Marine and terrestrial ecosystems of the Virgin Islands National Park and biosphere reserve

[From the U.S. Government Printing Office, www.gpo.gov]

MARINE AND TERRESTRIAL ECOSYSTEMS
OF THE VIRGIN ISLANDS NATIONAL PARK
AND BIOSPHERE RESERVE

BIOSPHERE RESERVE REPORT NO. 29

Prepared by:

Caroline S. Rogers
Virgin Islands National Park
St. John, U.S. Virgin Islands

and

Robert Teytaud
Island Resources Foundation
St. Thomas, U.S. Virgin Islands

r
Propecrty of ('CC Library

U.S. Department of The Interior
National Park Service
and
Virgin Islands Resource Management Cooperative
Virgin Islands National Park
P.O. Box 7789, St. Thomas
U.S. Virgin Islands 00801

[. S. DEPARTMENT OF COMMERCE NOAA
Local Contracting Agent CCASIAL SERVICES CENTER
*234 SOUTH HOBSON AVENUE
ISLAND RESOURCES FOUNDATIOI .ARLESTON, SC 29405-2413
Red Hook Box 33, St. Thomas
U.S. Virgin Islands 00802
(NPS Contract No. CX-0001-3-0048)

September, 1988

TABLE OF CONTENTS

TABLE OF CONTENTS i
LIST OF FIGURES iii
LIST OF TABLES iv
PREFACE v
FOREWORD vi
INTRODUCTION 1
ACKNOWLEDGMENTS 2
PHYSICAL SETTING AND CLIMATE 3
BRIEF HISTORIC OVERVIEW 6
MARINE SYSTEMS 8
HISTORY OF RESEARCH 8
DESCRIPTION OF MARINE SYSTEMS 9
Coral Reefs 10
Seagrass Beds 13
Mangroves 15
Salt Ponds 17
Algal Plains and Marine Algae 18
Beaches and Rocky Shores 18
STRESSES TO MARINE SYSTEMS 20
Sedimentation 20
Sedimentation studies within VINP/BR 21
Hawksnest Bay 22
Fish Bay 22
Reef Bay 22
Coral growth 22
Effects of sedimentation on coral
metabolism 23
Sedimentation rates 23
Recreational Uses 23
Anchor damage 25
Physical damage to Windswept and
Hawksnest Reefs 25
Oil Pollution 26
Natural Stresses 27
Storms 27
Coral diseases 27
Coral bleaching and other stresses 28
ENDANGERED AND THREATENED MARINE SPECIES 29
Sea Turtles 29
Marine Mammals 31
Other Species 31
FISHERIES 32
SOCIOECONOMIC AND CULTURAL ROLE OF FISHING IN VIBR 32
ASSESSMENT OF FISH AND SHELLFISH STOCKS 34
FISH COMMUNITY STRUCTURE 35
FISHERIES LANDINGS ON ST. JOHN 36
FISH POISONING 37
HABITATS OUTSIDE VINP OF ECOLOGICAL IMPORTANCE TO
BIOSPHERE RESERVE FISHERIES 37
LOBSTER AND CONCH FISHERIES 38

Conchs 39
Lobsters 40
OTHER FISHERIES 42
FISHERIES OF BUCK ISLAND REEF NATIONAL MONUMENT 43
LESSER ANTILLEAN REGIONAL FISHERIES 43
FISHERIES REGULATIONS IN THE USVI 44
RECOMMENDATIONS FOR FUTURE RESEARCH AND MANAGEMENT
OF MARINE RESOURCES 46
TERRESTRIAL SYSTEMS 48
HISTORY OF RESEARCH: FLORA AND VEGETATION 48
DESCRIPTION OF TERRESTRIAL SYSTEMS 49
Flora and Vegetation 49
soils 53
Groundwater 54
HISTORY OF RESEARCH: FAUNA 54
Invertebrates and Aquatic Fauna 55
Amphibians and Reptiles 56
Birds 57
Mammals 60
STRESSES TO TERRESTRIAL SYSTEMS 62
Soil Erosion and Runoff 62
Development 65
RECOMMENDATIONS FOR FUTURE RESEARCH AND MANAGEMENT
OF TERRESTRIAL RESOURCES 67
GEOLOGY 69
BIOGEOGRAPHY 73
RECENT GEOLOGICAL RESEARCH ON ST. JOHN 75
CONCLUSIONS 76
APPENDIX I 77
APPENDIX II 78
APPENDIX III 79
APPENDIX IV 82
INDEX 109

LIST OF FIGURES

Fig. 1. Location of St. John, U.S. Virgin Islands.
From: Putney (1982). 4
Fig. 2. Great House at Par Force Plantation, Reef Bay
Quarter (1988). Photo: S. Barry. 7
Fig. 3. Johnson's Reef at a depth of 60'.
Photo: C. Weikert. 11
Fig. 4. Map of major benthic communities, Hawksnest Bay.
From: Beets et al. 1986. 12
Fig. 5. School of grunts sheltering in a patch reef
off Scott Beach. Photo: L. McLain. 14
Fig. 6. Prop roots of red mangroves are important
nursery areas for fish. Drawing: M. Beath. 16
Fig. 7. Salt pond on St. John. Photo: L. McLain. 17
Fig. 8. Trunk Bay Beach on St. John. Photo: L. McLain. 19
Fig. 9. Recreational visits to Virgin Islands National
Park from 1957 - 1987. 24
Fig. 10. Boats in park waters from 1966 to 1987. 24
Fig. 11. Green sea turtle (Chelonia mydas) over seagrass
and sand bottom. Photo: L. McLain. 30
Fig. 12. Queen conch (Strombus gigas) in a seagrass bed
in Brown Bay. Photo: L. McLain. 39
Fig. 13. A spiny lobster (Panulirus argus) on
Johnson's Reef. Photo: L. McLain. 41
Fig. 14 Vegetation map of St. John.
From: Woodbury & Weaver 1987. 51
Fig. 15. Example of the Louisenhoj Formation, south end
of Trunk Cay. Photo: D. Rankin. 70
Fig. 16. Bedded siltstone of the Tutu Formation, point
between Francis and Maho Bays.
Photo: D. Rankin. 70

LIST OF TABLES

Table 1. Rare and endangered species of plants from
St. John. From: Woodbury & Weaver 1987. 50

PREFACE

The Virgin Islands Resource Management Cooperative (VIRMC)
is a relatively new, interdisciplinary grouping of resource
professionals concerned with island environments and the role of
parks and protected areas in sustainable development. After
nearly five years of continuous effort involving more than a
dozen institutions and over fifty researchers, graduate students,
interns, and peer reviewers, the ubiquitous administrators and
editors at Island Resources Foundation (IRF) have, in the name of
VIRMC, completed the twenty-nine part technical report series, of
which this "Synthesis' volume, by Dr. Caroline Rogers and Robert
Teytaud, is the last of the formal publications.

The story of how this extended series of research projects
got started and what it has to do with UNESCO and the Man and the
Biosphere program is touched upon by William Gregg in the
Foreword to this publication. There is no room to expand upon
the subject in this short Preface, especially since the details
and nuances lead into other more complex and sometimes unresolved
issues including the role of the Biosphere Reserve Center and its
longer term objectives for enlarging its training, education,
research and environmental monitoring activities.

To be fair, one should mention that not everybody agrees
that the idea of an eventual Eastern Caribbean multi-island,
multi-site, multi-national Biosphere Reserve is workable or
appropriate in the Eastern Caribbean. We certainly do not, as
yet, have enough management experience to know if the biosphere
reserve model, as it is being developed on St. John, has the
potential for being more flexible than, for example, a national
park as a resource protection and management strategy in selected
small island settings.

But for the moment, thanks to the VIRMC baseline research
effort and technical report series, and especially to this 'user-
friendly" synthesis, we do have a solid point of departure or
information baseline for the next stage of this grand
interdisciplinary experiment in island resource management. The
U.S. National Park Service, the Virgin Islands National Park, and
the Virgin Islands Resource Management Cooperative can be proud
of what they have accomplished. The Foundation is pleased to
have been associated with the adventure.

Dr. Edward L. Towle, President
Island Resources Foundation
and
Program Manager for VIRMC (1983-88)

FOREWORD

In 1983, resource planners, managers, and scientists from 13
Caribbean islands met on St. John in the U.S. Virgin Islands to
discuss the role of biosphere reserves and other protected areas
in the sustainable development of the Lesser Antillean province.
For most participants, the conference was the fir.st encounter
with biosphere reserves and their potential benefits to the
people of small tropical islands. For most, it was also their
first contact with UNESCO's Man and the Biosphere Program (MAB),
in which 114 nations are now participating.

The International Network of Biosphere Reserves is a cornerstone
of the MAB program. In biosphere reserves, the emphasis is on
developing and sharing information, locally, regionally, and
internationally, to achieve the purpose of MAB. The base of
operations for the ideal biosphere reserve includes one or more
natural areas where the region's characteristic ecosystems may be
studied as free as possible from human disturbance-- a baseline
against which to assess the effects of human uses occurring in
other parts of the biosphere reserve or nearby region. The
biosphere reserve also includes experimental research and
demonstration areas where scientists, managers, and local people
work together to plan, test, and implement economic uses and
activities which are culturally, ecologically, and
institutionally appropriate.

As the region lacked a good model of biosphere reserve concepts
in practice, the participants in the St. John workshop were
interested in seeing whether one could be developed in the Virgin
Islands so that they would have a better basis for evaluating the
benefits. With this challenge in mind, the Virgin Islands
Resource Management Cooperative, with substantial support from
the U.S. National Park Service, began a multi-year research
program to lay the foundation for building a biosphere reserve
program in the Virgin islands. A hub of the research effort was
the Virgin Islands National Park, which comprises over half of
the island of St. John, and is the only designated biosphere
reserve unit in the Lesser Antilles. The program's scope,
however, encompassed a much larger cultural and ecological region
including all of St. John and parts of the adjacent British
Virgin Islands ecosystem.

it was recognized from the start that building the biosphere
reserve "model'' would depend upon the cooperation of the park,
its neighbors and resource users, the region's community of
scientists and managers, and at least some measure of technical
assistance from outside the region. While the Virgin Islands
National Park had a large stake in building the model and
providing some institutional leadership, it could not conduct the
experiment or construct the edifice alone. Therefore a unique
vehicle was developed -- the "Virgin Islands Resource Management

Cooperative" which provided the means to marshall the technical
capabilities of more than a dozen local and regional agencies and
organizations concerned with conservation, research, and economic
development in the U.S. and British Virgin Islands and Puerto
Rico. Locally known by the acronym VIRMC, it has provided the
framework for carrying out research requiring interagency,
interdisciplinary, and transborder cooperation.

During the first five years of the VIRMC biosphere reserve
baseline research program, Dr. Edward L. Towle, President of
Island Resources Foundation (a charter member of VIRMC) served as
program manager, while Island Resources Foundation served as
prime contractor for VIRMC, coordinating 31 separate projects
which were primarily subcontracted to other active members of the
Cooperative. In addition to the financing support received from
the U.S. National Park Service, various projects received direct
or in-kind support from the College of the Virgin Islands,
Eastern Caribbean Natural Areas Management Program, Virgin
Islands Division of Fish and Wildlife, the New York Botanical
Gardens, the U.S. Forest Service, the U.S. MAB Project
Directorate on Biosphere Reserves, the West Indies Laboratory,
Yale University, and Island Resources Foundation.

This body of work reflects' unprecedented institutional
cooperation and has stimulated interest in biosphere reserves in
the region. in 1987, the National Park Service dedicated the
Virgin Islands Biosphere Reserve Center on St. John and is using
the new facility as a focus for scientific cooperation, training,
and interaction with the local community. The United Nations
Environment Programme, in cooperation with UNESCO, has approved a
project to assess the feasibility of building a Lesser Antillean
network of biosphere reserves. The project will be coordinated
through the Caribbean Conservation Association. The Virgin
Islands experience and the specific VIRMC recommendations on
institutionalizing the MAB Program in the Virgin Islands will be
important considerations.

This final synthesis volume is the twenty-ninth in the VIRMC
series. It summarizes what has been learned during the past five
years about ecosystems and socioeconomic environment of a cluster
of small tropical islands. It was prepared by Dr. Caroline
Rogers, research biologist at the Virgin Islands National Park
and executive officer of VIRMC, and Robert Teytaud. Dr. Rogers
has been an enthusiastic proponent of the biosphere reserve
approach, a principal investigator on several projects, and a key
architect of the MAB program.

It is our hope that this synthesis and other preceding reports in
t e VIRMC biosphere reserve research report series, will be used
b-y policymakers, scientists, managers, residents and other
resource users. They should lead to the development of practical
demonstrations of the value of biosphere reserve programs in

vii

understanding small island ecosystems and in establishing a
productive marriage of conservation and economic development in
the Virgin islands and other islands of the Lesser Antilles.

Dr. William P. Gregg, Jr.
U.S. National Park Service.

viii

INTRODUCTION

The history of research on St. John is a long and
fascinating one which encompasses early observations by Danish
settlers (Carstens c.1741) and Moravian missionaries (Oldendorp
1770). The following passage appears in Highfield & Barac's 1987
translation of Oldendorp (1770) and provides an example of some
of the earliest recorded observations of the island's animals:

The green turtle has a frog-like mouth, a short neck,
and four short feet, or paws, which are suitable both
for swimming and for moving on land. The hind feet are
shaped like a goose's foot, or like a flattened hand.
The front ones, however, are narrower and longer and
are shaped almost like wings. The entire body of the
creature is covered up to its neck with a hard shell
which consists of many-sided designs and encased by a
narrow, notched border ... Its head is decorated with
small squares, just as is the underside of the body,
where the shell, however, is soft in the middle. Its
throat is of a greenish color, the rest yellowish and
brownish. The head, as well as the legs, can be drawn
through openings located on both sides of the shell,
front and back. In that position, they are concealed
from any danger. They are from three to nine feet in
length and weigh from one hundred to eight hundred
pounds.

Since the 18th century, numerous reports by territorial and
federal government agencies, universities, private organizations,
and individuals, and publications in professional scientific
journals have described the natural resources of the island. The
nature of the research currently required for management of the
Virgin Islands Biosphere Reserve on St. John depends on building
on earlier research and multi-disciplinary and multi-
institutional cooperation. Since late 1983, an extensive amount
of research, primarily on marine resources, has been carried out
in the Virgin Islands under the auspices of the Virgin Islands
Resource Management Cooperative (VIRMC). The National Park
Service funded 30 projects which focused on Virgin Islands
National Park and Biosphere Reserve (VINP/BR) on St. John, U.S.
Virgin Islands (USVI) but included collection of data from the
British Virgin Islands (BVI) and Buck Island Reef National
Monument (BUIS), off St. Croix, USVI. The VIRMC studies of St.
John's marine and terrestrial ecosystems represent the most
intensive research effort undertaken for a protected area in the
Eastern Caribbean.

The Virgin Islands Resource Management Cooperative was
founded in 1982. Signatories to the Memorandum of Understanding
are: Virgin Islands National Park, the Department of Planning

are: Virgin Islands National Park, the Department of Planning
and Natural Resources of the USVI Government (Division of Fish
and Wildlife and Division of Natural Resources Management),
University of the Virgin islands, West Indies Laboratory, Island
Resources Foundation, Eastern Caribbean Natural Area Management
Program, U.S. Geological Survey, U.S. Fish and Wildlife Service,
Southern Forest Experiment Station, University of Puerto Rico
(Sea Grant Program and the Center for Energy and Environment
Research), Caribbean Fishery Management Council, the Ministry of
Natural Resources and Labor of the British Virgin Islands
Government, and the British Virgin Islands National Parks Trust.

Some of the VIRMC research has important implications for
resource management in VINP and for other areas in the Eastern
Caribbean. To date there has been no overall synthesis of
research with resource management implications for park managers
or others who have responsibility for managing Caribbean islands
or for scientists interested in studying terrestrial and marine
ecosystems of St. John or similar ecosystems in the Caribbean.

This synthesis/summary document has many objectives:
1) to present research which has been carried out by VIRMC and
other investigators in Virgin Islands National Park as well as
the extent of our knowledge of the marine and terrestrial
ecosystems within this protected area;
2) to identify areas where future research is required;
3) to synthesize information from selected documents with
implications for research and resource management within the
Caribbean and specifically within VINP/BR;
4) to present management recommendations for ecosystems within
VINP and the wider Caribbean; and
5) to provide a basis for making the VIBR more effective in
linking conservation and development (UNESCO Action Plan for
Biosphere Reserves 1984).

The report is organized into the following major sections:
marine Systems, Fisheries, Terrestrial Systems, and Geology of
St. John. Dr. Caroline Rogers, Research Biologist for VINP, was
largely responsible for the sections on marine systems and
fisheries, as well as overall organization of the report, while
Robert Teytaud was primarily responsible for the sections on
terrestrial systems and island geology.

ACKNOWLEDGEMENTS

Thanks to R. Boulon, J. Dammann and M. Goodwin for help with
the Fisheries Section, R. Norton for writing some of the section
on birds, and D. Rankin for supplying photographs and portions of
the text on geology. The following also provided constructive
comments: Dr. B. Brown, Dr. W. Gregg, A. Reilly, A. Webster, and
C. Weikert. Larry McLain and Jean-Pierre Badle assisted with

illustrations and figures. Lito Valls reviewed the section on
history. Ian Jones worked on the bibliography and Sandra Tate
handled the final assembly of the report. Thanks to all!

PHYSICAL SETTING AND CLIMATE

The Virgin Islands, with the exception of St. Croix, are a
part of the Puerto Rico Bank, a submerged plateau usually defined
by the 183 m depth contour, extending from eastern Puerto Rico to
Anegada. The islands are high parts of the plateau which extend
above sea level. Geologically the Virgin Islands are part of the
Greater Antilles, of which they form the eastern terminus, rather
than of the Lesser Antilles with which they are commonly grouped
culturally. Ocean depths between the northern Virgin .Islands and
Puerto Rico are generally less than 91 m, but the Puerto Rico
Bank itself is surrounded by steep slopes and deep water. The
Puerto Rico Trench with depths to 9,166 m lies to the north, the
Virgin Islands Basin with depths of 4,500 m to the south, and the
St. John and Anegada Passages with depths of 2,000 m to the east.

The island of St. John (18o 18'-18o 22'N; 64o 40'-64o 48'W)
lies in the Caribbean Sea about 88 km east of Puerto Rico (Fig.
1, Appendix I) and covers an area of about 48 km2. Its
topography is very rugged, with deep valleys and hills that rise
steeply from the shore. Over 80% of the slopes exceed 30% (CH2M
Hill 1979a). The highest point, Bordeaux Mountain, is 387 m
above sea level. There are numerous embayments along the
coastline which has a total length of about 52 km. St. Thomas
lies to the west and several of the British Virgin Islands are to
the north and east, across channels 1.5 to 5 km wide. St. Croix
is about 64 km to the south. On the northeastern shore of St.
Croix across a shallow channel 2.6 km wide, one finds Buck
Island, a National Monument managed by the National Park Service.
The island has an area of 71 ha, and its highest point is about
100 m above sea level.

