocarpus erecta Linnaeus, which 

is classified as a shrub. Assignments are consistent 

between islands, in spite of considerable differences in size and habit of individual plants in 

different places. 

Shrubs and trees show a very slight increase in 

numbers with area, because of the restricted size 

of the shrub species pool capable of colonizing 

such small islands; because some shrub species 

(notably Suriana maritima Linnaeus) may form 

extensive single-species stands on larger islands, 

thus excluding other shrub species; and because 

most of the islands now support coconut woodland rather than indigenous forest. Herb numbers 

increase more dramatically with area, probably 

as a result of the prevalence of weedy species and 

human introductions on larger islands, especially 

those that are inhabited, and by the provision of 

suitable habitat for these species on periodically 

cleared ground under coconuts. Abnormally low 

herb numbers on some larger islands can be 

FIGURE 227.—Relationship of number of species to logarithm of island area for plants on 

Belizean reef islands. 
532 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

TABLE 45.—Records of vascular plants from Belizean reef islands (listed north to south within 

each geographic feature) 

Island 

BARRIER REEF 

Sergeant's 

Goffs 

English 

Rendezvous 

Cay Glory 

Tobacco 

South Water 

Carrie Bow 

North Silk 

Middle Silk 

South Silk 

Round 

Pompion 

Ranguana 

North Spot 

Tom Owen's West 

Tom Owen's East 

Northeast Sapodilla 

Frank's 

Nicolas 

Hunting 

Lime 

Ragged 

Seal 

BARRIER LAGOON 

Buttonwood 

Hatchet 

LIGHTHOUSE REEF 

Sandbore 

Half Moon 

GLOVER'S REEF 

Northeast 

Long North 

Long 

Middle 

Southwest I 

Southwest II 

Total 

number 

of 

species 

20 

13 

23 

24 

7 

44 

70 

13 

13 

13 

9 

16 

21 

23 

7 

20 

15 

26 

31 

30 

54 

26 

7 

16 

27 

38 

24 

43 

33 

16 

22 

38 

20 

32 

Number 

of 

tree 

species 

4 

3 

5 

6 

1 

9 

11 

3 

3 

1 

2 

3 

7 

3 

2 

4 

3 

6 

7 

7 

13 

5 

2 

7 

6 

7 

4 

10 

9 

2 

5 

11 

4 

3 

Number 

of 

shrub 

species 

3 

0 

2 

3 

0 

7 

9 

1 

4 

4 

1 

3 

4 

3 

1 

2 

3 

4 

4 

6 

8 

3 

1 

1 

3 

4 

5 

6 

5 

4 

5 

5 

3 

3 

Number 

of 

herb 

species 

13 

10 

16 

15 

6 

28 

50 

9 

6 

8 

6 

10 

10 

17 

4 

14 

9 

16 

20 

17 

33 

18 

4 

8 

18 

27 

15 

27 

19 

10 

12 

22 

13 

26 

Area 

(ha) 

0.17 

0.32 

0.49 

0.18 

0.13 

2.50 

6.49 

0.39 

0.25 

0.40 

0.23 

0.31 

0.42 

1.12 

0.13 

0.24 

0.12 

5.06 

1.74 

3.86 

5.94 

2.07 

0.22 

0.14 

0.72 

1.30 

5.34 

16.50 

4.48 

0.53 

5.85 

5.97 

8.70 

6.23 

Distance 

from 

mainland 

(km) 

18.7 

25.7 

24.4 

26.4 

25.7 

18.2 

18.4 

18.7 

36.0 

36.0 

36.0 

32.6 

31.3 

32.6 

34.1 

33.3 

33.6 

36.0 

35.5 

35.7 

36.2 

36.2 

35.5 

27.8 

30.5 

32.1 

73.6 

78.3 

52.2 

51.5 

51.0 

49.5 

49.5 

45.5 

explained either by the persistence of dense 

thicket instead of coconut woodland (as on Nicolas Cay), or by the presence of extensive marshy 

areas within the cay, which reduce its effective 

area for dry-land plants (as on Northeast Sapodilla Cay and Southwest I Cay). The results of 

this analysis thus resemble those carried out for 

Aitutaki and Kapingamarangi (Stoddart 1975, 

fig. 33). 

Floristic diversity has also been analyzed in 

terms both of island area and of distance from 

the mainland coast of Central America. Linear, 

quadratic, and cubic trend surfaces have been 

fitted to species number data on coordinates of 
Area (ha) 

FIGURE 228.—Relationship of number of species to logarithm of island area for trees, shrubs, 

and herbs on Belizean reef islands. 

logarithmic island area (ha X 10) and logarithmic 

distance from mainland (km). This has been done 

both for total species and for total less introduced 

species. Values of R2 for the surfaces are given in 

Table 46. 

Equations for the linear surfaces are 

zi = 75.17702 + 18.96162* - 45.58615? 

Z2 = 56.09279 + 17.58632* - 33.31592? 

where z\ is number of all species, Z2 is number of 

indigenous species, x is logarithmic island area 

(ha X 10), andy is distance from mainland in km. 

Figure 229 plots the z\ surface for the islands 

studied and illustrates the relatively weak influence of distance compared with area, as might be 

expected for dominantly water-dispersed plants 

so close to continental shores. 

The use of a parameter such as distance from 

mainland to indicate isolation is not particularly 

useful. Currents in the region are dominantly 

north-south, transverse to the shortest-distance 

paths between islands and mainland, and since 

the area is affected by the Northeast Trades, the 

mainland in fact is leeward of the cays. Distances 
534 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

1-95 

Log Island Area (ha x IO) 

1 0 1-5 

1-75 

S 1-50 

1-25 

10 

Island Area (ha) 

FIGURE 229.—Linear trend surface for total species number in terms of island area and 

distance from mainland for Belizean reef islands. 

between the islands are probably insignificant for 

propagules arriving as drift from the eastern Caribbean. Circulation gyres in the Gulf of Honduras 

may also help to explain the remarkable absence 

on this coast of a species such as Scaevola pulmieri 

(Linnaeus) Vahl, which is widespread in the eastern Caribbean in similar habitats and which 

reaches the Cayman Islands (740 km to windward 

of Belize) and even the norhern coast of Yucatan 

(Sauer, 1959, 1967) and the Nicaraguan reefs 

(Milliman, 1969:9). Nevertheless, the data preTABLE 46.—Plant diversity related to island area and distance of island from mainland coast 

Trend surface 

Linear 

Quadratic 

Cubic 

All species Indigenous species 

0.591 

0.651 

0.681 

0.697 

0.742 

0.779 

sented in Figure 229 correspond with MacArthur 

and Wilson's prediction that highest species numbers will be found on large, near-to-mainland 

islands and lowest numbers on small, far islands, 

though this correspondence does not necessarily 

imply the operation of the mechanisms leading 

to dynamic equilibrium that they propose. 

The effect of distance is more dramatically 

shown in the comparison of species numbers between one of the atolls (Glover's Reef), Alacran 

Reef in the Gulf of Mexico, and Pedro Cays south 

of Jamaica (Figure 230). The higher diversities 

on Glover's Reef islands probably result from 

their proximity to the mainland, compared with 

the other more isolated reefs, although climatic 

differences may also be significant. Comparable 

rainfall records are not available for the three 

locations, but both Alacran and Pedro, with probably 1000 mm/y or less, are considerably drier 
NUMBER 12 535 

than Glover's Reef. A similar inference may be 

drawn from Figure 231, which plots generalized 

curves for species numbers on the Belizean barrier 

islands, those of Kapingamarangi (Niering, 1963), 

and the leeward Hawaiian Islands in the central 

north Pacific (Amerson, 1975): the curves are of 

similar form, reflecting the influence of area 

within each island group, but are displaced in 

magnitude, reflecting the effect of distance relative to source areas. 

SPECIES COMPOSITION CHANGES OVER TIME.— 

Species turnover is more readily determined for 

60 

40 

20 

0 1 

(•_. 

.^r\ 

y 

GLOVER'S 

\ 

\ 

' \ 

\ 

(* 

ALACRAN /__ 

/ V 

PEDRO CAYS 

"i—i—i—i—r 

10 

1 I I I I TT 

100 

Area (ha) 

FIGURE 230.—Relationship of number of species to logarithm of island area for plants on 

individual islands on Glover's Reef, Alacran Reef, and Pedro Bank. 