Winds

The Tradewinds (Easterlies) consistently approach the
islands from the east. Changes in the position and intensity of
the Bermuda High to the north and the Equatorial Trough to the
south are responsible for major seasonal variations. The winds
blow counterclockwise around the Equatorial Trough, a low
pressure zone.

When the Bermuda High is intensified from December through
February with only minimal pressure changes in the Equatorial
Trough, the Tradewinds are at a maximum, blowing 11-21 knots
about 60% of the time (Towle et al. 1977). During this period,

700 650 60�

ATLANTIC OCEAN

Dominican Republic

Anegada

Puerto Rico Tortola
-_ f Virgin Gorda
St. T John ,hn Angullla
_ St.;Thomass 4 St. Martin
S St. Barthelemy
Saba
St. Croix Barbuda
St. Eustatius *
St. Kltts
Nevis * Antigua

Montserrat &

Guadeloupe ~
Marie Galante

CARIBBEAN SEA Dominica

Martinique I

,',"~ ........-~-_ ,=St. Lucia

St. Vincent Barbados
aQ < \_ SztJ-~ tJ 4\Bequia ,
Hawksn'est Bay c C
_i c.~ c,.... �-1.. ~: 4/ VL (- .' The Grenadines
Ii J ST JOHN ; t IJ J "~ . Carrlacou ;

Grenada

Fish Bay

,___.____<~~~~~~~~~~~~~~~ 0 fsy= ;Tobago

""~ ! ~ sTrinidad

Map of St. John, USVI, showing locations of study sites. -10
Broken line indicates boundary of Virgin Islands National Park.

NAUTICAL MILES

I I I I I I
o 50 100 150 200 250 300

Fig. 1. Location of St. John, U.S. Virgin Islands.
From: Putney (1982).

the "Christmas winds", strong winds from the north, may appear as
a result of weak cold fronts moving through the Caribbean. A
decrease in the pressure of Equatorial Trough results in a
reduction in wind velocities from March through May. From June
through August, wind speeds approach those in December through
February as the Bermuda High strengthens with a concomitant
decrease in pressure in the Equatorial Trough. The lowest wind
velocities occur from September through November as the Bermuda
High weakens with only a small decrease in pressure in the
Equatorial Trough.

Although hurricanes have occurred in every month of the
year, most storms pass through the Eastern Caribbean between
August and October with peak activity in September. Twenty-four
hurricanes have passed within 50 miles of the Virgin Islands
since 1900 (U.S. Army 1975). Major hurricanes occur about once
every 33 years.

Rainfall

National Park Service records for Cruz Bay, St. John, from
1872 to 1986 indicate an average annual rainfall of 44 inches
(112 cm) and a range of 23 to 74 inches. Rainfall varies
considerably not only from year to year but from month to month.
Although most rain typically falls from September to November,
occasional storms, particularly in the Spring, can bring large
amounts of rain. One storm in April 1983 dropped 15.6 inches of
rain over a 24 hour period. Cosner (1972) estimated that
evapotranspiration accounts for over 90% of the rainfall.

Waves and Currents

The mean tidal range in the islands is only 0.8-1.0 ft.
(0.24-0.30 in), and tidal currents are usually weak (Towle et al.
1977). Diurnal tides predominate around St. Croix and on the
south coasts of St. John and St. Thomas. Semi-diurnal tides
occur on the north coasts of Puerto Rico, St. Thomas and St. John
because of exposure to open water. Waves come primarily from the
east in the winter and the southeast in the summer, in response
to seasonal shifts in wind direction. Waves are 1-3 ft. (0.3-0.9
m) about half the time. southeasterly swells, occasionally
reaching 10-12 ft. (3-3.7 m) nearshore, develop in winter. The
northern swells create strong longshore currents. Wave energy is
concentrated on projecting headlands, and shoreline configuration
reflects variations in the resistance of specific rock types to
erosion in these headlands.

BRIEF HISTORIC OVERVIEW

Archaeological finds at Lameshur Bay, St. John, provided
evidence that seafaring Arawaks occupied the island as early as
770 B.C. (R. Reeves pers. comm.). These aborigines practiced a
mixed subsistence economy that involved harvesting of shellfish
and other marine organisms and some cultivation of manioc along
with other foodstuffs. Most of the known settlement sites as in
most of the Eastern Caribbean, are near the coast (Tyson 1987).

In 1493, Christopher Columbus claimed St. John for Spain
during his second voyage to the New World. Columbus' men saw no
human beings on the island but there is some evidence that the
island was uninhabited from sometime before 1550 for over 100
years, and between 1671 and 1717 occupied only intermittently.

Colonization of St. John began in March 1718 as Danish
authorities parcelled out land for plantations. All of the
island, with the exception of some of the rockiest headlands, had
been subdivided into plantations by 1728 (Larsen 1986). The
plantation economy depended on the importation of slaves from
Africa and other Caribbean colonies.

After 1765, the plantation system, which had been based on a
diversity of crops including sugar cane, cotton, coffee and
indigo, gave way to an economy based primarily on sugar. This
economy was undermined after 1840 by loss of soil fertility,
rising costs, falling prices and the 1848 emancipation of the
slaves. Between 1805 and 1915, pressure on the island's
resources declined substantially as the population fell from 2577
to 905 (Tyson 1987). The last commercial sugar production was at
Par Force plantation in Reef Bay Quarter in 1897 (Fig. 2). After
this, sugar was produced for local consumption, only. By 1915,
much of the island had reverted to secondary forest. The
plantation system gave way to small scale agriculture as peasants
raised livestock, grew fruit, and harvested and processed bay
leaves (Tyson 1984, Olwig 1986). A distinct Afro-Caribbean
culture evolved around these agricultural pursuits, fishing, boat
building, and charcoal production (Olwig 1986).

The United States took possession of St. John in 1917,
leading to an economy based on tourism. Much of the old
plantation lands were incorporated into Virgin Islands National
Park which was established in 1956 with donations of land by
Laurance Rockefeller. The park occupies approximately 50% of the
island. The island's fascinating history has resulted in a
dynamic, cultural landscape which consists of a mosaic of
protected land and private and public lands which are rapidly
being developed.

Fig. 2. Great House at Par Force Plantation, Reef Bay Quarter
(1988). Photo: S. Barry.

MARINE SYSTEMS

HISTORY OF RESEARCH

Kumpf & Randall (1961) produced the first map of the benthic
communities around St. John in 1959, showing the location of the
following bottom types: coral or rock bottom, coral patches on
sand, seagrass, sand or coral rubble, and mud (Appendix II).
From 1958 to 1961, several studies focused on fish taxonomy and
ecology (listed in Randall 1961). Randall established an
artificial reef in Little Lameshur Bay in 1960 (Randall 1963).
The Tektite I and Tektite II underwater habitat projects took
place in Greater Lameshur Bay from 1969 - 1971 (Collette & Earle,
1972, Earle & Lavenberg 1975). These studies included research
on lobsters (Cooper et al. 1975, Herrnkind et al. 1975, Olsen &
Koblic 1975, Olsen et al. 1975), algae (Earle 1972, Mathieson et
al. 1975), fishes (Collette & Talbot 1972, Clifton & Hunter 1972,
Smith & Tyler 1972) and coral reef biomass (Lee et al. 1975).

Alan Robinson, the VINP scientist from 1970 to 1975,
reported on recreational uses of the park (1973, 1976), salt
ponds of St. John (1974), the soft bottom community in Caneel Bay
(1972), and briefly summarized research in the park (1975).
Small (1982) reported on sea turtle nesting within VINP and BUIS.
Faculty and students (for example, from the School for Field
Studies and Gustavus Adolphus College) based at the Virgin
Islands Ecological Research Station, which is operated by the
University of the Virgin Islands, have produced numerous reports,
primarily on marine organisms in the Lameshur Bays. Studies of
marine areas outside of park waters include those by Grigg (1978)
on Cruz Bay, Brody et al. (1969, 1970) on Chocolate Hole and Cruz
Bay, and by EPA (1988) on Turner Bay and in surrounding waters.
Some abstracts on marine research in the USVI can also be found
proceedings of meetings of the Association of Island Marine
Laboratories in the Caribbean (e.g. AIMLC 1986).

Little research has taken place on physical oceanography.
Dammann (1969) presents information on water temperature,
salinity, transparency, and currents around the USVI. Towle et
al. (1977) prepared a compendium on oceanography, seismic
activity, and climatology in the islands. VanEepoel et al.
(1971) present detailed oceanographic data and statistics on
tropical storms and hurricanes (see also Bowden et al. 1974).
Water quality data (dissolved oxygen, turbidity, temperature,
salinity, and other parameters) for St. John's waters from 1973
to 1986 are available from the Dept. of Planning and Natural
Resources (formerly Dept. of Conservation and Cultural Affairs).

The most comprehensive research on marine systems has taken
place under the auspices of the Virgin Islands Resource

Management Cooperative, beginning with studies in 1983. Phase I
of the VIRMC studies focused on obtaining baseline data on
critical benthic habitats and fisheries around the island. Phase
II studies emphasized an interdisciplinary approach to selected
watersheds on St. John and the initiation of long-term monitoring
of some of the ecosystems. Some Phase I and Phase II projects
concentrated on areas in the BVI which potentially could join the
VIBR to form a larger, yet to be designed, international
biosphere reserve. In addition, two studies examined zoning and
management of the Virgin Islands Biosphere Reserve. The third
Phase included studies of the consequences of increasing
recreational uses of marine resources, further work on long-term
plots established in St. John' s forests and watershed processes,
an analysis of Buck Island fisheries, and compilation of this
synthesis document (see Appendix III).

DESCRIPTION OF MARINE SYSTEMS

Environmental conditions on small Caribbean islands such as
St. John are conducive to development of closely interrelated
mangrove, seagrass, and coral reef systems (Towle et al. 1977).
Fish and shellfish depend on all of these systems for food and
shelter and link them to each other through their migrations and
the resultant transfer of nutrients. Some fishes migrate daily
from the shelter of the reefs to feed in adjacent seagrass beds
at night. The seagrass beds are therefore subsidizing the reefs
in terms of energy flow. Seagrass blades (particularly
Syringodium) float for days after detachment by rough seas or by
grazing animals such as parrotfishes and sea turtles. Drifting
over reefs and seagrass beds, they are an important source of
nutrients.

Juveniles of many organisms depend on mangroves and seagrass
beds, moving to deeper waters and offshore reefs as they mature.
Destruction of red mangroves with submerged prop roots decreases
the habitat for juvenile fishes (Thayer et al. 1987). Cutting of
mangroves which entrap sediments can result in excessive
siltation for nearby seagrass beds and reefs from runoff after
heavy rains. Runoff can be especially severe where there has
been careless shoreline development or poor agricultural
practices.

Fish can affect these ecosystems in a number of ways, some
of them very complex. They can affect reefs by 1) grazing on
dead coral, breaking down the skeleton and producing large
quantities of sediment, 2) providing nutrients in their wastes,
and 3) grazing on coral larvae, thereby influencing the reef's
structure. Randall (1974) suggested that the continuous rain of
sediments from parrotfish and surgeonfish feces adversely affects
reef organisms. However, fish feces may enhance coral growth

(Meyer et al. 1983) and supply essential nutrients (Szmant-
Froelich et al. 1981). In a recent synthesis of studies of the
effects of fish grazing on coral reef structure, Hixon (1986)
noted that damselfish which defend algal territories directly
influence coral recruitment and growth, as well as bioerosion.
Numerous studies demonstrate the effects of fish grazing on reef
algae (e.g., Ogden & Lobel 1978). Economically important fish
(e.g., snappers, grunts) and conchs feed on the benthic
microalgae and invertebrates in reef sediments. Most fish seek
coral colonies for shelter from predation, and a few species eat
living coral. Fish eat other sessile animals of the reef
including sponges.

Mangrove forests are not extensive on St. John because in
many places the island's steep slopes extend down to the water's
edge, and flat land for mangrove growth is limited. The absence
of rivers or permanent streams reduces nutrient input for
mangrove development, but the resultant water clarity also favors
development of coral reefs close to shore. Seagrass beds occur
in shallow waters which receive protection from heavy seas and in
deeper water seaward of the reefs.

Coral Reefs

Most of the reefs around St. John are shallow fringing reefs
which grow near shore or extend out from the headlands at
entrances to the numerous bays which indent St. John's coastline
(Fig. 3). Many of the reefs near shore slope down to about 12 m
where there is an abrupt transition to a relatively barren, sandy
plain. Scattered patch reefs with high coral diversity occur on
geological features offshore at greater depths. True, extensive
barrier reefs with well-defined lagoons do not occur around the
island, although the reef in Newfound Bay could be considered a
combination of a fringing and a barrier reef. Bank reefs which
do not break the surface occur offshore. Isolated patch reefs
can be found in Hawksnest Bay, Reef Bay and elsewhere. Elkhorn
coral (Acropora palmata) is the most abundant species on the
shallow reefs. Brain corals (Diploria spp.), lettuce corals
(Agaricia spp.), and fire corals (Millepora spp.) are scattered
throughout the elkhorn framework.

Many of the reefs are true coral reefs established on a
framework of coral skeletons deposited over thousands of years.
However, St. John has many coral communities comprised of coral
colonies and other reef organisms growing on submerged boulders
and rock ridges near shore such as occur at Leinster Bay.

Beets et al. (1986) produced 16 maps, based on aerial
photographs and field observations, and described all of the
island's bays. Each of their maps indicate major ecosystems and
their subzones (Fig. 4). They observed a total of 36 hard coral
species around St. John. Beets & Lewand (1986) collected over

Fig. 3. Johnson's Reef at a depth of 60'.
Photo: C. Weikert.

864� aMarine Benthic Communities

St. John, USVI

Bs N

0 100 200 300

Scale in Meters
Profile 3.6
Aer i a I Photo9 rap hs, NOS 1983

Rfp
Br ~~~Ra ;>mgoat~R >iB

SSR ss~~~~~~~~~~~~~~~~~~~R

ZX~~~~ ~~~~~~~~ gAW\ A~~Peace Hill
/(//. RP / Hawksnest

Pofile 37

Hawksnest Bay

S..

Fig. 4. Map of major benthic communities, Hawksnest Bay.
SBg-$$

From: Beets, et al. 1986.

12
RII, ~ ~ 1

500 specimens of common marine organisms within VINP to
supplement park collections by J. Randall, A. Dammann, and others
in the 1960's. Hubbard et al. (1987) and Rogers & Zullo (1987)
described reefs in Hawksnest, Fish, and Reef Bays. Dammann
(1969) provided descriptions of reefs in Chocolate Hole and
Mary's Creek.

In recognition of the need for quantitative baseline data to
provide information for effective management of VINP coral reefs,
Rogers & zullo (1987) initiated a long-term monitoring program in
1984. They established transects in Reef, Fish, and Hawksnest
Bays. The watersheds for these 3 bays have different development
histories. Reef Bay has remained undisturbed since the decline
of the sugar plantation there in the late 1800's. The Hawksnest
watershed is the site of a new medical clinic. The Fish Bay
watershed is currently undergoing subdivision, roadbuilding, and
construction of houses. Although the lands are outside the park,
the study reefs are in park waters and could deteriorate with
increased siltation from erosion associated with rapid
development of this area. Tropical Storm Klaus in Nov. 1984
resulted in a statistically significant decrease in the mean
percent cover of the dominant coral Agaricia agaricites at the
study site in Fish Bay. Future measurements along these
transects should allow tracking of their condition over time.
Hubbard et al. (1987) studied reefs in these three bays as well.
(See discussion of their work under Sedimentation).

Seagrass Beds

Seagrass beds are highly productive ecosystems which are
extensive in the shallow waters around the USVI. Turtle grass
(Thalassia testudinum), manatee grass (Syringodium filiforme),
and shoal grass (Halodule wrightii) are the most abundant
species. Seagrasses are true flowering plants, spreading
primarily through growth of roots and rhizomes. These extensive
root systems help to stabilize sediments and reduce shoreline
erosion. Calcareous algae such as Halimeda and Penicillus, which
have calcium carbonate or limestone in their tissues, grow in the
grass beds and surrounding sand bottoms. Most beach sands are
primarily composed of particles which arose from the
decomposition and disintegration of these plants.

Several species of commercially important fishes, such as
snappers and grunts, find shelter in coral reefs during the day
and feed in the seagrass beds at night (Fig. 5). Seagrass beds
are the primary habitat for the commercially important queen
conch (Strombus gigas) which feeds on algae growing on the grass
blades. The food web in grass beds is based both on direct
harbivory (grazing) on grass blades and algae, and on detritus
from the decomposition of these plants. Conspicuous halos or
bare sandy zones around coral patch reefs within grass beds are

Fig. 5. School of grunts sheltering in a patch reef off Scott
Beach. Photo: L. McLain.

the product of physical factors like currents but also of grazing
by fishes and sea urchins (Ogden et al. 1973, Randall 1965).

Seagrass beds around St. John are not as extensive as those
around St. Thomas and St. Croix. Beets et al. (1986) mapped and
briefly described seagrass beds in many park bays while
Williams(1988) mapped seagrass beds in Francis and Maho Bays.
None of the northern bays have dense seagrass beds, and the best
beds occur in Rendezvous and Reef Bay on the south side of the
island.

Examination of aerial photographs from 1975 to 1983 suggest
a serious decline in the seagrass beds in Francis and Maho Bays.
Williams (1988) noted that anchors from the increasing number of
boats visiting these bays are ripping up seagrass roots and
rhizomes. (See also Rogers et al. 1988 and Recreational Uses
section below.) The population of 50 individuals of the
threatened green sea turtle Chelonia mydas grazes heavily on the
turtle grass in these bays. Thalassia plants inside small fenced
areas (4 m x4 m) which excluded turtles showed some recovery
after 3 months, indicating that recovery or at least healthier

growth of these plants can occur. (Anchoring was also prohibited
in the plots.)

Williams (1988) noted that the stressed seagrasses in
Francis and Maho had extremely low productivity and fragile,
narrow leaves. She recommended prohibition of anchoring in these
bays. Anchors may be partly responsible for deterioration of
Thalassia beds in Caneel Bay (Beets, personal communication),
although effluent from a desalination plant may have caused most
of the damage. Robinson (1972) previously described the
seagrasses in Caneel as "extensive".

Mangroves

Mangroves are very productive ecosystems which support a
high diversity of fish, birds, and other wildlife. The mangrove
food web, based largely on the release of nutrients from the
decomposition of mangrove leaves, supports nearshore fisheries.
Mangroves are vital feeding, nesting and roosting areas for
several species of birds including ospreys, pelicans, egrets,
herons and ducks. Many of the species are rarely seen outside
this habitat.

Red mangroves (Rhizophora mangle) grow along the shoreline
often in monospecific stands. In well-developed forests, red
mangroves form the seawardmost zone, and black (Avicennia
germinans) and white (Laguncularia racemosa) mangroves grow
behind the red mangroves. The red mangroves are the most
distinctive of the three major tree species, with their
characteristic aerial prop roots (Fig. 6). These roots are
usually partially submerged and support a diverse community of
sponges, ascidians, algae and sometimes even corals. Juvenile
reef and pelagic fishes as well as lobsters are abundant in these
root communities (Thayer et al. 1987). The roots also trap
sediments and over a period of several years can extend the
shoreline seaward. Mangroves can reduce the amount of runoff
(and associated sewage and pollutants) which reaches offshore
seagrass beds and coral reefs.

Two of the four major physiographic types of mangrove
forests described by Cintron & Schaeffer-Novelli (1983) occur in
St. John. Fringe forests are found along protected shores such
as those in Coral Bay, on shoals or spits, or around the margins
of some salt ponds. In such areas steep gradients in topography,
salinity, turbulence and tidal amplitude occur over a short
distance. The inner parts of the fringe are subject to less
variation in tidal amplitude and turbulence, and soil elevations
ate higher resulting in less flooding by seawater.

Fig. 6. Prop roots of red mangroves are important nursery areas
for fish. Drawing: M. Beath.

Basin forests develop where there are sheet flows of water
over areas of very small topographic gradients, such as flat
areas where guts reach the coasts, and drainage depressions.
Basins receive and store water annually and do not develop strong
salinity gradients. Best development occurs in areas of weak but
constant flows, whereas forests in stagnant basins have reduced
growth. Examples of this type of forest can be found at Mary's
Creek, Reef Bay, or Fish Bay. Cintron (1987) discussed the
adaptations of mangroves and other systems to episodes or
"pulses" of storms and provided management recommendations for
mangrove ecosystems.
Salt Ponds

Most salt ponds are formed when mangroves or fringing coral
reefs grow across and isolate a portion of a bay (Fig. 7).
Hurricanes and other storms also can pile up coral fragments to
create barriers to the sea. St. John has excellent examples of
the many stages of this process (Robinson & Feazel 1974). Large
ponds include Southside Pond on the southeast end of the island,
Francis Bay salt pond, and Mandal Pond. At Newfound Bay a salt
pond may form adjacent to the existing one if the reef continues
to grow across the mouth of the bay.

Fig. 7. Salt pond on St. John. Photo: L. McLain.

These ecosystems support a specialized biota which varies
with fluctuations in pond water salinity. Rapid evaporation
results in extremely saline water, and salt crystals sometimes
form along the shores of the pond. The most saline ponds support
a less diverse fauna than those ponds where storm waves breach
the berm and create channels which allow seawater of normal
(lower) salinity to rush in. Barracudas, mullets, and crabs can
be found in ponds which are open to the sea.

Mangroves frequently fringe the edges of these ponds, and
many of St. John's birds are associated with them. Several
species of insect-eating and fish-eating birds, including
kingfishers, herons, ospreys, stilts, and sandpipers feed on
insects, brine shrimp, and fishes in these ponds.

Many salt ponds situated between an upland watershed and its
associated bay function as settling or catchment basins, trapping
the runoff from the land. Especially in areas where watershed
development has occurred, the ponds reduce the amount of silt-
laden water which reaches the bay, contributing to maintenance of
high water quality. Analysis of salt pond sediments can provide
insights into historical land use in a watershed (Nichols & Brush
1988).

Algal Plains and Marine Algae

Algal plains around St. John have not been well-studied.
Beets et al. (1986) note this habitat on several of their maps
for St. John. In her study of herbivory in Great Lameshur Bay
carried out from the underwater habitat Tektite, Earle (1972)
listed 154 species of plants and 35 species of plant-eating
fishes, many of them associated with the algal plain near the
habitat. Earle concluded that herbivorous fish had a greater
influence than invertebrates on plant distributions and
abundance. In the "Marine Algae of the Danish West Indies"
published from 1913-1920, Borgesen described 327 species of
algae.

Beaches and Rocky Shores

St. John's beaches are composed primarily of calcium
carbonate sands formed from fragments of corals, shells, and
algal particles. Some beaches in high-energy environments are
composed mostly of large chunks of coral torn loose by the waves.
Beach materials of terrigenous origin typically consist of light-
colored quartz or feldspar sand, or larger dark-colored gravel,
cobbles, and boulders. Terrigenous materials are either eroded
from cliffs at the shoreline or from inland soils, and
transported to the shore by runoff. Many beaches also have
ledges composed of beachrock which run parallel to the beach

face, either exposed or buried under a layer of sediment.
Beachrock forms fairly rapidly below the beach surface by a
natural process in which calcium carbonate precipitates out of
seawater, cementing together sediment particles.

Beaches are constantly changing under the driving influence
of wind, waves, tides and currents. Seasonal and longer-term
changes in these variables significantly affect beach dynamics.
High energy storm waves erode beach sediments and produce a
steeply sloping foreshore and narrow berm. Beach sediments are
transported offshore and may be deposited as a bar at the limit
of the outer breakers. When more average waves return, the beach
will typically recover to near-original profiles as the less
energetic waves transport sand shoreward again. During this
exchange of sand between the beach face and offshore storage,
some of the sediments will be carried laterally along the coast
by longshore currents, some will be lost permanently to deeper
water, and some will be blown or washed across the berm to become
part of the backshore or beach ridge.

On St. John, most beaches are found along the margins of
deeply indented bays, resulting in little exchange of sediment
from bay to bay around the enclosing headlands (Fig. 8). Most
sediment transport is onshore and offshore within the bay itself,

Fig. 8. Trunk Bay Beach on St. John.
Photo: L. McLain.

and the sediment budget is more or less in equilibrium. one
exception is the beach at Cinnamon Bay, which is exposed to storm
swells from the north in winter and lacks a large headland on the
western side to trap sediments within the bay. This bay
configuration results in significant losses of sediment by
longshore transport, and beach erosion has been and continues to
be a problem. Dolan (1973) and Hoffman (1974) studied beach
dynamics at Cinnamon, Trunk, and Hawksnest Bays, Young & Young
(1968) studied sand movement at Lameshur Bay, and Hubbard (1980)
reported a study of storm-related versus seasonal changes at Buck
Island beach, St. Croix.

A number of crabs, mollusks, worms, and other invertebrates
live in the intertidal zone of St. John's beaches, and species of
ghost crabs (Ocyp2ode) and land crabs (Geocarcinus) make their
burrows above high tide. Many shorebirds visit St. John's
beaches, and sea turtles use the beaches for nesting.

Rocky shores are steep slopes, cliffs, or outcrops composed
of more or less consolidated rock substrate. They are areas of
high wave energy and constitute a severe environment for life.
Between the level of the lowest spring tides and the highest
elevation normally wetted by spray, they are inhabited by a
distinctive assemblage of animals and plants, including the
edible topshell or "whelk" (Cittarium pica) discussed below.
Some remote rocky shores are used by seabirds (such as the red-
billed tropic bird) for nesting and roosting habitat.

STRESSES TO MARINE SYSTEMS

Sediment at ion

Sedimentation from dredging and runoff probably constitutes
the biggest potential source of reef degradation in the
Caribbean. In a recent survey (Rogers 1985), resource managers
and scientists in the Caribbean indicated that siltation had
caused deterioration of reefs off Florida, Martinique,
Guadeloupe, Costa Rica, and elsewhere.

Sediments that settle on coral colonies and cause high
turbidity in suspension have sublethal and lethal effects on
corals. Dredging near coral reefs and accelerated runoff of
eroded soils increase turbidity, thereby cutting down light
available for photosynthesis and increasing sediment load on reef
organisms. Aside from the localized consequences of removal and
cutting of substrate, dredging affects extensive areas by putting
fine sediments into suspension where currents carry them to other
reef areas. Deterioration of the reef can continue for several
years after dredging ceases because of continual resuspension and
transport of dredged materials (Brock et al. 1966). Dredging has

destroyed large areas of seagrass beds particularly off Florida
(Zieman 1985).

Hubbard (1987) reviewed the effects of sedimentation on
marine systems, particularly coral reefs in the Eastern
Caribbean, and evaluated the current federal and territorial
legislation and programs to control sedimentation in the USVI.
Rogers (in prep.) reviewed the effects of sedimentation on coral
reefs at the organism and ecosystem level and discussed the ways
sedimentation can influence fish populations and consequently the
fisheries upon which they are based.

Sedimentation studies within VINP/BR

A major environmental concern on St. John is siltation of
nearshore marine habitats from accelerated erosion following
watershed development. This concern prompted many of the VIRMC
II studies. Erosion and runoff are potentially serious problems
on St. John because of its extremely steep hillsides and the
recent, dramatic increase in development.

Rapid, and in some cases, uncontrolled development on St.
John has led to increases in terrigenous runoff. After heavy
rains, bays within and outside the park have highly turbid water.
St. John residents believe that corals in Hawksnest and Cinnamon
Bays on the north shore of the island were killed from siltation
following exceptionally heavy rains in April 1983 (457 mm in 24
hr). Development of private lands within the park boundary and
development outside the boundary can contribute to turbidity in
park waters which encompass 2,286 ha around the island. New
subdivision roads continuously being carved into the hillsides
are creatingj the potential for accelerated erosion and runoff.

Using a theoretical approach, Hubbard et al. (1987)
calculated peak discharge rates and volumes assuming "natural"
conditions of total forest cover for St. John. The distribution
of reefs around the island appears to follow a pattern consistent
with the assumptions of their model. They concluded that the
primary controls of reef development are watershed size, runoff
(primarily associated with 10 - 25 yr storms), bay geometry, and
exposure and that the recent dramatic increase in the island's
development exerts less control. The location of the well-
defined "guts" through which runoff flows into the bays strongly
influences the distribution of reef organisms.

The most turbid nearshore waters are associated with high
runoff and poor flushing ability because of bay geometry and low
wave energy, e.g., inner Fish, Coral, Cruz, Great Cruz, and
eastern Hawksnest Bays. Greater exposure in western central Reef
Bay correlates with higher coral cover than the reefs further

east. Hubbard et al. (1987) were primarily examining presence
and absence of reefs in their model, and further research on reef
structural parameters, e.g., species diversity, in relation to
runoff would be worthwhile.

In their comprehensive and detailed study of Hawksnest,
Reef, and Fish Bays, Hubbard et al. (1987) briefly describe the
reefs and provide profiles showing the different reef organisms.
They present grain size characteristics and percent terrigenous
material for surface sediments and sediment cores; document
growth rates of M. annularis colonies; and provide useful
recommendations for future coastal development.

Hawksnest Bay There is less reef cover in Hawksnest than in Fish
and Reef Bays. The easternmost patch reef has less than 5%
cover, reflecting low wave action and proximity to a large gut.

Examination of annual growth rings for 12 M. annularis heads
in Hawksnest revealed a significant short-term decrease in 1981
and 1983 which coincided with heavy rainfall following the
construction in 1980 of the clinic in the upper watershed.

Fish Bay The highest concentrations of terrigenous material were
found in sediments from inner Fish Bay. Sediment cores from Fish
Bay showed higher concentrations of terrigenous matter in the
upper layers, possibly the effects of runoff associated with
recent bulldozing and road construction. On the other hand, reef
development to the southeast may reduce wave energy and trap
fine sediment in the inner area of the bay. The best coral cover
in Fish Bay occurs along forereef slopes.

Reef Bay Although Reef and Fish Bays have similar watershed and
drainage areas, Reef Bay sediments contain less terrigenous
matter, presumably because the bay has greater exposure and
circulation. As in Fish Bay, the best coral cover was found in
sloping lower forereef environments.

Coral growth Cores of large coral heads provide historical
information on water quality as smaller growth bands indicate
less favorable conditions for growth. Hubbard et al. (1987)
hypothesized that reef development (specifically coral growth)
would have been greatest following the sugar cane plantation era
and less extensive prior to and subsequent to that period,
because of adverse effects associated with land clearing.

Although the results of the coring were ambiguous, Hubbard et
al. (1987) were able to document a 10 - 20% reduction in growth
between the post-plantation era and the present in 5 separate
cores through M. annularis colonies. The growth rates were
consistent with t-hose found by others (Dustan 1975, Gladfelter et
al. 1978, Hudson et al. 1976, Hubbard & Scaturo 1985). The only
other coral growth rates for St. John appear to be those for A.

palmata and A. cervicornis reported in Feazel (1975).

For all three bays, Hubbard et al. (1987) found a gradual
reduction in coral growth over the last 100 - 200 years (based on
limited data from coral heads). They suggest the decrease in
growth may be a response to the intensive agriculture of the
1800's, but the growth decline in some cases corresponded to the
period following intensive cultivation. Recovery of the forests
may have been accompanied by a decrease in the ability of the
land to hold and retain sediment and water as more extensive
vegetation (e.g., grasses) were shaded out by trees. Breakdown
of the terracing systems from the plantation era could also lead
to an increase in near shore siltation.

When trying to analyze effects of historical and current
land clearing practices it should be remembered that modern
practices of clearing land with bulldozers that can remove
topsoil down to underlying bedrock and completely uproot trees
are potentially far more destructive than past methods.

Effects of sedimentation on coral metabolism Porter & Rogers (in
prep.) studied the effects of terrigenous sediments on oxygen
production and respiration by A. palmata, D. strigosa, and M.
annularis colonies at Henley Cay (west of St. John) using a self-
contained chamber equipped with oxygen probes and stirrers.
Different concentrations of sediments were applied to coral
colonies to determine reduction in photosynthesis and stimulation
of respiration resulting from this stress.

Sedimentation rates Nichols & Brush (1988) studied Reef Bay
mangrove swamp and Mandal Pond on the south coast of St. John to
examine the relationship of land use history in the watershed to
natural sedimentation rates from erosion. Using sediment cores,
they found that the deposits effectively provided a historical
record of sedimentation rates in the two ponds. In spite of
extensive clearing of the vegetation in these areas during the
plantation era, Nichols & Brush (1988) concluded that human
activity had not significantly altered the natural processes of
sedimentation. At Reef Bay, in contrast to Mandal Pond,
infilling is exceeding submergence.

Recreational Uses

Throughout the Caribbean, tourism and coastal development
are exerting severe pressure on the natural resources of many
islands and countries. More and more tourists are visiting parks
and protected areas in the region. Robinson (1973, 1976) drew
attention to environmental damage associated with recreation in
VINP in the 1970's prior to the dramatic increase in visitation.
Rogers et al. (1988) recently documented the trends in
recreational uses of the waters and beaches of VINP and assessed

RECREATIONAL VISITORS
1957-1987
800

700 -

600 -

800 -

400-

300 -

200 -

100 -

0-
57 59 61 63 65 67 69 71 73 75 77 79 81 83 85 87
YEAR

Fig. 9. Recreational visits to Virgin Islands National Park from
1957 - 1987.

BOATS IN PARK WATERS

so90

70 -

80 -

40-

30-

20 -

10 -

0-
67 69 71 73 75 77 79 81 83 85 87
YEAR

Fig. 10. Boats in park waters from 1966 to 1987.

the degradation of the marine resources attributable to
recreational activities (exclusive of fishing).

Recreation in the park has increased dramatically in the
last 8 - 10 years. Visits to the park have risen from less than
100,000 prior to 1967 to almost 800,000 in 1987 (Fig. 9). Annual
visitation to Trunk Bay beach, the most heavily used beach in the
park, has risen from under 20,000 people in 1966 to almost
170,000 in 1986. The average number of boats per day in park
waters increased from less than 10 in 1966 to about 80 in 1986
(Fig. 10).

Some of the coral reefs and seagrass beds along the north
shore of St. John which receives the heaviest use have suffered
degradation. Anchor damage and damage from boats striking or
grounding on reefs is evident. Seagrass beds in Francis and Maho
Bays have deteriorated (Williams 1988).

Anchor damage

Between January and March 1987, Rogers et al. (1988)
examined anchor damage in northern and northwestern bays of St.
John, including Francis, Maho and Leinster. Of the 186 boats
surveyed, 32% were anchored in seagrass and 14% in coral
communities with the remainder on sand, mud, or pavement. In
general, the deeper the anchor, the less damage it was causing as
sensitive coral and seagrass communities tend to occur near shore
in most of the bays surveyed. An estimated 30,000 boats per year
anchor in park waters. Mini cruise ships often set 2 anchors,
each weighing I ton. Anchor chains are often more devastating
than anchors themselves because they scour out a large area of
the bottom as a boat shifts around with wind and current changes.

Physical damage to Windswept and Hawksnest Reefs

Monthly observations were made at Windswept and Hawksnest
Reef from Feb. 86 through April 1987 to record physical damage to
these especially vulnerable sites from anchors, boat groundings,
careless snorkelers, or other causes. Installation of marker
buoys in may 1986 to warn boaters of the presence of these reefs
resulted in a marked decrease in damage from boats. While heavy
swells can fracture fragile coral branches, boat groundings can
do more damage, in some cases even cracking the reef's framework.

Recognition of the severity of damage associated with
increased recreational uses of marine resources in the Caribbean
is growing. Tilmant (1987) recently reviewed adverse effects of
recreation on coral reefs, citing examples from Australia and
Florida. In a survey of stresses affecting Caribbean reefs, many
resource managers and scientists reported damage from anchors
(e.g., from cruise ships and dive boats), from boat groundings,
and from people walking on reef flats and removing corals as

souvenirs (Rogers 1985). Construction of marinas and boatyards
to support recreational boating have had serious environmental
consequences.

Damage from recreational activities is superimposed on
destruction from natural causes. VINP Research staff tagged and
monitored 50 elkhorn colonies in Hawksnest Bay in July 1987 and
found that only 10 of these corals appeared undisturbed 7 months
later. Eleven colonies had been completely detached from the
substrate apparently during heavy seas. Other tagged corals
exhibited breakage, algal growth, or signs of predation.

In response to increased visitation and degradation of the
park's marine resources, VINP developed a shoreline use plan in
1987. The plan calls for establishment of mooring buoys to
protect sensitive seagrass and reef areas, and seeks to regulate
use of park beaches and waters by commercial tour groups. Looe
Key National Marine Sanctuary in Florida has a very effective
mooring system (Halas 1985). Billy Causey, the Sanctuary
Manager, concluded that "the installation of the buoys has been
the most beneficial effort that we could have undertaken to
protect our reefs". Similar moorings will be installed in VINP.

Oil Pollution

Cintron et al. (1981) reviewed the impact of oil pollution
on the structure and function of tropical marine systems.
Grounding of vessels often results in release of oil near reefs,
mangroves and seagrass beds. The environmental effects can be
dramatic, or sublethal and delayed (Birkeland et al. 1976).
Further research is required on toxicity of dispersants and
oil/dispersant mixtures. Use of dispersants can actually
increase damage, especially to reefs and beaches (Cintron et al.
1981).

Corals appear to be more susceptible to detergents used as
dispersants and to detergent/oil mixtures than to oil alone
(Elgershizen & Kruijf 1976). Chronic low levels of oil pollution
endanger coral reproduction (Loya & Rinkevich 1979). In general,
oil poses the greatest threat to shallow reef flats where it
actually contacts coral colonies. Little is known of the effects
of oil pollution on seagrass beds.

Mangroves are especially vulnerable to oil spills as oil can
coat the prop roots of red mangroves and the pneumatophores of
black mangroves, cutting off oxygen. Death of the trees is
inevitable if oil heavily coats the intertidal root areas.
Toxic products can remain in the soil resulting in further die
off. Lugo et al. (1980) noted sublethal responses, including
leaf deformities and partial defoliation.