Area (ha) 

FIGURE 231.—Generalized species-area relationships for islands of the Belizean barrier reef, 

Kapingamarangi Atoll, and the Leeward Hawaiian Islands. 
536 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

plants than for some other kinds of organisms 

(compare the criteria for avifaunal turnover discussed by Lynch and Johnson, 1974), but the 

problems of insuring completeness of the record 

are such that we do not think our data can 

support inferences of colonization or extinction 

from new records of plants made during repetitive 

surveys between 1960 and 1972. The single exception refers to small islands devastated by a major 

hurricane in 1961, where it has been possible to 

follow the course of plant colonization and revegetation (Stoddart, 1974). Data in Table 47 refer 

to four small islands on the northern barrier reef, 

the first three of which had their pre-existing 

plant cover entirely or almost entirely removed 

by the storm; the fourth retained a number of 

species, although all were much damaged. These 

data suggest that at least a decade is required for 

a small island to be re-stocked, and it would be 

useful to monitor the future development of these 

floras (compare also Riitzler and Ferraris, herein: 

77). 

It is easier to infer local extinction from failure 

to find plants recorded during earlier surveys 

than it is to infer colonization. Extinctions are of 

two kinds: catastrophic extinction resulting from 

severe storm effects, of the kind envisaged by 

MacArthur and Wilson (1967) for the smaller 

Dry Tortugas; and local random extinction. In 

the case of beach-crest shrubs, the latter may 

result from small-scale beach retreat destroying a 

particular habitat to which a species is confined. 

This has occurred in the extinction of Sophora 

tomentosa Linnaeus on Tobacco Cay between 1965 

and 1972, and of Tournefortia gnaplhalodes (LinTABLE 47.—Post-hurricane recovery of vegetation on four 

small barrier reef islands (data for each year include number 

of species and island area (ha) in parentheses) 

Island 

Sergeant's Cay 

Goffs Cay 

English Cay 

Rendezvous Cay 

1962 1965 1972 

4 (0.20) 

2 (0.23) 

2 (0.34) 

16 (0.22) 

12 (0.16) 

4 (0.29) 

5 (0.35) 

16 (0.20) 

14 (0.18) 

10 (0.32) 

18 (0.49) 

17 (0.18) 

naeus) R. Brown on Nicolas, Hunting, and Lime 

cays between 1960 and 1972. Extinctions of the 

first type occur only on smaller islands, generally 

smaller than 1 ha, but those of the second may 

be independent of island size. Since total species 

number is low on small islands, percentage turnover resulting from catastrophic damage is correspondingly high. Figure 232 plots extinctions 

on barrier reef islands determined in 1972 by 

comparison with records from 1960, 1962, and 

1965, both as absolute numbers of extinct species 

against logarithmic area, and as percentages of 

the total number of species recorded from the 

island against logarithmic area. These data emphasize that very small islands (less than 0.5 ha) 

are particularly prone to extinction of their 

plants, as MacArthur and Wilson (1967:54-55) 

hypothesized. 

Discussion 

The data reported here supplement those for 

Kapingamarangi and Aitutaki; they include data 

on many very small islands, over which controversy has centered; and they apply to islands with 

much higher species numbers than those previously discussed. 

Total recorded species number varies directly 

with area over the whole size range of islands 

considered, down to 0.1 ha; there is no indication 

of the 1.4 ha threshold in area attributed by 

Wiens to the minimum size required for freshwater lens formation, and by MacArthur and 

Wilson to the instability of substrates below this 

size. We have no records on fresh-water lens 

formation on Belizean islands (although freshwater wells are not found on islands smaller than 

2.5 ha). Our data suggest, however, that presence 

or absence of a lens may be less critical than 

Wiens supposed, especially in explanation of so 

crude a parameter as total species number and 

especially so far as herb species numbers are 

concerned. Further, hurricane activity on these 

islands is so regular and occasionally so intense 

that if a threshold of instability does exist, it may 

be higher than the 1.4 ha previously suggested. 

Nevertheless, there is no indication in the Beli-
NUMBER 12 537 

Q 

— • • 

4 — 

— • • • 

2— • • • • • • 

— ••• • • • • • • 

0 1 1 1 1 1 1 r—i—i 1 1 1 1 , — 

0-1 1-0 

1-0 

Area (ha) 

FIGURE 232.—Species extinctions on Belizean barrier reef islands between 1960 and 1972: a, 

absolute number of species extinct; b, extinctions as a percentage of the total recorded flora of 

the island. 
538 SMITHSONIAN CONTRIBUTIONS TO THE MARINE SCIENCES 

zean data of any marked discontinuity in the area 

control. 