An environmental sensitivity index atlas was prepared for
the USVI coastal areas by Resource Planning Institute with
assistance from VIRMC members in 1986 (RPI 1986) . Coastal
segments for St. John, St. Thomas, and St. Croix were mapped and
ranked in order of increasing susceptibility to oil pollution.
In general, low energy shorelines with sheltered mangroves are
the most vulnerable.

Natural Stresses

Storms

Storms have undoubtedly caused the greatest destruction to
coral reefs off the south side of the island. Hurricanes David
and Frederic (1979), known to have caused considerable damage to
reefs off St. Croix, are most likely responsible for the
widespread fragmentation of the dominant branching coral species
seen in Fish Bay, Reef Bay, Coral Bay and elsewhere. Tropical
Storm Klaus hit the U.S. Virgin Islands on Nov. 6-7, 1984, with
winds gusting to 50 knots and very heavy sea swells. About 229
mm of rain fell during the storm, but the damage was primarily
from the heavy swells rather than turbidity. Rogers & Zu11o
(1987) describe the physical damage to corals in Fish Bay.

Coral diseases

Coral diseases are affecting different hard coral species in
the waters around St. John. White band disease primarily affects
the acroporid corals, Acropora palmata and A. cervicornis, the
same species which are the most vulnerable to storms. Acropora
palmata is the dominant shallow water coral in the Virgin
islands, and throughout the Caribbean. The disease is
characterized by a distinct white strip about I cm wide where
live coral tissue has been completely removed from the skeleton.
It usually kills the entire coral colony. Peters (1983) suggests
bacteria may be responsible, but no causal agent has been
specifically identified. This disease can have a significant
impact on reefs if the coral colonies weaken and eventually
collapse because of an increase in carbonate-boring organisms.
Decreases in structural complexity of the reef could affect
associated fish communities (Gladfelter 1982, Sano et al. 1987).

White band disease has been found on many reefs throughout
the Caribbean, although in many cases the impact has been
negligible (Gladfelter 1982, Rogers 1985). However, the disease
has affected over 5 ha of the A. palmata reef at Buck Island Reef
National Monument, St. Croix (Gladfelter 1982), and has recently

killed large areas of A. palmata on other St. Croix reefs. Davis
et al. (1986) photographed the progression of the disease in
elkhorn corals at Buck Island and recorded a rate of advance of
4-5 mm/day. Some of the Buck Island corals are regenerating and
new colonies have become established (W. Gladfelter, pers.
comm.).

Beets et al. (1986) found colonies with white band disease
in the following locations around St. John: Brown Bay, Whistling
Cay, Cinnamon, Mennebeck Bay, Windswept and Jumbie Bay. A
relatively large number of standing dead A. palmata colonies at
Jumbie Bay may reflect the presence of white band disease in the
past.

Anderson et al. (1986) concluded that white band disease is
widespread throughout the British Virgin Islands. They estimated
that 0 to 25% of the elkhorn colonies in their study areas
exhibited white band disease, with most sites having less than
10% of the colonies affected. It is impossible to determine how
severe the disease was in the past, although in some areas a
large amount of "standing dead" coral was seen.

Black band disease infects several hard coral species
including Montastrea annularis, M. cavernosa, Diploria strigosa,
and D. labyrinthiformis and some of the common gorgonian species
in the Caribbean (Antonius 1985). The pathogen is the blue-
green algae Phormidium corallyticum. Black band disease occurs
on corals in VINP and BUIS.

White band and other coral diseases have not been directly
correlated with human activities, although Peters (1984) suggests
that injuries to corals from snorkelers and divers, and adverse
environmental conditions, such as high turbidity and sediment,
could increase the frequency of occurrence. National Park
Service photographs from the early 1970's at Buck Island Reef and
a diagram appearing in Robinson (1973) indicate presence of white
band in the Virgin Islands at that time. Gladfelter (1982)
studied elkhorn corals with white band in 1976 at Buck Island.
It is not known if visitation to Caribbean coral reef areas and
damage resulting from recreational activities have been
accompanied by increases in coral diseases.

Coral bleaching and other stresses

In the summer of 1987, many corals throughout the Caribbean
began to expel their zooxanthellae and lose their normal
coloration (Williams et al. 1987). Changes in salinity,
turbidity, exposure, and temperature can cause this coral
bleaching. Glynn (1984) documents an extreme case in Panama
where bleaching was followed by death of the primary framework
corals. Scientists suspect warm temperatures were at least
partially responsible for the zooxanthellae expulsions in 1987.

This episode was the most severe on record for Florida reefs, but
corals which bleached in July had begun to recover by November
(Jaap 1988).

VINP staff observed bleaching in several hard coral species
and in the zoanthid Palythoa around St. John in October 1987. It
was not as severe as that reported for Florida. The brain corals
D. labyrinthiformis and D. strigosa were most severely affected
although not all colonies of these species turned pale.
Bleaching seemed most conspicuous in relatively protected,
shallow bays and apparently less extensive in deep areas with
strong currents. Agaricia lamarcki colonies as deep as 27 m were
bleached. Portions of some affected corals have died and become
covered with algae, but other colonies have recovered entirely.

Another Caribbean-wide epidemic occurred in 1983 when over
90% of the population of long-spined black sea urchins (Diadema
antillarum) died (Lessios et al. 1984). The die off was
followed, at least on some reefs, by extensive growth of algae
which the urchins normally feed upon (Hughes et al. 1987). The
mass mortality occurred in December 1983 around St. John.
Research in Lameshur Bay suggests that although the population is
recovering, densities may not return to previous levels for
several decades. However, the documented increase in average
urchin body size may result in a return to pre-mortality urchin
and algal biomass levels much sooner (Levitan 1988).

Witman (1988), also working in Lameshur Bay, found that
predation by the fireworm Hermodice carunculata on shallow reefs
limited branch growth of the fire coral Millepora complanata and
opened up lesions on the colony surfaces. These lesions were
rapidly colonized by crustose corallines and other algae. Fire
corals only regenerated about 25% of the spaces cleared by
predation in one year.

ENDANGERED AND THREATENED MARINE SPECIES

Sea Turtles

In 1980, NPS began monitoring sea turtle nesting activities
on St. John and Buck Island, collecting data on seasonality,
nesting success, clutch sizes, incubation/emergence, and nest
predation. Hawksbill (Eretmochelys imbricata), leatherback
(Demochelys coriacea) and green (Chelonia mydas) turtles all
nested on USVI beaches in the past (Towle et al. 1978).
Hawksbills and leatherbacks are considered endangered, while the
green turtle (Fig. 11) is on the federal list of threatened
species. Currently, hawksbills nest far more frequently than the
other species.

Sandy Point, on the west end of St. Croix, is one of two
Caribbean sites where leatherbacks nest in abundance. Greens do
not appear to nest on St. John anymore, and only a few greens
have nested on Buck Island since 1980 when rangers began keeping
records. Four leatherbacks came up on Trunk Bay beach in 1985
and one appeared on Cinnamon in 1984, the first documented
observations of this species on St. John since the 1950's.

Mongooses are the primary predator at Buck Island and St.
John, preying on eggs and hatchlings. In Small's 1980-81 study
they were responsible for destroying an estimated 23% of all
turtle eggs on St. John (Small 1982). Extensive trapping of nest
sites reduced predation from 15% of nests in 1982 to 6% in 1983.
Prior to total eradication of mongooses at Buck Island (completed
in 1987), mongooses were destroying up to 55% of all nests on the
island. Inundations by heavy sea swells destroys some nests, and
poaching is a problem on both islands.

On St. John, nesting of hawksbills generally takes place
between June and December with peaks from August through
November. At Buck Island, records indicate nesting from April
until late December with peak activity from late July through

Fig. 11. Green turtle (Chelonia mydas) over seagrass and sand
bottom. Photo: L. McLain.

October. Leatherbacks can nest as early as March. Given the 60-
70 day incubation period, nesting activity can occur year-round
on St. John and Buck Island.

Yearly fluctuations in nesting activity at Buck Island and
St. John probably reflect both different levels of monitoring and
natural variation in the 2-3 year nesting cycle of sea turtles
(Zullo 1986). Although 20 of St. John's beaches supported
nesting between 1980-1985, only Cocoloba had nests every year.
In spite of its small size (71 ha) Buck Island has more activity
than St. John, and the turtles using its beaches deserve special
protection. Starting in 1987, NPS staff intensified the
monitoring of sea turtle nesting on Buck Island. In 1987, they
recorded 120 activities (crawls, nests, aborted nests) and
confirmed 65 nests. Most nests were those of hawksbills,
although a few leatherbacks and greens also nested.

Marine Mammals

A variety of marine mammals have been seen in USVI waters
including bottlenose, spinner, and spotted dolphins, and
humpback, killer, pilot, finback, and sperm whales. The National
Marine Fisheries Service began keeping statistics on sightings in
1980. Humpbacks travel south to Caribbean waters to mate and
bear their young. They are seen in the USVI from November to
May. The largest number (about 250 animals) of humpbacks on
record were sighted in the 1984-1985 season.

Other Species

Park waters contain limited populations of endangered black
corals (Antipatharians), primarily off the northeast coast of St.
John in depths over 15 m. Beets (pers. comm.) mapped and
measured 35 colonies of three species in Haulover Bay.

FISHERIES

A series of VIRMC fisheries projects done from 1984-1987 had
the following objectives: to study the role of fishing in the
culture and economy of the VIBR, to assess the fish and shellfish
stocks, to determine an ecologically realistic definition of
fishery management zones around St. John, to determine if
management zones should include the BVI or other areas outside
VIBR, to see if fish and shellfish stocks within the VIBR or at
Buck Island are declining, and to initiate an effective program
for long-term monitoring of fisheries. Lesser Antillean
fisheries were also characterized. In this section, these VIRMC
studies and other pertinent reports are discussed. Mahon et al.
(1986) prepared an annotated bibliography of references on
fisheries in the Lesser Antilles.

SOCIOECONOMIC AND CULTURAL ROLE OF FISHING IN VIBR

Studies of fishing on St. John indicate that few people on
the island made or make a living from fishing, but it is a very
important part of island culture (Dammann 1969, Koester 1986).
Most people catch fish for their families and sell or give away
the surplus.

Fiedler & Jarvis (1932) interviewed fishermen on St. Thomas
and St. John in 1931. At that time 78 out of a total of 405
fishermen were from St. John, and all boats, with the exception
of one motorboat, were powered by oar or sail. The island's
population was about 765. Fish sold at 6 cents a pound.
Fishermen used whelk, conch, and lobster to bait their traps.
Ninety tangle nets were in use for catching turtles. The most
extensively fished waters were those between St. Thomas and St.
John.

In 1959, there were 11 full-time fishermen and less than 75
part-time fishermen out of a population of about 800 (Idyll &
Randall 1959). Idyll & Randall (1959) estimated 500-560 traps
were set by local and British Virgin Islands fishermen. Most of
the catch was from traps, followed by handlines and beach seines.
Apparently, only 5 seines were used around St. John, but these
were about 360 feet long and 12 feet deep.

In 1959, demand for fish on St. John was not met.
Apparently, most St. Thomas and BVI fishermen fishing around the
island took their catch to Charlotte-Amalie, St. Thomas. At the
time of this survey, residents did not feel fish populations had
declined.

3 2

Dammann (1969) estimated about 400 people were involved in
commercial fishing in the USVI in 1967, with 120 of them full
time. He concluded that the reeffish populations were not being
overfished at this time because while the number of fishermen was
about the same as in 1930, the number of fish pots had decreased
by about half, and the total catch had increased. Clavijo (1984)
reported 454 licensed fishermen in 1982-1983. Olsen (1979)
described recreational fishing in the USVI and concluded that the
estimated combined landings from commercial and recreational
fishing probably exceeded the maximum sustained yield.

Koester (1986) described the role of fishing in the economy
and culture of St. John based primarily on interviews in March
1984, with 16 fishermen (13 of whom resided on the island) and 7
government agency employees. He estimated a total of 25 to 30
artisanal fishermen lived on St. John, ranging from those who
fish parttime for their own families to a few individuals who
consider fishing their main occupation and sell their catches.
All use relatively low technology methods. At the time of this
study, only two people were fishing commercially, and they fished
outside park waters.

An unknown number of people from the British Virgin Islands
and St. Thomas use the waters around St. John for fishing. Some
St. John residents go to the BVI to fish where lobsters, conchs
and whelks are more abundant.

Fishermen primarily use fish pots or traps to catch reef
fish and lobsters, followed by hook and line methods for pelagic
and bottom fish. Trapping has traditionally been the preferred
fishing technique (Dammann 1969, Boulon & Clavijo 1986). People
also Scuba dive for conchs and lobsters.

The demand for fish on St. John consistently exceeds the
supply. Fishermen believe that populations of lobsters, conchs,
whelks and reeffish have declined and cite a variety of causes:
1) adverse environmental changes associated with tourism;
2) increased boat traffic, resulting in cutting and dragging of
trap lines;
3) theft of fish traps; and
4) restrictions imposed by the park.

It is difficult to assess the magnitude of the impact of
park regulations on fishing. Trap fishing and hook and line
fishing are still permitted. However, the fishermen are
constrained by a variety of restrictions. For example, Title 36
CFR (Code of Federal Regulations) establishes quotas on lobsters,
conchs, and whelks, prohibits spearfishing, restricts use of nets
to those under 20' long, and prohibits all fishing in Trunk Bay.

3 3

The Federal Endangered Species Act of 1973 prohibits
catching of sea turtles. other regulations, e.g., against
cutting of certain plants used in fish trap construction, also
limit traditional activities.

The restrictions on the use of gill nets and longer seines
for catching baitfish perhaps has had the greatest impact. The
relative abundance of baitfish inside and outside park waters is
not known.

The tourist industry has provided new employment
opportunities, replacing the island's traditional subsistence
economy. The park probably has had more of an effect on the
island because of its role in increasing tourism and related
industries than from its restriction of traditional uses of the
land and sea. Koester (1986) recommends integration of fishermen
into the management of VINP/BR.

In spite of protective legislation in the USVI and
specifically within VINP, and a gradual decrease in the amount of
fishing around the island as people have given up fishing for
jobs in tourism and elsewhere, fish populations have declined
drastically.

ASSESSMENT OF FISH AND SHELLFISH STOCKS

Estimation of most fishery stock parameters is extremely
difficult. Stocks are a function of recruitment, growth, natural
mortality, and fishing mortality. Stock estimates can be based
on 1) estimates of productivity in terms of carbon transfer
through trophic levels, 2) length-weight frequencies of
individual species, and 3) age, growth, and mortality rates.
Fisheries information on catch per unit effort (CPUE) is also
useful.

Dammann (1986) notes that although both biological and
fisheries data are routinely used in stock assessment, the
resulting estimates are often not entirely suitable for tropical
shallow-water fisheries. it is especially difficult to assess
stocks in shallow tropical seas because of the large number of
species and the complex interrelationships of the coral reef,
seagrass bed, and mangrove ecosystems which they inhabit.

Large assemblages of species often occur on narrow insular
platforms separated from one another by deep ocean passages. In
the Eastern Caribbean, each island platform is usually considered
to contain its own fisheries stock despite the fact that there is
obviously a great deal of gene exchange throughout the region.
Most species in the Virgin Islands, and Puerto Rico fisheries have
pan-Caribbean distributions. There is recent evidence that
marine planktonic larvae can maintain their position relative to

3 4

a particular insular shelf without being swept downstream. Each
insular platform apparently recruits new individuals both from
within and outside its own system. There are no natural
boundaries to the present VIBR, and, in light of daily and
seasonal migrations, it is not appropriate to treat the resources
within the area as distinct stocks.

Useable fisheries data are derived from the entire
geological platform or relatively large sections of it, and at
present are not suitable for estimating sustainable yields from
such a small arbitrarily defined area as the waters within VINP.
It is probably not appropriate to treat the marine resources
within the biosphere reserve as distinct stocks.

Fisheries yields from shallow tropical reef, seagrass bed,
and mangrove areas perhaps average about 4-5 metric tons/sq.
kilometer/yr (Munro & Williams 1985). Larger yields (up to 15
mt/km2/yr) come from well-developed reef areas in the Pacific
where fishermen harvest almost all species of living organisms
(Munro 1984).

The Caribbean Fisheries Management Council represents the
governments of the USVI, Puerto Rico, and the U.S. mainland. It
works in cooperation with the BVI government to produce Fisheries
Management Plans (FMP) for the local fisheries resources.

An analysis of the data contained in the FMPs of shallow-
water reef fish (CFMC 1985) and lobsters (CFMC 1981) reveals that
1) local shallow water resources are heavily, and in some cases
over-exploited, 2) fisheries productivity of the shelf is
comparatively low (3.3 mt/km2/yr), and 3) maximum sustainable
yield (MSY) estimates are low (shallow water reeffish for PR and
the USVI, 3500 mt/yr; spiny lobsters for PR and USVI 373.5 kg/yr;
conchs, 163.8 kg/yr for St. Thomas-St. John.)

FISH COMMUNITY STRUCTURE

Randall (1968) described 300 common reeffish species from
the West Indies. He examined the food habits of 212 species
based primarily on specimens from Puerto Rico and the USVI
(Randall 1967). For two study sites in St. John, the fringing
reef in Lameshur Bay and one at Ram Head Bay, herbivores made up
about 24%, omnivores about 16%, and carnivores 60% of the total
fish biomass. Randall (1963) installed an artificial reef in
Lameshur Bay in 1960 and found that it supported 11 times more
fish than a nearby natural reef, probably because of the food
provided by a nearby seagrass bed.

Boulon et al. (1986a) studied approximately 50 commercially

important species from 20 different habitat types within VIBR
using a visual census technique. They recorded number of
individuals of each species and minimum/maximum length for each
species. Grunts, groupers, goatfishes, snappers, triggerfishes,
mackerels, squirrelfishes, and parrotfishes are some of the most
important commercial groups they observed.

Variations in the number of censuses done by Boulon et al.
(1986) for each habitat type made comparisons difficult. In
general, however, high species diversity was correlated with high
habitat complexity. Of all habitats sampled, the lower forereef
(the seaward extension of most reef systems) stands out because
of a high number of species and the relatively larger size of
individual fishes. Lobsters are also associated with reef areas
with more complex structure, shelter in crevices and overhangs.

Bank pavement areas, often deeper than 20 m, also tend to
have larger individuals and may be where they find the principal
habitat for lobsters and queen triggerfish. Juvenile fishes tend
to be associated with shallow mangrove areas while fish with
planktonic eggs reproduce in deeper, offshore habitats. There is
some evidence that deepwater algal plains and shallow grass beds
serve as nursery areas as well. Migration of coastal pelagic
fish species, lobsters, and conchs present sampling and
monitoring difficulties. However, most reeffish are non-
migratory which allows more management options.