The evidence also indicates that with the Belizean reef system, distance from mainland is a 

relatively weak control of species number on islands. Both Ridley (1930:689) and Carlquist 

(1974:45-96) have noted that many island species 

are water-dispersed with high efficiency, and that 

many are pan-tropical or at least extremely wideranging in their distribution. There is also negative evidence of the unimportance of distance as 

a diversity control in the absence of many species 

on coral islands, even when source reservoirs exist 

only a few kilometers away. This suggests that 

ecological controls are more important than location (Stoddart, 1975:93). Nevertheless, speciesarea curves for Belize, Alacran, Pedro Bank, Kapingamarangi, and the Leeward Hawaiian Islands, which vary widely in degree of isolation 

and also in floristic diversity, suggest that distance 

imposes a gross regional control by limiting numbers on the most distant islands. 

The Belizean cays, however, lie close to continental shores, and we infer that their floras are 

not distance limited, as are those of the Leeward 

Hawaiian Islands. It follows that species numbers 

are not restricted by any reduction in the rate of 

propagule arrival, but rather by the ability of 

species to establish themselves once they have 

arrived, that is, by an ecological control. Although quantitative data are lacking, casual observation of germinating seeds indicates that a 

constant stream of propagules, many of which 

fail to develop into mature plants, is arriving on 

the islands. If it were not for this ecological control 

rendering many colonization attempts abortive, 

the floras of the islands could not be in equilibrium since known local extinction rates are probably low, except on the smallest hurricane-devastated islands. 

This argument parallels that of Lack (1976:1-

14) for West Indian birds. Lack documented 

many cases of the arrival of birds on islands where 

they failed to establish themselves (for example, 

humingbirds on the Cayman Islands). He ascribed this phenomenon to ecological control 

("ecological proverty"), while admitting that it 

was unclear how such control operated. We 

would like to draw attention to the need to study 

the mechanisms, notably environmental control 

and competition, which similarly prevent the establishment of plants. In contrast to the enormous 

literature on dispersal, which in our case is so 

effective as to be almost irrelevant in differentiating the floras of the cays, virtually no attention 

has been given to this crucial problem. Without 

such study, theoretical explanations of species 

numbers must remain speculative. 

It is also possible that the closeness of the 

control of species numbers by area is much exaggerated by using aggregate numbers rather 

than numbers of species existing at any one time. 

Under conditions of high turnover, competition 

may lead to replacement of species rather than to 

packing of more species into a limited area, that 

is, the effects are temporal rather than spatial, so 

that species numbers remain constant at any 

time, but over time, the aggregate of species 

recorded increases up to a maximum set by the 

available species pool. The fact that many species 

often form extensive single-species stands on coral 

islands (for instance, Scaevila taccada (Gaertner) 

Roxburgh in the Indo-Pacific and Suriana maritima 

Linnaeus in the Gulf of Mexico) supports this 

suggestion. We have no means of estimating the 

effect of using aggregate species numbers in the 

Belizean case, but it may be noted that on Cayo 

Ahogado, Puerto Rico, admittedly an extremely 

small island (0.065 ha) with continuously changing topography, Heatwole and Levins (1973) recorded a total of 51 plant species; the maximum 

present at any one time was 21, and the number 

that might be considered established was only 9. 

Finally, we would like to sound a note of 

caution. In spite of many years of activity by 

several workers on these islands, we would be 

reluctant to undertake a more sophisticated analysis, particularly of colonization and extinction, 

without a great deal more investigation. Our data 

will, however, serve as a basis for continued monitoring of floristic change on these islands in the 

future. 
NUMBER 12 539 

Literature Cited 

Amerson, A. B., Jr. 

1975. Species Richness on the Nondisturbed Northwestern Hawaiian Islands. Ecology, 56:435-444. 

Asprey, G. F., and R. G. Robbins 

1953. The Vegetation of Jamaica. Ecological Monographs, 

23:359-412. 

Bowman, H.H.M. 

1918. Botanical Ecology of the Dry Tortugas. Papers from 

the Department of Marine Biology, Carnegie Institution 

of Washington, 12:109-138. 

Carlquist, S. 