Boulon et al. (1986) selected four long-term monitoring
sites for fish: Fish Bay, Lameshur, Hawksnest, and Trunk Bay.
Fish Bay was selected because of potential problems from land-
based sediment associated with watershed development. Lameshur
has little impact from coastal development but has some artisanal
fishing pressure. Previous scientific research carried out here
can be used for some comparisons. Concern over development of
the watershed (particularly building of the St. John clinic) and
observation of heavy runoff led to selection of Hawksnest Bay.
Trunk Bay was selected in an attempt to study visitor impact.
The V.I. Division of Fish and Wildlife and VINP recently signed a
cooperative agreement to continue monitoring at Lameshur and Fish
Bays.

FISHERIES LANDINGS ON ST. JOHN

Dammann (1969) and Olsen (1979) provide landings data for
fish and shellfish in the USVI. More recently, Boulon & Clavijo
(1986) looked at 6 months of fishing landings for commercial and
subsistence fishermen specifically on St. John. It is impossible
to determine the amount of catch landed on St. John that actually
comes from within VINP waters. Data came from 28 commercial and
10 subsistence fishermen who landed catch on St. John. (Only 12

St. John fishermen had commercial licenses in 1984.) For the
first 6 months of 1984, commercial fishermen landed over 16,000
lbs of fish and shellfish.

About 1400 lbs of lobsters were caught adjacent to or within
VINP in a 6 month period in 1984. Fishing for lobsters is
concentrated around Fish Bay, Reef Bay and Johnson's Reef (north
side). Only 3 fishermen reported conch catches landed on St.
John. Most of the conchs came from the south shore where
seagrass beds are more extensive. Subsistence fishermen take the
same species as found in the commercial catches, with each
individual landing less than 50 lbs of fish per month. Most of
their fishing takes place outside the park.

Further work on the intensity of trap fishing within park
waters and around the island of St. John is necessary.
observations by park staff indicate that the number of traps
which lack buoys at the surface and other required identification
may exceed those which are marked correctly. Within the park,
Lameshur appears to be the the most intensively fished bay.

FISH POISONING

Fish toxicity is a serious marketing concern in the USVI.
Over half of the total weight of fish landed and half of the
species in the fishery have some risk of carrying the ciguatera
toxin which is associated with a benthic dinoflagellate (Olsen et
al. 1982). About 3.5% of the species are very likely to contain
the poison, including several of the jacks, the dog snapper, and
the barracuda (Olsen et al. 1982, Dammann 1969). Toxic fish are
apparently unaffected and do not exhibit any signs to indicate
they are poisonous. No practical test for toxicity has yet been
developed. Ciguatera occurs throughout the Caribbean but most
frequently in the islands north of Martinique. One of the areas
of highest risk is south of Norman and Peter Islands in the BVI.
Doctors in the Marshall Islands (Pacific) have recently found
that mannitol, a drug used to reduce swelling of the brain,
dramatically and quickly relieves symptoms of ciguatera in some
victims.

HABITATS OUTSIDE VINP OF ECOLOGICAL IMPORTANCE TO BIOSPHERE
RESERVE FISHERIES

Boulon (1986b) carried out a study to determine which
habitats outside VIBR are of ecological importance to fisheries
within it. VIBR can not be considered a closed system because of
the migration of organisms and the production of planktonic eggs
and larvae by most food fishes. The generally westerly flow of

3 7

the major currents near St. John suggest that planktonic eggs and
larvae reach the island from sources "upstream", or to the east.

Boulon (1986b) sampled the following areas: Pillsbury
Sound, SW St. John, bank patch reef off Reef Bay, Hurricane Hole,
Norman Islands, NE St. John bank pavement, Great Thatch, and
Soper's Hole. Of these, he concluded that only a few could have
importance for the VIBR. Coral Bay is the largest and most
protected bay on St. John, and the most extensive mangrove
communities on the island occur here. This area may serve as a
recruitment source for lobsters and fish for many reefs and near
shore communities along the south coast of the island. Bank
pavement areas northeast of St. John in the Narrows and off SW
St. John provide good lobster habitat and support large
populations of adult fish. The large bank patch reef off Reef
Bay apears to be a good spawning location. Soper's Hole
(Tortola) has suffered from dredging, but may still be an
important nursery habitat.

LOBSTER AND CONCH FISHERIES

DuBois (1985) summarized the history of lobster and conch
fisheries in the Caribbean, depletion of the stocks, and the
constraints on effective management of the fishery. Lobsters and
conchs are currently the Caribbean's most economically important
fishery exports. Valencia (1978, cited in DuBois) estimated that
the fisheries contributed less than 1% of the Gross National
Product of the USVI. However, fishing is an important tradition
in the islands, and its cultural and social value should not be
overlooked.

In the past, the USVI exported lobsters to other countries.
Lobsters were apparently so abundant that they were hard to give
away and often served as bait for fish traps. The USVI was
exporting lobsters to Puerto Rico as late as 1960 (Caribo 1961).
Since the late 1960's, the USVI has been importing conchs and
lobsters, primarily to meet the demands of tourism.

DuBois (1985) noted that attempts to reverse the decline of
lobsters and conchs throughout the Caribbean have been largely
ineffective. DuBois (1985) states: "Within the region, the USVI
perhaps represents the most dramatic example of depletion of
stocks due to excessive demand associated with growth and tourism
development". He concluded that Belize and Turks and Caicos have
been relatively successful in managing lobster and conch
fisheries largely because of effective fishermen cooperatives and
environmental education. The USVI fishermen are not currently
organized into effective cooperatives.

Conchs

Randall (1964) studied the basic biology of the conchs in
St. John in the late 1950's and early 1960's. Conchs occur over
a large depth range and in a wide variety of habitats including
sand, rubble, and seagrass but most often in seagrass beds (Fig.
12). They are herbivores which primarily eat algae but also
ingest seagrass. Randall (1964) found that conchs tended to
spawn in warm months, ceasing in November or December and

Fig. 12. Queen conch (Strombus gigas) in a seagrass bed in Brown
Bay. Photo: L. McLain.

beginning again in January, February, or March. Predators of
conchs include spotted eagle rays, lobsters, queen triggerfish,
and sharks. While some predators are only capable of consuming
the animal after it is removed from its shell, a few species can
crush small conch shells to get at the animal inside.

Deterioration of seagrass beds around St. John (Williams
1988) could be partly responsible for the observed depletion of
conch populations. Randall (1964) referred to hundreds of conchs
in Salt Pond Bay, but populations this large have not been seen

there in the last several years.

Boulon (1987) monitored juvenile and adult conch populations
in Reef Bay, Outer and Inner Fish Bay, Hawksnest Bay, and
Threadneedle Bay monthly in 1985, finding very low numbers of
individuals and widely fluctuating population sizes. He also
used a diving sled to survey conchs around St. John in 1985,
repeating surveys done four years earlier (Wood & Olsen 1983).
Numbers of conchs in 1985 were not significantly different than
in 1981, although Boulon (1987) suggests caution in drawing
conclusions from these results because of small sample sizes.
Spot checks at 14 other sites around the island which appeared to
be good habitat (e.g., Inner Fish Bay and Reef Bay seagrass beds)
revealed 5 sites with no conchs and the remainder with less than
15 each (Boulon 1987). Piles of empty shells were found at
several locations.

Absence or low numbers of conchs at sites known to have
supported conch populations in the past (Hawksnest, Inner Fish
Bay) could reflect migration or harvesting of the animals. The
southern park waters, at least above 20 m, have more conchs that
northern waters, with Outer Fish Bay having the largest
population (Boulon 1987). The number of conchs in this bay were
significantly less in the summer months, and Boulon (1987)
suggested conchs migrate offshore into deeper water during the
summer reproductive season to mate and deposit eggs.

In contrast, Hesse (1979) and Coulston et al. (1985)
described migrations in the fall months in the Bahamas and St.
Croix, respectively. Coulston et al. (1985) saw reproductive
activity from March through November in depths of 15 m to 21 m.
Deepwater populations may move inshore into adjacent shallow bays
such as Threadneedle. Coulston et al. (1985) and Boulon (1987)
agree that deepwater reproductive populations, which escape
fishing pressure, may be essential to maintenance of the species
in USVI waters. Boulon (1987) suggests that the continuing take
of immature conchs around St. John will result in near
elimination of local populations. Recent research suggests that
the flared lip on conchs may not necessarily indicate
reproductive maturity (Wilkins et al. 1985), which means that
regulations which only permit taking of conchs with flared lips
are not protecting the reproductive populations as intended.

Lobsters

Boulon (1987) surveyed areas in Fish, Reef, and Hawksnest
Bays for spiny lobsters (Panulirus argus) (Fig. 13) and spotted
lobsters (Panulirus guttatus) in 1985 and 1986. Alarmingly low
numbers of lobsters were found at all sites, although the
presence of ledges, overhangs, and holes in these reef areas
would appear to provide good lobster habitat. For example, in
Reef Bay, only 0 to 4 lobsters were observed in each of 9

Fig. 13. A spiny lobster (Panulirus argus) on Johnson's Reef.
Photo: L. McLain.

censuses, with all of these having carapace lengths less than 3.5
inches (legal size). At Hawksnest, only 1 spiny lobster was seen
in 11 monthly surveys. Spot checks for lobsters at 9 additional
sites in the summer of 1985, turned up exceptionally low numbers
of both species as well.

Knowledge of basic lobster biology is a prerequisite for
effective management. Tektite studies provided information on
population dynamics (growth and mortality), ecology, and behavior
(Cooper et al. 1975, Herrnkind et al. 1975, Olsen et al. 1975,
Olsen & Koblic 1975). Olsen et al. (1975) studied lobster
distribution around all of St. John using Scuba. In their
survey, lobsters were rare from Hawksnest to Francis Bay and on
the east end of the island. Mean lobster densities ranged from 0
to 19.4 individuals/ha. This contrasts with an estimated mean
density of 64.8 lobsters/ha in the Dry Tortugas in the early
1970's (Davis 1977). Their observations were confined to waters
18 m deep or less, and substantial lobster populations could
occur deeper. Lobsters tended to inhabit areas near dropoffs to
deep water.

The reproductive period for lobsters appears to be in the
spring and fall according to Olsen et al. (1975) or just in the
spring according to Davis (1977). The planktonic phase of the
spiny lobster's life lasts an estimated 6 to 7 months (Sims &
Ingle 1966, cited in Olsen et al. 1975). Mortality may be over
99% (Buesa 1969; cited in Olsen et al. 1975). Juveniles settle
in shallow mangrove and seagrass bottoms. The long-spined black
sea urchin Diadema antillarum commonly provides shelter for young
lobsters (Davis 1971). Olsen et al. (1975) suggest most juvenile
lobsters in USVI waters occur in mangrove lagoons. Juveniles
leave the mangroves after 2 years, after reaching a carapace
length of about 80 mm. Some data suggest offshore movement of
lobsters during fall and winter out to the reefs.

Olsen & Koblic (1975) found that female lobsters take up to
4 years to reach market size after settlement from the plankton
while males take 2 years. Survivorship was estimated at 52 to
66% per year. These authors felt that even within park waters
the lobster population showed the effects of fishing.

Herrnkind et al. (1975) studied den occupancy and nocturnal
movements of spiny lobsters. One study site in Greater Lameshur
had population densities ranging from 4.8 to 15/ha. Lobsters
leave dens at night to feed in reefs or sand-algal plains,
returning before dawn. These authors suggest resident lobsters
have home ranges of 200 to 300 m in diameter. Herrnkind et al.
(1975) found indications that intensive handling of lobsters
altered their distribution patterns after release (see also Davis
1977).

Gut analysis revealed lobsters eat primarily mollusks
(including small conchs) as well as Diadema, Echinometra, and
hermit crabs through scavenging and active predation (Herrnkind
et al. 1975). In his study at Dry Tortugas, Davis (1977) also
found that lobsters eat mollusks and echinoids, including
Tripneustes and Lytechinus.

Davis (1977) showed the effects of intense recreational
harvest of lobsters at Dry Tortugas (Fort Jefferson National
Monument). Following prohibition of harvesting for over 2 years,
sport fishing for lobsters was permitted for 8 months. Although
the control population did not change, the trap catch rate in the
experimental area decreased 58% immediately after harvest. One
year later, the catch rate had recovered to 78% of its former
level. Full recovery could take several years. Davis (1977)
documented seasonal movement between shallow and deep waters that
appeared to be related to mating and reproduction.

OTHER FISHERIES

The whelk or West Indian topshell, Cittarium pica, inhabits
rocky coasts a few feet above and below the water line. Randall
(1964) studied the general biology of the whelks in Europa Bay,
St. John. Boulon (1986, 1987) studied density, seasonal
distribution, and size classes of whelks between Windswept Beach
and Peter Bay on St. John's north shore. The conspicuous lack of
individuals larger than 6.9 cm suggests high levels of
harvesting.

Several species of fish are used as bait in local reef and
pelagic fisheries, including dwarf herring, redear sardine, false
pilchard, flying fish, ballyhoo, and mackeral scad. The Division
of Fish and Wildlife surveyed baitfish in the USVI from 1984 to
1987. In St. Thomas and St. John fishermen primarily use haul
seines to catch bait, while St. Croix fishermen generally use
cast nets. Some species of baitfish may show seasonality in
abundance.

FISHERIES OF BUCK ISLAND REEF NATIONAL MONUMENT

Tobias et al. (1988) studied lobsters, conchs and reeffish
at Buck Island from November 1985 to June 1986 to determine the
effectiveness of NPS regulations in protecting fish and shellfish
populations and to assess the impact of commercial trap fishing.
Censuses similar to those carried out previously by Gladfelter et
al. (1977) and Simpson (1979) at the same locations led them to
conclude that reeffish are decreasing in abundance. Park rangers
and concessioners feel there has been a severe decline. The
average density of conchs in seagrass beds west of the island was
0.14 individuals/m2 which compares to a density of 0.17/m2
reported by Boulon et al. (1986) for VINP. Less than 2% of the
animals censused were adults. The mean density of lobsters at 3
small reef areas was only 0.008/m2.

Fewer species and fewer fish overall were caught at Buck
Island in experimental traps than at Teague Bay, an unprotected
area south of Buck Island. Commercially important reef fishes do
not appear to be any more abundant than at Teague Bay or
elsewhere around St. Croix.

LESSER ANTILLEAN REGIONAL FISHERIES

Goodwin (1986) provides information on regional trends in
Lesser Antillean fisheries and the implications of these trends
for management of the VIBR and other protected areas in the

region. Sustainable conservation of regional ecosystems requires
effective inter-institutional and international cooperation.

Few data exist on the abundance and sustainable yields of
Lesser Antillean fishery resources. Shallow-water reeffishes are
apparently overfished in most of the Lesser Antilles (Goodwin et
al. 1985). Habitat destruction associated with coastal
construction will become an increasingly important threat to
fisheries. However, current landings could be moderately
increased through attention to underexploited resources (deep
water snapper-grouper stocks, migratory pelagic fishes).
Management of the resources is presently secondary to
considerations of improved local income, employment and
nutrition. Biosphere Reserves could benefit fisheries by 1)
including protected breeding and nursery areas, 2) managing zones
to increase local availability of fish, and 3) developing
strategies for use of underused resources.

The Organization of Eastern Caribbean States has drawn up a
draft set of fishery regulations which provides for the
establishment of marine reserves, thereby providing a legal
foundation for designating components of a multi-site reserve.
Strategies which involve the participation of local people have
been successful in the BVI, Dominica, St. Lucia, and elsewhere
and may provide a valuable starting point for development of a
regional biosphere reserve network. Case studies of successful
marine resource management strategies and research projects which
result in clear benefits are needed. The Draft Fisheries
Regulations include specific management measures for fish,
lobsters, conchs, sea turtles and coral.

FISHERIES REGULATIONS IN THE USVI

The Division of Fish and Wildlife in the Dept. of Planning
and Natural Resources has responsibility for managing the
fisheries in the USVI. Territorial Act 3330, in 1972,
established many regulations, including minimum size of lobsters
and gear restrictions. The Territorial Government also adopted
federal regulations in the Exclusive Economic Zone (EEZ), 3 to
200 miles offshore, which, (beginning in 1982), established a
minimum size limit of 8 inches for yellowtail snappers (with a
one inch per year increase to 12 inches) and 12 inches for Nassau
grouper (with a one inch per year increase to 24 inches). There
is also a closed season each year for Nassau groupers from
January I to March 31 to protect spawning individuals.

"Fisheries in Crisis" was the title of a conference held by
the Division of Fish and Wildlife in St. Thomas in Sept. 1987.
This conference provided momentum for the creation of Fishermen's
Committees of Overseers. The St. Thomas-St. John Committee
submitted a proposal to the Governor to declare a 5 year

moratorium on the taking of all conchs and a size limit and
closed season for whelks. With the Governor's signature, these
regulations went into effect February 1988. A size limit of 2
and seven-sixteenth inches (6.2 cm) for whelks became effective
on Feb. 1, 1988 as well as a closed season from April I to Sept.
30 of each year. Prior to this, there had been no regulations at
all on conchs or whelks, with the exception of those inside VINP.
Inside the park, people were allowed to take 2 conchs and 1
gallon of whelks per person per day. The National Park Service
and the local government have concurrent jurisdiction over park
waters, and the park has adopted the more restrictive
regulations.

In addition to regulations discussed above, the park
prohibits seines longer than 20', allows use of traditional
design fish traps, permits the taking of 2 lobsters per person
per day, and prohibits all fishing within Trunk Bay (Title 36
CFR).

RECOMMENDATIONS FOR FUTURE RESEARCH AND MANAGEMENT
OF MARINE RESOURCES

Effective long-term assessment programs must be established
for the marine ecosystems of St. John. The National Park Service
is providing funds for long-term monitoring of coral reefs at
Virgin Islands National Park, Buck Island Reef National Monument,
Biscayne National Park, and Fort Jefferson National Monument (Dry
Tortugas). Routine gathering of data on environmental
parameters, such as temperature, will be combined with tracking
of benthic reef organisms using photographic and transect methods
at permanent study sites (Rogers 1988) . Fish censuses will also
be carried out. The new Cooperative Agreement between the NPS
and the V.I. Divison of Fish and Wildlife should help to
facilitate cooperative research, particularly on local fisheries.

Conch and whelk populations should be systematically
monitored to determine the effectiveness of recent regulations.
Fish and lobster populations should be monitored to determine the
need for further management actions, for example, area closures,
gear restrictions, or increase in minimum size at harvest (see
Boulon 1987). Data must be collected in a statistically rigorous
way.

Local fishermen must be an integral part of the management
process and the decisions which are made. Numerous people have
pointed out and emphasized not only the desirability but the
necessity of bringing fishermen into the process of research,
information exchange, and management (e.g. DuBois 1985, Goodwin
1986, Koester 1986).

DuBois (1985) points out the critical need for research on
the linkages between habitat degradation (such as destruction of
mangroves) and fisheries (stock depletion). Overfishing has also
depleted lobster and conch populations in many Caribbean nations
(DuBois 1985).