1974. Island Biology. 660 pages. New York. 

Chapman, V. J. 

1944. 1939 Cambridge Expedition to Jamaica, Part 1: 

A Study of the Botantical Processes Concerned in 

the Development of the Jamaican Shoreline. Journal of the Linnean Society, Botany, 52:407-447. 

Davis. J. H. 

1942. The Ecology of the Vegetation and Topography 

of the Sand Keys of Florida. Papers from the Tortugas 

Laboratory, Carnegie Institution of Washington, 33:113-

195. 

Fosberg, F. R. 

1962. A Brief Survey of the Cays of Arrecife Alacran, a 

Mexican Atoll. Atoll Research Bulletin, 93:1-25. 

Heatwole, H., and R. Levins 

1973. Biogeography of the Puerto Rican Bank: SpeciesTurnover on a Small Cay, Cayo Ahogado. Ecology, 

54:1042-1055. 

Lack, D. 

1976. Island Biology Illustrated by the Land Birds of Jamaica. 

445 pages. Oxford. 

Lynch, J. F., and N. K. Johnson 

1974. Turnover and Equilibria in Insular Avifaunas, 

with Special Reference to the California Channel 

Islands. Condor, 76:370-384. 

MacArthur, R. H., and E. O. Wilson 

1967. The Theory of Island Biogeography. 203 pages. Princeton. 

Milliman, J. D. 

1969. Four Southwestern Caribbean Atolls: Courtown 

Cays, Albuquerque Cays, Roncador Bank and 

Serrana Bank. Atoll Research Bulletin, 129:1-26. 

Millspaugh, C. F. 

1907. Flora of the Sand Keys of Florida. Field Columbian 

Museum Publication, Botantical Series, 2, 118:191-245. 

Niering, W. A. 

1963. Terrestrial Ecology of Kapingamarangi Atoll, 

Caroline Islands. Ecological Monographs, 33:131-

160. 

Ridley, H. N. 

1930. Island Floras. In The Dispersal of Plants Throughout 

the World, pages 575-590. London. 

Sauer, J. D. 

1959. Coastal Pioneer Plants of the Caribbean and Gulf 

of Mexico. 21 pages 53 figures, 11 tables; Appendix, 20 pages. University of Wisconsin, Departments of Botany and Geography, Madison. [Mimeographed.] 

1967. Geographic Reconnaissance of Seashore Vegetation along the Mexican Gulf Coast. Louisana State 

University Coastal Studies Series, 21:1-59. 

Stoddart, D. R. 

1962. Three Caribbean Atolls: Turneffe Islands, Lighthouse Reef, and Glover's Reef, British Honduras. 

Atoll Research Bulletin, 87:1-151. 

1963. Effects of Hurricane Hattie on the British Honduras Reefs and Cays, October 30-31, 1961. Atoll 

Research Bulletin, 95:1-142. 

1965. British Honduras Cays and the Low Wooded Island Problem. Transactions of the Institute of British 

Geographers, 36:131-147. 

1969. Post-Hurricane Changes on the British Honduras 

Reefs and Cays: Re-Survey of 1965. Atoll Research 

Bulletin, 131:1-25. 

1974. Post-Hurricane Changes on the British Honduras 

Reefs: Re-survey of 1972. In A. M. Cameron, B. 

M. Campbell, A. B. Cribb, R. Endean, J. S. Jell, 

O. A. Jones, P. Mather and F. H. Talbot, editors, 

Proceedings of the Second International Coral Reef Symposium, 2:473-483. Brisbane, Australia: The 

Greater Barrier Reef Committee. 

1975. Vegetation and Floristics of the Aitutaki Motus. 

Atoll Research Bulletin, 190:87-116. 

Stoddart, D. R., and F. R. Fosberg 

In prep. Topographic and Floristic Change, Dry Tortugas, Florida, 1904-1977. 

Whitehead, D. R., and C. E. Jones 

1969. Small Islands and the Equilibrium Theory of Insular Biogeography. Evolution, 23:171-179. 

Wiens, H. J. 

1962. Atoll Environment and Ecology. 532 pages. New Haven. 

Zans, V. A. 

1958. The Pedro Cays and Pedro Bank: Report on the 

Survey of the Cays 1955-57. Bulletin of the Geological 

Survey Department, Jamaica, 3:1-47.