The lack of sufficient information on the coastal and
inshore currents off St. John impedes research regarding
dispersal of larvae, sediment transport, and transport and
dilution of pollutants such as oil, nutrients and toxic
materials. Effective management of fisheries requires more
knowledge of larval dispersal patterns. As Munro & Williams
(1985) point out, "Effective management strategies will vary
greatly depending on the extent to which recruitment to a reef is
derived from within the fished population (self-recruiting) or
was spawned outside the system". Other research needs include:
migration patterns of coastal pelagics; seasonality of fish and
shellfish populations; and the population dynamics/habitat
requirements of baitfish.

Implementation of specific management recommendations for
the marine systems and fisheries of VINP could help to reduce the
pressure on these resources and perhaps reverse the deterioration
of benthic areas and the decline of fish and shellfish
populations.

The Shoreline Use Plan should be implemented as soon as
possible by installing mooring and marker buoys in critical bays
and near vulnerable reefs. Minimum depth or minimum distance
from shore requirements for anchoring should be considered in
areas where anchoring is allowed. Zoning of park waters for
fishing or other activities should be evaluated as a means of
reducing conflicts among resource users (see Kelleher 1985 on
zoning for Great Barrier Reef, Australia; see Putney 1987 on
resource users in VINP).

The water quality monitoring program started by the park's
research staff in 1988 to look at basic parameters (oxygen,
turbidity) should be continued and expanded to include collection
of data to help determine if sewage, oil, and fuel from boats are
causing deterioration of water quality in park waters.

TERRESTRIAL SYSTEMS

HISTORY OF RESEARCH: FLORA AND VEGETATION

The Virgin Islands were among the first Caribbean islands to
be explored by botanists. Several early floras (West 1793, von
Eggers 1879, Millspaugh 1902) dealt mainly with St. Croix.
Borgeson (1898, 1909, 1923) described the vegetation of the
Danish (later, United States) islands in a series of papers.
Britton published annotated systematic lists of plants of Anegada
(1916), the USVI (1918), and a narrative of collecting on Puerto
Rico and the USVI (1923). Britton & Wilson (1923-1930) prepared
a flora of the vascular plants of Puerto Rico and the Virgin
Islands, covering the area from Mona to Anegada and St. Croix.
Liogier (1965, 1967) made nomenclatural changes and additions to
Britton & Wilson's flora, and Fosberg (1976) revised the flora of
St. Croix.

Beard (1945) described the flora of the BVI. D'Arcy
prepared annotated checklists of the flora of Tortola (1967) and
Anegada (1971). Little (1969) studied the trees of Jost van
Dyke, and Little et al. (1976) treated the flora of Virgin Gorda.
Little & Wadsworth (1964) and Little et al. (1974) produced a
comprehensive two-volume guide to the identification and
distribution of the trees of Puerto Rico and the US and British
Virgin Islands. Oakes & Butcher (1962) discussed the poisonous
and injurious plants of the Virgin Islands. Woodbury & Little
(1979) prepared a flora of Buck Island Reef National Monument.

The works of Beard (1944, 1949, 1955) constitute the basic
references on tropical American vegetation, including the
Windward, Leeward, and the British (but not the US) Virgin
Islands. The plant ecology of Puerto Rico was studied by Gleason
& Cook (1927). Velez (1957) described the herbaceous angiosperms
of the Lesser Antilles. Ewel & Whitmore (1973) mapped the
ecological life zones of Puerto Rico and the USVI and described
their plant associations. Howard (1973) discussed the vegetation
of the Antilles, including biogeographical aspects. Lugo et al.
(1981) and Brown (1982) provide an overview of the status of
Caribbean forests in the early 1980's. Many other references on
the plant ecology of the West Indies can be found in Rundel
(1974). Mosquera & Feheley (1984) provided a bibliography of
forestry in Puerto Rico and the USVI.

While the above references contain much valuable
information, few works deal exclusively with the botany and plant
ecology of St. John. Woodworth (1943) prepared an annotated list
of the economic plants of the island. Robertson (1957) prepared
a preliminary map of plant formations as part of an initial

biological survey for the National Park. Forman & Hahn (1980)
gave a quantitative description of the spatial patterns of trees
in the Reef Bay forest. A list of vascular flora in VINP is
contained in the NPS NPFLORA computerized data base. Teytaud
(1988) mapped environmental factors and potential natural
vegetation in St. John. Woodbury & Weaver (1987) described and
mapped the existing vegetation types of St. John and Hassel
island, provided a species listing by vegetation type, and
collected herbarium specimens to be deposited at the Virgin
Islands Biosphere Reserve Center.

Weaver & Chinea-Rivera (1988) established sixteen long-term
monitoring plots in the Cinnamon Bay watershed in 1983 to explore
species-site relationships and structural characteristics of the
vegetation at different elevations. Stem density, mean height
and diameter of stems, basal area, number of species, and
diversity indices were calculated. The trees in these plots were
remeasured in 1988. Earhart et al. (1988) and Matuszak et al.
(1987) describe a long-term monitoring study, begun in 1984, of
vegetation on three plots in the Reef Bay, Hawksnest Bay, and
Fish Bay watersheds.

DESCRIPTION OF TERRESTRIAL SYSTEMS

Flora and Vegetation

According to the life zone system of Holdridge (1967) most
of St. John has a subtropical climate which is transitional
between dry forest and moist forest (Ewel & Whitmore 1973,
Teytaud 1988). A true moist forest life zone occurs only in a
small area on the highest and wettest parts of the island, and a
true dry forest life zone occupies only the driest areas at the
eastern end and some points of land along the shorelines. In
Beard's (1955) system of vegetation classification, upland areas
with this type of climate would normally support "climax" plant
formations consisting of semi-evergreen forest in the moist
forest life zone, and deciduous seasonal forest elsewhere. In
St. John the "normal" climatic factors are overridden by the
effects of steep topography, shallow non-zonal soils, and drying
tradewinds. These conditions shift the potential natural
vegetation to Beard's dry-evergreen formation-series, with dry
evergreen forest in the moist forest life zone and dry evergreen
woodland and thicket in other areas (see Teytaud 1988).

Recurrent hurricanes, human disturbances (chiefly vegetation
clearing and development, Tyson 1984), and the presence of exotic
feral animals such as pigs, goats, and donkeys have all had an
effect on the vegetation of St. John (Lazell 1980, Nellis et al.
in prep). Existing vegetation is almost entirely secondary and
therefore difficult to relate to Beard's descriptions of climax

formations. Harris (1963), Lazell (1980) and Byrne (1980)
provide comparative data from other West Indian islands subjected
to similar kinds of human disturbance.

Woodbury & Weaver (1987) mapped and classified the existing
vegetation of the uplands as follows: (a) moist forest; 10 to 30
m tall, with 2 or 3 tree strata (layers); trees over 70%
evergreen; has characteristics of both Beard's seasonal and dry
evergreen formation-series but not typical of either type;
subdivided into uplands gallery, and basin types; (b) dry
evergreen woodland (described by some authors as deciduous type);
5 to 10 m tall with two tree layers; leaves small; (c) dry
evergreen thicket or scrub 0 - 3 to 5 m tall; with small leaves;
(d) thorn and cactus, less than 5 m tall; dominated by tree cacti
and other thorny species; (e) recently disturbed secondary
vegetation in various stages of recovery; and (f) pasture
(Fig. 14). The most advanced seral stages occur in some moist
upland forests (e.g., Bordeaux Mountain summit) and gallery
forests (e.g., Reef Bay Gut and Battery Gut) but still contain
introduced species. The dry forests between the littoral and the
summit forests are made up of a mixture of secondary species and
show no pronounced altitudinal zonation.

Woodbury and Weaver listed nearly 800 species of plants in
116 families from St. John. Eighteen species were considered
rare and endangered, and six of these may be new to science
(Table 1). Of the 750 species of trees of Puerto Rico and the
Virgin Islands which were described in Little & Wadsworth (1964)
and Little et al. (1974), 547 (73%) are native and 203 (27%) were
introduced. Matuszak (1985) lists 14 endemic species for the
Virgin Islands. Woodbury & Weaver (1987) note that the areas of
vegetation under Park Service jurisdiction are unique because
similarly protected areas in these vegetation types do not exist
elsewhere in the Caribbean.

Table 1. Rare and endangered species of plants from St. John
(From: Woodbury & Weaver 1987)

Machaonia sp. (new) Erythrina eggersii
Malpighia sp. (new) Galactia eggersii
Eugenia sp. (new) Malpighia linearis
Byrsonima sp. (new) Malpighia woodburyana
Psidium sp. (new) Zanthoxylum thomasianum*
Apocynaceae (new, vine) Ilex urbanii
Peperomia myrtifolia Calyptranthes thomasianum
Schoepfia schreberi Solanum mucronaturm
Cypselia humifusa Tillandsia lineatispica

* On the Federal endangered species list.

Z3 Z

Earhart et al. (1988) established three permanent plots in
the forest of St. John in the Reef, Fish, and Hawksnest Bay
watersheds. They identified and measured all plants having a
diameter at breast height (dbh) greater than or equal to 5 cm,
measured a sub-sample of tree heights, mapped stem locations, and
determined the distribution of each species within each plot.
For each species, they calculated basal area and relative
density, dominance and frequency values.

The Bordeaux plot (1.0 ha), in upland moist forest, has 62
species in 33 families. Farming probably ceased on this plot 100
years ago. Located in gallery moist forest, the L'Esperance plot
(0.5 ha) has 39 species in 26 families and has been recovering
for an estimated 60-70 years. The Hawksnest plot (0.5 ha), in
dry evergreen woodland, has 54 species in 26 families and was
abandoned more recently than the other plots. The regenerating
forest in these 3 plots are composed of native and exotic
species. Matuszak et al. (1987) provide interesting historical
and ethnobotanical data on eight of the most important species in
the plots.

Tyson (1987) reported on land use in the Reef, Fish and
Hawksnest Bay watersheds during the period from 1718-1950.
Agriculture in all three watersheds peaked between 1755 and 1780
but continued at a relatively high level until 1820 and then
declined during the 19th century. Tyson delineates areas which
were subjected to intensive use, as well as some areas which may
have remained free from direct human impact.

Sage Mountain in Tortola supports a vegetation type that is
similar to the upland moist forest in St. John in some respects
but unique in others (see Little & Wadsworth 1964). Beard (1949)
classified this forest as "xerophytic rain forest,' thus giving
it an undeserved reputation for wetness (Bowden et al. 1970). In
his revised classification scheme Beard (1955) placed this type
in dry evergreen forest; it represents merely a somewhat wetter
example of the moist forest life zone found on St. John.
Although the Sage Mountain forest is nominally protected by the
BVI Government, the relatively "natural" portion of it has shrunk
steadily over the years. Efforts should be made under the aegis
of the Biosphere Reserve to help the British Virgin Islands to
protect the remaining forest, both as a valuable resource in its
own right and because of its importance for comparative research.

At the time Beard published his 1949 paper on the vegetation
of the Lesser Antilles, he could not find a single undisturbed
example of either dry evergreen or seasonal vegetation in the dry
forest and moist forest life zones of those islands. Very little
is known about the structural or functional characteristics of
natural vegetation in these life zones. Lugo et al. (1978)
studied the structure, productivity, and transpiration of Guanica
Commonwealth Forest, a subtropical dry forest on the southwest

coast of Puerto Rico characterized by secondary growth with scrub
forest, deciduous forest and semi-evergreen forest types. Data
on nutrient dynamics (Lugo & Murphy 1986) and biomass (Murphy &
Lugo 1986) for this forest are also available. This site is
somewhat drier than St. John and is underlain by shallow
limestone-derived soils, but is similar enough that the results
of these studies may serve as an approximation of the functional
characteristics of St. John's secondary dry forest. Lugo et al.
(1978) reported that productivity of the Guanica forest is low
compared to that of other tropical ecosystems. As in Cruz Bay
(Teytaud 1988), soil moisture deficit occurs for ten months of
the year and low soil moisture availability is the dominant
factor controlling forest productivity, growth, litter
decomposition, water loss, and physiognomy.

soils

Rivera et al. (1966, 1970) described and mapped the soil
types of the USVI and provided information on suitability of
soils for various uses. Most of St. John is covered by soils
belonging to the Cramer series; other soils are generally found
only in valleys and along the coasts (Rivera et al. 1970).
Cramer soils occur on steep mountainsides, and the surface layer
is gravelly or stony clay loam. Between 50% and 70% of the
surface may be covered with rock outcrops, boulders and stones.
These soilIs were formed in place from basic volcanic rocks and
are red or reddish-brown and moderately permeable. They are
well-drained, slightly acidic to neutral in the surface layer,
clayey, moderately fertile, shallow, and dry. The subsoil may be
at or below the wilting point for 3 to 6 months per year. Runoff
is excessive, permeability and infiltration are moderate, and
erosion hazard is severe. Most St. John soils fall in the U.S.
Department of Agriculture's hydrologic group D, indicating that
they have a high potential for runoff.

Buck Island is almost entirely covered with soils belonging
to the Southgate Rockland Complex (Rivera et al. 1970). These
soils formed in place on steep slopes from "granitic" rocks.
They are greyish-brown to dark brown clay loams that are 50% to
70% rock outcrops, with soil material less than 13 cm thick
between the outcrops. Loose stones and boulders are common on
the surface. These soils are well-drained, slightly acidic in
the surface layer, clayey, moderately fertile, very shallow and
have a low water-holding capacity. Collins & Craft (1988) note
that the classification of Rivera et al. (1966, 1970) should be
modified to reflect the conditions which prevail on tropical
islands with complex geological history.

Soils near the periphery of each of the long-term plots
established by Earhart et al. (1988) in the Fish, Reef, and

5 3

Hawksnest Bay watersheds were sampled in 1986 (Matuszak et al.
1987). Samples were analyzed for macronutrients (P, K, Ca, mg,
and 5) and micronutrients (Zn, Fe, Cu, Mn). Organic matter,
texture, sodium content, and pH were also examined.

Brown & Scheffel (1985) investigated the recovery of soil
organic carbon during forest succession on St. John. They found
that conversion of forest to agriculture results in almost a
three-fold decrease in soil carbon. However, recovery of soil
carbon in the moist forest life zone of St. John is relatively
fast and seems to occur within 50-60 years, a rate similar to
that found for dry forest and wet forest life zones in Puerto
Rico. The carbon content of the mature forest in St. John is
high and more typical of sites in wetter climates.

Groundwater

A reconnaissance survey of groundwater was carried out by
McGuinness (1946). Cosner (1972) prepared a more extensive
report, including alternatives of water supply.

About 1 to 3 inches of water per year enters the soil and
stream-chann,el deposits. The most important aquifer is the
fractured volcanic rocks throughout most of the island. The
depth to the water table averages less than 50 feet in the hills
of St. John, but it is close to the land surface along the
shoreline. Some of the groundwater may enter the sea, but much
of it is used by the vegetation in the valleys and along the
shore (Cosner 1972).

Jordan (1972) presents evidence which indicates that the
base flow of streams on the three main islands of the USVI has
diminished. Streams that were once perennial now dry up in the
dry season, except for a few spring-fed pools in the stream
channels that are sustained except in severe drought. Jordan
attributes this decline mainly to changes in land use, i.e.,
reversion of land from crops and pasture to dense brush and
forest. Recharge to the aquifer has been reduced because
rainfall that once infiltrated below the shallow root zone of
crops and grasses has been increasingly intercepted and
transpired by deep-rooted secondary brush and trees. A
contributing factor to the decline in base flow appears to be a
long-term decrease in rainfall since the early 1900's.

HISTORY OF RESEARCH: FAUNA

Much of the literature on the terrestrial fauna of the
Virgin Islands appears in works on biogeography and systematics

of various taxonomic groups. More information exists on birds,
reptiles, mammals, and amphibians than on invertebrates.

The volumes of the New York Academy of Science's Scientific
Survey of Puerto Rico and the Virgin Islands, issued at irregular
intervals from the 1920's to the present, are a basic reference
including many citations of earlier zoological work. Highfield
et al. (1985) produced a preliminary bibliography of holdings in
the Virgin Islands National Park libraries which contains
pertinent references. Work is continuing on a very comprehensive
annotated bibliography of the Virgin Islands (Highfield et al, in
prep.). The endoparasites, arachnida, insects, brochyuran crabs,
fresh water fishes, and mammals of St. Croix, were catalogued by
Beatty (1944). Philibosian & Yntema (1977a) published an
annotated checklist of the birds, mammals, reptiles and
amphibians of the Virgin Islands and Puerto Rico. MacLean (1982)
gives keys and species accounts of the reptiles and amphibians of
the Virgin Islands. Lazell (1980, 1983a, 1983b) gave an account
of the terrestrial fauna and the biogeography of the herpetofauna
of the British Virgin Islands. Riley's (1975) Field Guide to the
Butterflies of the West Indies is the basic reference for this
group. Raffaele's Guide to the Birds of Puerto Rico and the
Virgin Islands (1983), is more specific to the Virgin Islands
than the earlier standard, Bond's Birds of the West Indies
(1979), and contains information on endangered species. Chace &
Hobbs' (1969) systematic account of the freshwater and
terrestrial decapod crustaceans of the West Indies includes the
species present in the Virgin Islands.

Invertebrates and Aquatic Fauna

Muchmore (1987) investigated the invertebrates of St. John,
establishing many new distribution records and discovering a
considerable number of new species. Until recently only a few
records have existed in the literature of onychophora, isopods,
millipedes, scorpions, spiders, ants and a scattering of other
insects from St. John. Before Muchmore's work there were
apparently no records of snails, annelids, freshwater crustacea,
centipedes, most arachnida, and most groups of insects, though
some of these have been reported from St. Croix, St. Thomas, and
Tortola (Muchmore 1982). Curry (1970) prepared an entomological
checklist for the Virgin Islands Ecological Research Station at
Lameshur Bay and collected many specimens. Her checklist
includes more than 75 taxa in 10 orders, Coleoptera being the
best represented. Curry & Curry (1971) studied the Chironomids
(midges) in the USVI and on Anegada. Coleopterans (beetles) and
other insects from the USVI are described by Ivie (1983, 1985a,
1985b), Ivie & Miller 1984, Ivie & Triplehorn (1986), and Ivie &
Nickle (1986).

The freshwater stream habitat has apparently not been well
investigated in St. John. Besides insects and frogs, at least 8
native species of fishes, 9 shrimps and several kinds of crabs
are known to inhabit freshwater streams in St. Croix (Beatty
1944, Clavijo et al. 1980). Woodbury & Weaver (1987) found very
small areas of freshwater swamp vegetation in a few places on the
leeward side of St. John where there were large inputs of fresh
water. Typical tree species here are the pond apple (Annona
glabra), escambron (Machaerium lunatum), and Enallagma latifolia.
Examples of this uncommon habitat occur at Fish Bay and Coral
Bay.

An interesting and not uncommon freshwater habitat occurs in
the leaf axils of various species of bromeliads, which retain
moisture after rainfall. Miller (1971a, 1971b) investigated the
invertebrate communities living in leaf axils and found
scorpions, cockroaches, isopods, centipedes, two species of
frogs, various dipteran larvae, algae and protozoans. He studied
dipteran larvae in two species of bromeliads, Tillandsia
urtriculata from the xerophytic vegetation near the coast and
Aechmea lingulata from the moist highlands of Bordeaux Mountain.
The bromeliad leaf axil micro-habitat represents one of the most
widely distributed freshwater sources on the island, and it
enables dipterans to successfully inhabit areas which would
otherwise be too dry to support their larvae.

Amphibians and Reptiles

Of the 45 living and 6 extinct terrestrial species of native
amphibians, reptiles, mammals and birds listed by Philibosian &
Yntema (1977a) as breeding on St. John, none are endemic to that
island alone, but 11 of the non-flying forms are endemic to the
Puerto Rico Bank. These authors list a total of 111 species of
living vertebrates on St. John, including introduced and
non-breeding species (but not accidental birds or freshwater
fishes).

Four species of frogs, a land tortoise, 2 geckos, 3 Anolis
lizards, the common tree iguana, an Ameiva ground lizard, an
Amphisbaena worm lizard, and a Typhlops worm snake are found on
St. John today (Philibosian & Yntema 1977a, MacLean 1982). The
systematic literature on Virgin Islands amphibians and reptiles
is growing (Williams 1969; MacLean 1982; Nellis et al. 1983;
Lazell 1980, 1983a, 1983b). Philibosian & Yntema (1976, 1977,
1978) along with MacLean et al. (1977) provide recent
herpetological summaries for the USVI. Philibosian (1975)
studied the territorial behavior and population regulation of
Anolis acutus in St. Croix and A. cristatellus in St. John.

Birds

St. John plays an increasing role as a research site in the
tropics. One of the earliest lists of birds from St. John may be
found in Oldendorp's monumental cataloging from 1732 to 1768 of
life on St. John during the influence of the Moravian Church
(Oldendorp 1770). Successive lists of birds and new additions to
the bird fauna (Ridgway 1884) of the Virgin Islands, and St. John
in particular, have been published (Newton 1860; Nichols 1943;
Mortenson 1909, 1910). Bond's Supplements to the Check-list of
Birds of the West Indies (1940) and the revised field guide (Bond
1985) specifically mention St. John as site locations of range
extensions. The 6th edition of the American Ornithologists
Union's Check-list of North American Birds (1983) included an
expanded geographical range, including Central America, and the
West Indies. Again, St. John is frequently mentioned for its
range extensions for migrant avifauna. Norton (1983, 1986) has
prepared checklists for VINP, and Leck and Norton (in prep.) are
working on an updated checklist. Wauer (1988) recently produced
an illustrated guidebook to the more common birds in the USVI.

Lamb (1957) and Kepler (1971) provided information on
endangered species and their habitats. The Brown Pelican
(Pelecanus occidentalis) gained attention on St. John because of
sufficient numbers and accessible colonies for research (Boulon
et al. 1982, Collazo & Agardy 1982, Coblentz 1986). This
endangered species nests, roosts and feeds adjacent to and within
park boundaries. The recent listing of the Roseate Tern (Sterna
dougallii) as threatened (Federal Register, Nov. 2, 1987) in the
West Indies can in part be attributed to work on colonies near
St. John (Halewyn & Norton 1984, Norton 1988).

For example, the Puerto Rican (Stolid) Flycatcher (Myiarchus
stolidus) on St. John played a role in describing the status and
speciation of the Myiarchus group in the West Indies (Hall 1965,
Seaman 1957, Lanyon 1967). The type locality of the Caribbean
(American) Coot was St. John (Ridgway 1884), and the recent
discovery of the Lesser Antillean Bullfinch on St. John (Raffaele
& Roby 1977) illustrates that the bird life of St. John is
dynamic. New data may be expected as a result of the island's
location in the West Indian archipelago and because of its
regenerating habitats (Norton & Hobbs 1987).

International concern for the role of forest habitats as
wintering areas for migrant and resident birds in the tropics is
focusing on the island of St. John. Comparative studies
conducted on St. John and St. Thomas (Askins et al. 1987) are
expected to provide insight on the importance of forest habitats
on developed and relatively undeveloped islands in the West
Indies. Annual Christmas bird counts conducted on St. John for
the last 10 years and other islands in the Caribbean also provide

a data base for further comparative work (Norton in prep. a).

St. John has 24 living and 2 extirpated breeding species of
land birds (Leck & Norton, in prep; Norton 1986). Land birds are
here considered to be those in the families of vultures and
raptors, and pigeons through passerines (Bond 1979). There are
19 living and 3 extirpated (no longer breeding in the Virgin
islands) native breeding water birds, seabirds, waders and
shorebirds. One hundred nineteen (119) species are listed as
migrants or visitors not known to breed, of which 3 introduced
species may be found on St. John. The total list of avifauna as
of 1988 is 170 species.

Robertson (1962) laid the foundation of serious modern
ornithological study in St. John and gave a summary of previous
work. He investigated the quantitative distribution of landbirds
in relation to moist and dry forests. The heavy second growth
forest on St. John is especially attractive to non-breeding,
wintering wood warblers (Askins et al. 1987). Worm-eating,
Black-throated Blue, and Hooded Warblers were widely distributed
in the moist forest habitat in St. John (Robertson 1962).

The Virgin Islands lack most Greater Antillean species of
birds. Fourteen species of St. John's breeding land birds are
widely distributed in the Caribbean and 10 species reach range
limits in the eastern Greater Antilles. of the latter, 5
apparently reached the Virgin Islands from the Lesser Antilles
and 5 from Puerto Rico and the Greater Antilles. The Mocking
bird has recently extended its range to the Virgin Islands from
Puerto Rico; the other Greater Antillean species are rare and
appear to be relict. The Lesser Antillean species seem to be a
more recent and more aggressive element in the Virgin Islands
avifauna, and all except the Bridled Quail Dove are common and
extending their ranges (Robertson 1962).

Robertson noted that he had no satisfactory explanation for
the absence from the northern Virgin Islands of 22 out of the
total of 53 species in the native land avifauna of Puerto Rico.
Judging from the vegetation of St. John, he assumed that habitat
(i.e., heavy forest) suitable for many of the missing species was
available in the Virgin Islands under original conditions, and a
land connection to Puerto Rico is generally thought to have
existed during glacial periods in the Pleistocene. Only two
species of birds in St. John are now more or less restricted in
distribution to the heavier (moist) forest areas. Robertson
speculated that forest obliteration early in the historical
period may partially account for the present impoverishment of
forest birds, but admits that evidence to support this view is
lacking since the earliest information on Virgin Islands birds
dates from the 1850's, long after forest destruction and near the
close of the plantation era.

Only a few of the forest-adapted birds in the West Indies,
including the two found on St. John, are good overwater
dispersers. On St. John, poor overwater dispersal ability alone
is probably not a sufficient mechanism to account for the paucity
of forest species, because a land connection to Puerto Rico would
have afforded a direct route during the Pleistocene. According
to Pregill & Olsen (1981) however, dry scrub and savanna were the
predominant vegetation types on the emergent portions of the
Puerto Rico Bank whenever sea level was lowered by glaciation.
Species narrowly adapted to mesic conditions would have been
confined to reduced windward upland habitats on the higher
mountains of Puerto Rico. Most forest birds would have been
prevented from colonizing the Puerto Rico Bank by lack of habitat
during glacial eras and by water barriers during interglacials.
Thus, contrary to Robertson's speculations, the present poor
representation of forest species in the Virgin Islands may not be
a consequence of forest destruction by man.

Small, low islands tend to have fewer species than larger
and/or higher ones, since the size of the species pool is to some
extent a function of both the total area of the island and the
number of habitats available. With 24 extant breeding species of
land birds and 2 extinct, St. John has slightly more species than
the average for an island of its size and number of habitats, as
can be determined from Terborgh & Faaborg's (1980a) regression
analysis of species number on area and habitat diversity.

Terborgh & Faaborg (1980a) also found a striking difference
in the trophic organization of West Indian bird communities
compared to the mainland: in the Antilles the largest guilds (in
terms of biomass) are of frugivores rather than gleaning and
hovering insectivores. Since insects as a resource can be
partitioned in many more ways than fruit, communities low in
insectivores and high in frugivores will necessarily contain
fewer species than those in which the reverse is true. A partial
explanation for the paucity of insectivores in the Antilles might
be the finding that insect densities appear to be lower on
Caribbean islands than on the mainland, but paradoxically, the
biomass of insectivorous birds is greatly increased every winter
by large numbers of migrant North American warblers (Terborgh &
Faaborg 1980b). As many of its forested watersheds recover under
the protection of the National Park, St. John could become an
important site for the testing of theories on insular land bird
ecology (see Abbott 1980, Strong et al. 1984).

Seaman (1949-1963) authored a series of reports on his
studies of Virgin Islands wildlife, including various species of
birds. He studied the food habits of the Bridled Quail Dove
(Geotrygon mystacea) (Seaman 1966). A study of the biology of
this species in the Virgin Islands, including field work in St.
John, is presently in preparation by Nellis et al. Another study
of G. mystacea on Guana Island in the British Virgin Islands,

using radio tracking techniques, is in progress (Chipley in
prep.). Leck (1973) studied dominance relationships among
nectar-feeding birds (2 hummingbirds and bananaquit) in St.
Croix. LaBastille & Richmond (1973) gave an account of the birds
of Anegada, with descriptions of habitat and comparisons to
records from the USVI. Mirecki (1976) reported the results of
the Cambridge ornithological expedition to the British Virgin
Islands. Philibosian & Yntema (1977b) discussed the status of
endangered, threatened, and extinct birds of the Virgin Islands.

Norton (1979, 1981) reported several new records of birds
for the Virgin Islands, and has annually edited the West Indies
region seasonal summaries of American Birds (1980-1988),
including data from the Virgin Islands. Norton has reported on
the status of Pelecaniformes (Norton in press) and Sterninae
(Norton in prep. b) in the Virgin Islands. Norton (1984) noted
the expansion of the Cayenne Tern in the British and USVI and on
the Puerto Rico Bank (Schaffner et al. 1986).

Norton et al. (1986) discuss some aspects of distribution
and ecology of Virgin Islands anatids. Norton & Seaman (1985)
described the status of fledgling white-crowned pigeons banded in
the Virgin Islands. Nellis et al. (1984) discussed the
population status of Zenaida Doves and other Columbids in the
Virgin Islands. Lazell (1980, 1983) reported on the status of
endangered birds in the British Virgin Islands.

Mammals

The only native mammals now known from St. John are five
species of bats (fruit bat, Artibeus jamaicensis; fisherman bat,
Noctilos leporinus; red fruit bat, Stenoderma rufum; cave bat,
Brachyphylla cavernarum; velvety free-tailed bat, Molossus
molossus). The Brazilian free-tailed bat is present but not
known to breed (Philibosian & Yntema 1977a). Hayward (1969) made
brief notes on species of bats collected at Lameshur Bay. Nellis
(1971) and McManus & Nellis (1972a, 1972b) studied bats found on
St. John, although their work was done in other locations. An
extinct Capromyid rodent, the Antillean Hutia (Gsolobodon
portoricensis), is known from bones in aboriginal kitchen
middens.

Five other species of wild mammals found on St. John today
were introduced by man. These are the roof rat (Ratus rattus),
the Norway rat (Rattus norvegicus), the house mouse (Mus
musculus), the white-tailed deer (Oedocoilus virginianus), and
the Indian mongoose (Herpestes auropunctatus). These animals
have all established breeding populations on the island, and the
mongoose in particular has attained large population sizes and
ubiquitous distribution. In addition there are feral populations

of a few domestic animals such as pigs, cats, and burros. Of
these the burros have by far the largest numbers. Other
publications dealing with Virgin Islands mammals are Lazell
(1980, 1983b), Seaman (1949-1963), Philibosian & Yntema (1977b),
and LaBastille & Richmond (1973).

Coblentz (1983) studied the impact of exotic animals in VINP
and reported that the abundant burros and mongooses clearly exert
a significant influence on the island's biota. Burros have a
detrimental impact on vegetation and increase erosion through
their trail-forming activities. Mongooses are destructive to
many small ground-dwelling native vertebrates and invertebrates
(Coblentz & Coblentz 1985a, Seaman 1952, Seaman & Randall 1962,
Nellis 1979, Nellis & Small 1983), and also prey on eggs and
hatchlings of the endangered hawksbill turtle (Small 1982).
Lazell (1980) also discusses the impact of feral and domestic
goats on vegetation.

Coblentz & Coblentz (1985b) studied the reproduction of the
mongoose in St. John. Seaman (1952) and Seaman & Randall (1962)
discussed mongoose predation on wildlife in the Virgin Islands,
as did Nellis (1979) and Nellis & Small (1983). Nellis & Evarard
(1983) presented a monographic treatment of the biology of the
mongoose in the Caribbean. A study of the distribution, home
ranges, behavior, physical adaptations, food habits, and
population biology of feral burros in St. John, including
management options, was recently carried out by biologists from
the Virgin Islands Division of Fish and Wildlife. A population
size of about 300 was estimated by these workers (Nellis et al.
in prep.), and exclosures were used to assess impact on
vegetation. Radio tracking techniques were employed to monitor
movements.

Coblentz & Coblentz (1985a) tested the feasibility of short
duration (5 day) high-intensity trapping to control mongooses in
localized sensitive areas, in particular near sea turtle nesting
beaches. They estimated that an average of 86-87% of the
trappable mongooses in localized areas could be removed in 5
days, and stated that hawksbill turtles nesting at beaches where
trapping was done experienced no known loss of eggs or young,
compared to as much as a 23% loss to mongooses previously (Small
1982). They recommended such a trapping program as a
satisfactory alternative to complete eradication of mongooses,
which is preferable but difficult and costly. VINP Resource
Management staff are currently trapping mongooses near park
beaches.

STRESSES TO TERRESTRIAL SYSTEMS

Originally, St. John was almost completely forested, but
approximately 90% of the forests were removed during the
plantation era in the 1700's and 1800's. AS slavery was
abolished, large plantations gave way to small-scale intensive
farming and grazing. As this agricultural phase came to a close,
the natural vegetation began to recover. Virgin islands National
Park now protects many of the resultant secondary forests which
are representative of an ecosystem in which humans have had
significant impact (Tyson 1987). St. John entered a new
development phase in the late 1960's and 1970�s. At the present
time, many areas outside park boundaries (and some private lands
within the authorized boundary) are again being cleared of
vegetation to accommodate the increasing demands of development.

Soil Erosion and Runoff

As discussed previously in the section on "Stresses to
Marine Systems", sediments from upland watersheds can have
adverse impacts on the three major Caribbean coastal ecosystems--
coral reefs, seagrass beds, and mangroves. The steep topography
of most Caribbean islands, plus the high clay, slow-infiltration
capacity of many soils, can cause very high natural erosional
losses even under forested conditions (Liegel 1985). While it
appears to be true that Caribbean islands in general have a
higher population density, higher intensity of land use, lower
forest cover, and higher rates of sediment yield than continental
areas either in the Caribbean or the United States, understanding
of tropical watershed function remains relatively incomplete, and
quantitative information is scarce. Land use and the intensity
of rainfall seem to be the most important variables controlling
variations in sediment yield (Lugo 1985). Williams & Hamilton
(1982) have recently reviewed the literature on the behavior of
tropical watersheds, and Dissmeyer (1985) discusses practical
techniques for watershed management in the Caribbean.

An excellent general presentation of the theoretical and
practical aspects of soil erosion and sedimentation, as well as a
proposed sediment control program for the Virgin Islands, is
given in CH2M Hill (1979a). This report contains useful
background information on climate, soil characteristics, major
causes of erosion, and effects of sedimentation. Detailed
topographic maps showing major watersheds and nearshore marine
communities, as well as major tidal flooding areas and water
resources facilities, are included. In another report, CH2M Hill
(1979b) discuss the use of the Universal Soil Loss Equation
(USLE), developed by the U.S. Soil Conservation Service for
agricultural applications, in predicting soil erosion in the

Virgin Islands. Attempts (e.g., U.S. Department of Agriculture,
undated) have been made to adapt the USLE for tropical
conditions, but in this form it is useful mainly as a crude indi-
cator of relative erosional impact rather than an accurate
predictor of actual soil loss.

Virgin Islands soils tend to have high percentages of clay
and silt, and to be coarsely granular when dry. The soils are
hi,-hly permeable until saturated, after which they swell up and
be�come poorly permeable, with the result that runoff then
increases dramatically. The clayey topsoil has a high percentage
of colloidal-sized soil particles which are difficult to dislodge
from the soil matrix, but once dislodged, they will remain in
suspension for a very long time before settling out. Such
particles are an important factor in causing turbidity of stream
and coastal waters of the Virgin Islands (CH2M Hill 1979a).

High-intensity rainfall events in the Virgin Islands occur
in association with easterly waves, polar air, or tropical storms
and hurricanes, mostly between the months of September through
November (Bowden et al. 1970). Information on the maximum
24-hour rainfall (both observed and calculated), the probability
of occurrence for rainfalls of various intensities, and some data
on recorded stream flow are available for St. John (CH2M Hill
1979b, Cosner 1972, Jordan 1972, Robison et al. 1973). Due to
rainfall variability, small watershed area, actual
evapotranspiration-to-rainfall ratio near unity (Teytaud 1938),
and low groundwater storage capacity, streamflow on St. John is
generally meager, variable and intermittent (CH2M Hill 1979b).
There are often long periods between flows in major watercourses!
broken by periods of short and sometimes very high flows. This
large variability in streamflow causes high variability in
erosion and sediment transport to the sea. Most watercourses on
St. John are free of undergrowth indicating that while stream
flow is periodic, it is frequent enough and violent enough to
prevent the growth of an understory.

Man's land-use activities in a watershed tend to increase
the natural variation in suspended sediment by creating new
sediment sources and increasing runoff. Several studies indicate
that land use is the single most important factor affecting
sediment losses in tropical watersheds (Gottfried & Ingles 1985).
Most states have adopted water quality standards which include
limitations on the amount of sediment, usually expressed as
turbidity, that a land-use activity can add to a stream. Virgin
Islands law prohibits discharge without a permit of any pollutant
into streams but provides no standards; the definition of
"pollutant" does not specifically refer to soil or suspended
sediment, although the definition of "pollution" includes
"changes" in turbidity of any waters (Virgin Islands Department
of Conservation and Cultural Affairs 1979, pp. 17-23).

Concentrations of dissolved nutrients in streams draining
undisturbed watersheds are generally quite low. Nutrient inputs
come mainly from surface runoff, sub-surface percolation, and the
atmosphere; outputs are in the form of streamflow, groundwater
seepage, and losses to organisms and burial of organic sediments.
As in the case of suspended sediment, nutrient concentrations
tend to vary widely for different runoff events and for adjacent
watersheds with similar land uses. Other factors being equal,
nitrogen and phosphorus concentrations correlate best with land
use and cover, in particular to the proportion of agricultural
and urban land in a watershed (Marsh 1983).

Purcell (1980) studied the effects of nutrient and sediment
inputs on the plankton populations of two relatively undeveloped
bays in St. John. Two VIRMC projects have addressed the fate of
sediments after they enter the sea. As mentioned previously,
Hubbard (1987) reviewed the literature on sedimentation in
tropical marine environments and made recommendations on
watershed management and priorities for future research on
sedimentation. He lists the following land-use activities as
major causes of erosion and/or sedimentation in the Virgin
Islands: road construction, land clearing, agriculture,
alteration of watercourses, filling in of coastal ponds or
opening them to the sea, and urban development. Hubbard et al.
(1987) studied the effects of runoff on the marine environment by
calculating theoretical values for peak discharge rates and
volumes under "natural" conditions of total forest cover.

The vital role of watercourses in providing a linkage
between the upland and marine components of a coastal ecosystem,
and the importance of preserving their integrity and natural
function has been discussed by many authors (e.g., Clark 1977).
Streamside buffer strips reduce erosion, preserve the stream
channel's stability, retard runoff, trap sediments and nutrients,
maintain suitable water temperature for aquatic life, and provide
habitat for birds and other wildlife. The optimum width for a
streaniside buffer zone varies with stream width, topography, soil
type, type of impact, and other factors. Title 12 of the Virgin
Islands Code (Virgin Islands Department of Conservation and
Cultural Affairs 1979, pp. 3-4) prohibits the cutting or injury
of any vegetation within 30 feet from the center or 25 feet from
the edge of any natural watercourse. Recent research has shown
that this may not be a sufficient width of buffer area to protect
a stream from the effects of sedimentation when adjacent land is
cleared, especially on steep slopes. Several studies in the
United States (cited in Brinson et al. 1981) recommend a minimum
buffer width of 75-100 feet on each side. For the temperate
U.S., Dissmeyer (1985) gives widths varying from 25 feet at 0%
slope to 105 feet at 40% slope and 185 feet at 80% slope. Lugo
(1985) gives a recommended width of at least 165 feet for
tropical watersheds. Since most of St. John has slopes in excess
of 40% (Teyt-aud 1988), and is at times subject to more intense

rainstorms than many temperate areas, it seems clear that a 165
feet (50 m) width is not excessive. In addition to adopting this
standard within the park, cooperative agreements should be sought
with local government to observe the same standard outside park
boundaries whenever sedimentation might impact park lands or
waters.

Development

The impact of the explosively growing human population of
St. John may very soon become the overriding ecological force in
many of the island's ecosystems, as it once was during the days
of plantation agriculture. The purpose of this section is to
briefly review some of the pertinent literature on terrestrial
resource management in small islands, emphasizing an
interdisciplinary "human ecology" approach.

Towle (1985) recently reviewed the environment/development
dilemma of small islands, giving a number of case histories. He
discusses various conceptual models and methodologies developed
over the years to deal with the problems of island systems and to
bridge the gap between the natural and social sciences in
minimizing human-environment conflict. Coulianos (1980)
researched concepts for ecodevelopmental tourism for small
Caribbean islands. Towle & Teytaud (1982) prepared a useful
annotated bibliography of source materials on Caribbean island
resource management, focusing on the USVI. Clark (1977) prepared
a comprehensive general work on coastal ecosystem management, and
McEachern & Towle (1974) and Odum (1976) discuss ecological
guidelines for island development in the tropics. Westman (1985)
provides an excellent synthesis of ecology and planning.

McElroy (1978) reported on economic and social impacts of
the Virgin Islands' coastal zone management program, and Pflaum
(1980) gives data and historical trends on population, economy,
and social characteristics. Posner et al. (1981) analyzed the
economic impact of VINP on St. John and developed an economic
assessment methodology for insular areas.

Many studies have attempted to transfer the concept of
carrying capacity from animal ecology to environmental planning
and resource management, but most efforts have met with limited
success. There are a number of difficulties connected with such
use of carrying capacity concepts. First, lack of theoretical or
empirical knowledge about the interactions among the multiple
factors involved requires many value judgments and simplifying
assumptions, which are not always clearly specified. Secondly,
in human systems, many limits to growth can be technologically
circumvented, albeit at ever-increasing economic, environmental
and social costs. Finally, planners must consider at least four

different kinds of carrying capacity: ecological, relating to
human impact on the ecosystem; physical, relating to available
land area and/or scope of facilities; psycho-social, relating to
human satisfaction and societal function; and recreational,
relating to some combination of the preceding or to the lowest
threshold of any one. In each case the estimation of thresholds
of overload is essentially the process of making an intelligent
guess as to the allowable level of human use, based on the goals,
policies and objectives of the managed area. in zones where the
goal is resource protection, the thresholds will differ from
those in zones where the goal is maximization of recreational
opportunities. Since the limits to growth will be set by the
lowest threshold point, conflicts inevitably arise in multiple
use areas (Yapp & Barrow 1979).

Lewsey (1978) studied the environmental effects of tourism
development on the carrying capacity of small island systems,
using Barbados as a case study. Brookfield (1980) edited a
report on a UJNESCO/MAB study of the carrying capacity of the
small islands of Eastern Fiji, using a complex simulation model.
The government of Hawaii (Anonymous 1979) prepared a research
plan with valuable discussions of procedures and problems in the
use of carrying capacity concepts in growth management of
islands. A planning study for a workshop on Caribbean island
resource management and carrying capacity, based on the adoptive
environmental assessment and management approach of Holling
(1978), was carried out for U.S. MAB by Towle et al. (1982).

As Westman (1985) has pointed out, so far most attempts to
use the carrying capacity-concept in environmental planning have
been theoretically flawed. Some studies have simply attempted to
establish appropriate levels of use given different perceptual
and economic constraints, with no attempt to determine the true
ecological carrying capacity. Other studies have used
mathematical manipulations which are not empirically justified,
have disregarded fundamental ecological concepts, or have failed
to use detailed ecological data. Nevertheless, the application
of carrying capacity concept to environmental management remains
an important goal worthy of further research. One recent study,
by Shelby and Heberlein, published in 1986 and, entitled "Carrying
capacity in Recreation Settings" is especially useful in the
context of St. John and has some useful "lessons" for management.
Future efforts will require similar but broader empirical studies
of the nature of the interrelationships between variables, and
must take into account all different kinds of carrying capacity
in various ecological and developmental contexts and under a wide
variety of impacts affecting the "system".

RECOMMENDATIONS FOR FUTURE RESEARCH AND MANAGEMENT
OF TERRESTRIAL RESOURCES

The works of Earhart et al. (1988) and Matuszak et al.
(1987) serve as the basis for a long-term study of St. John's
forest dynamics. The New York Botanical Garden has made a
commitment to maintain their plots and carry out intensive
investigations over a 30-year period with some logistical support
from VINP. The long-term objectives include:

1. analysis of secondary forest development (including data
on recruitment, growth and mortality);
2. analysis of the influence of exotic species on forest
development;
3. evaluation of the role of major storms and other
disturbances;
4. assessment of the effects of historical land use on
forest regeneration; and
5. identification of management actions which will preserve
endangered species and promote conservation of biological
diversity.

The New York Botanical Garden also initiated work on a Flora
of St. John in 1987. Researchers will conduct a thorough
botanical survey of the entire island and prepare a manual of the
plants. The manual will include keys and illustrations of about
300 species, vegetation maps, descriptions of the ecological
characteristics of the island, a brief land use history and a
section on indigenous use of native plants. Researchers from the
University of Wisconsin have recently initiated studies of the
dry forests of St. John and plan to study regeneration,
germination of native and exotic species, influence of soil
moisture on germination, and the role of grazing animals in
forest dynamics (Brown 1988).

Studies of the terrestrial plant communities in St. John
have begun to move beyond the initial phase of cataloging and
classification into studies of ecosystem function,
development/regeneration, and response to natural and human
disturbance. much interest has been aroused within the last 10
to 15 years by the shift from the older paradigm of the natural
environment as relatively benign, free of disturbances and
supporting mainly steady-state ecosystems, to the present
viewpoint of nature as a shifting landscape of mostly
non-equilibrium patches. Such patches originate from a wide
range of natural disturbances and typically incorporate a variety
of mechanisms for recovery (Reiners 1983, Lugo 1978, Pickett &
White 1985, Forman & Godron 1981, Holling 1985). One important
consequence of this change in viewpoint is that the focus of
research is no longer on comparisons between disturbed and

undisturbed ecosystems, but rather on the effects of natural
versus human-caused disturbances.

in general, human-induced disturbances contrast sharply with
natural ones in terms of kind, scale, intensity, and frequency.
Functional studies need to be carried out for each major
vegetation type in St. John as a baseline against which to judge
successional changes and the effects of different kinds of
disturbance on the whole ecosystem (Lugo 1978, Odum 1985, Sprugel
1985, Reiners 1983). Careful studies are necessary to determine
whether there are any differences in ecosystem response to
natural disturbances versus human disturbances. The importance
of such studies lies in increasing our ability to recognize
symptoms of stress and manage ecosystems in ways that recognize
their holistic nature (Risser 1985).

At the organism and population levels, detailed information
on life histories, reproduction and mortality needs to be
collected for dominant, rare and endangered species.
observations of successional sequences initiated by various kinds
and intensities of disturbance, both human and natural, should be
made. Given enough information of this type, predictive models
of forest succession can be constructed which incorporate
variable succession pathways depending on a stand's age and
species composition when disturbed (Ewel 1980).

More information is needed on the adverse effects of
careless watershed development and their mitigation, including
practical information on stabilization of cleared hillsides and
other means of erosion control. Hubbard (1987) presents many
excellent recommendations and guidelines for development. He
notes that the following activities have severe consequences and
should be prohibited or discouraged: dredging in nearshore
waters, filling of salt ponds, removal of seagrasses and
mangroves, and development in major water courses.

GEOLOGY

St. Thomas and St. John, the two large northern islands of
the USVI, are composed of stratified volcanic and volcaniclastic
rocks with minor limestone of Early Cretaceous (Albian) to
possible Late Cretaceous age (Donnelly 1966). These rocks are
intruded by a wide range of dikes and plutons of gabbroic to
granitic composition, some of which may be as young as Tertiary
(Kesler & Sutter 1979).

The oldest rocks on St. John are submarine lavas
(keratophyre and spilite), beds of volcanic debris and chert and
associated intrusive rocks of the Water Island Formation
(Donnelly 1966) which underlie the southeastern half of the
island including the East End. Fossils in cherts of the Water
Island Formation on St. Thomas indicate that the unit is of Early
Cretaceous (Albian) age. The Water Island Formation is overlain
by andesitic volcanic and volcaniclastic rocks of the Louisenhoj
Formation (Fig. 15) which underlies much of northwestern St.
John. Donnelly (1966) suggested that the Louisenhoj Formation
was deposited unconformably on the Water Island Formation after a
period of emergence, tilting and erosion, on the slopes and
environs of a subaerial volcanic island located roughly between
St. Thomas and St. John, an area now occupied by Pillsbury Sound.
Recent work by D. Rankin (written communication, 1988) suggests
that the Louisenhoj Formation is conformable on the Water Island.
A period of quiescence followed, and limestones and calcareous
shales and sandstones of the Outer Brass Limestone were deposited
on the Louisenhoj. This unit is relatively thin (a few tens of
meters thick) and forms a band of rock underlying the lowland
from Maho Bay to Leinster Hill.

The youngest layered deposits on St. John are volcaniclastic
rocks of the Tutu Formation (Fig. 16) which make up Whistling Cay
and much of Mary Point. Fossils contained in the Tutu Formation
on St. Thomas suggest that those deposits are of Early Cretaceous
(Albian) age (Donnelly et al. 1971). Thus the volcanic and
volcaniclastic rocks of St. John all appear to have been
deposited in a relatively short period of time spanning perhaps
10 to 15 million years about 100 million years ago (D. Rankin,
written communication, 1988).

Following deposition of the Tutu Formation, the layered
rocks of St. John were folded, faulted and intruded by dikes and
plutons of gabbroic to granitic composition. The plutons are
prominent along the north coast of St. John where their heat has
recrystallized the surrounding strata. The layered rocks now
form a monocline dipping to the north with the steepest dips
along the north shore of St. John. The monocline is in turn
warped into a broad north plunging anticline whose form is more
or less indicated by the broad curve of the north shore of St.

Fig. 15. Example of the Louisenhoj Formation, south end of Trunk
Bay. Photo: D. Rankin.

Fig. 16. Bedded siltstone of the Tutu Formation, point between
Francis and Maho Bays. Photo: D. Rankin.

John from Hawksnest Bay to the East End (D. Rankin, written
communication, 1988).

The geology of the British Virgin Islands is essentially
similar to that of St. Thomas and St. John, although some
Tertiary volcanic and volcaniclastic rocks are exposed and
plutonic rocks are prominent (Helsley 1960). Anegada is a low
sandy spit formed of carbonate reef sediments.

St. Croix lies on a somewhat isolated, submerged ridge
separated from the Puerto Rico Bank by the Virgin Islands Basin.
Geologically it is related to the islands of the Puerto Rico
Bank. If St. Croix was ever connected to the northern Virgins,
it may have been separated from that group by either block
(Meyerhoff 1927, Whetten 1966) or shear faulting (Adey 1977,
Turner 1971).

The oldest rocks exposed on St. Croix are epiclastic
volcanic sandstone and mudstone of the Caledonia Formation
(Whetten 1966). These weakly metamorphosed, uplifted, folded and
faulted rocks were derived from volcanic and other narrow-trench
sediments originally deposited by turbidity currents on the deep
ocean floor about 70 to 80 million years ago (Adey 1977). Buck
Island is an emergent part of the St. Croix shelf.

Somewhat later in the Cretaceous, one or more volcanoes
formed on the sea floor to the south or southeast of St. Croix.
Volcanic debris was shed northward to form the Judith Fancy
formation, composed of tuffaceous sedimentary rocks, which occur
on St. Croix but not on Buck Island.

St. Croix was uplifted above sea level in the Oligocene
(Whetten 1974), originally as two islands. The East End Range
(including proto-Buck Island) and the Northside Range were
separated by a trough several miles wide. The trough was
subsequently filled in by the deposition of the Kingshill marl
formation. There then followed a period of mild deformation,
post-Miocene uplift, and erosion to form the present-day
topographic features.

The Virgin Islands are near the northeastern corner of the
present Caribbean Plate, a relatively small trapezoidal-shaped
plate which is moving eastward relative to the North and South
American continents carried on the American Plate. The arc of
the Lesser Antilles is an active volcanic arc above a subduction
zone in which Atlantic oceanic crust of the American Plate is
carried downward under the Caribbean Plate. The closest volcano
to the Virgin Islands of this presently active arc is Saba, about
160 km to the east. The Caribbean Plate is sliding past North
and South American plates along east-west trending northern and
southern boundaries (an extreme oversimplification) (see Dillon
et al. 1987). The western boundary is a subduction zone in which

the Cocos Plate is being driven northeastward and down under the
edge of the Caribbean Plate west of Central America.

The deep basins of the Caribbean Plate, the Colombian and
Venezuelan basins, are probably floored by oceanic crust, that
is, crust formed at a spreading ridge. Two hypotheses dominate
current thinking for the origin of the Caribbean (Dillon et al.
1987). One suggests that a branch of the Atlantic spreading
system extended to the Pacific Ocean through the Caribbean area.
By this hypothesis new oceanic crust formed between North and
South America as the two continents separated during their
westward migration from the Mid-Atlantic ridge. The other
hypothesis is that the Caribbean oceanic crust formed in the
Pacific Ocean and was carried eastward between North and South
America as they separated. According to Dillon et al. (1987) the
latter hypothesis has had the most adherents in recent years, but
the first hypothesis of ocean-opening in place, seems to simplify
many Caribbean geologic problems and is gaining support.

The present northern boundary of the Caribbean Plate extends
from the Gulf of Honduras, along the Cayman Trough and the north
coast of Hispaniola to the Puerto Rico Trench. Thus the Greater
Antilles straddle the present plate boundary. Cuba lies to the
north and the rest of the Greater Antilles lie to the south
within the Caribbean Plate. The origin of the Greater Antilles
is closely tied to the site of origin of the Caribbean oceanic
crest.

One group of authors believes that the Greater Antilles
evolved as a mid-Caribbean archipelago which never had any close
connections with Central America (Pregill 1981). These islands
are thought to have resulted from volcanism due to subduction of
oceanic lithosphere along a broad arc extending from western
Central America east to Puerto Rico, beginning in the late
Cretaceous. The islands became fully uplifted and continuously
emergent sometime in the late Oligocene or early Miocene (about
23 million years ago). The Lesser Antilles formed much more
recently by volcanism at the eastern plate boundary.

In contrast, another group of authors believes that the
Greater Antilles are the remnants of a "proto-Antillean"
archipelago which originated as a series of volcanic islands
between southern Mexico and South America during the first half
of the Cretaceous, 135 to 90 million years ago (Coney 1982, Rosen
1975). Beginning about 90 million years ago this archipelago was
rafted eastward with the relative movement of Caribbean
lithosphere. The northernmost islands became the backbone of
Cuba, the central ones other parts of the Greater Antilles, and
the southernmost elements became the coastal range of Venezuela
and the islands of Aruba, Bonaire, and Curacao. Some fragments

7 2

may also have been displaced to the region of the Lesser
Antilles. By the Eocene (40 million years ago) the core of the
Greater Antilles had reached their present positions and a new
archipelago, the backbone of present-day Central America, had
formed along the trailing edge of the Caribbean plate. An arc of
new volcanic islands -- the forerunners of the Lesser Antilles --
had appeared at the leading edge of the Caribbean plate as it
dived under the adjacent plate. During the last 40 million years
these original Lesser Antilles were consumed in the adjacent
trench. A new volcanic arc, the present-day Lesser Antilles, has
been created (Brown & Gibson 1983, Pielou 1979). At the present
time it is not clear which hypothesis on the evolution of the
Greater Antilles is most nearly correct, but the question has
important implications for the Virgin Islands.

BIOGEOGRAPHY

The different interpretations of Caribbean plate tectonic
history have spawned a lively controversy regarding the reality
of the hypothesized "proto-Antilles" and their role in the
distribution of the Greater Antillean biota. Some authors (e.g.,
Pregill 1981) argue for continued acceptance of the classical
view that all organisms which colonized these islands did so by
some form of long-distance overwater dispersal. Others (e.g.,
Rosen 1975) attribute the origin of the entire biota to transport
from the region of central America aboard the drifting islands of
the "proto-Antilles."

MacFadden (1980, 1981) emphasizes that the two "opposing"
biogeographic models may actually be complementary. The present
biota of the Antilles could be the result of an early plate
tectonic rafting event "over-printed" with numerous events of
later overwater dispersal to, and among, Caribbean Islands. A
critical, and at present unresolved issue in the controversy is
whether large land masses in the Greater Antilles have been
continuously above sea level and capable of sustaining a flora
and fauna (Pregill 1981, MacFadden 1981). Few geological studies
have been directly concerned with just when the islands were
above water, and caution is necessary in using plate tectonic
reconstructions to support biogeographic hypotheses (Hedges 1982,
Khudoley & Myerhoff 1971).

At the time of the Wisconsinan glacial maximum (24,000 to
17,000 years ago), eustatic sea level was lowered by as much as
120 m and the Caribbean climate was drier than at present
(Bonatti & Gartner 1973). During the previous interglacial of
65,000 years ago, however, sea level was 8-10 m above the present
level and the climate was wet. The successive glaciation events
which were responsible for cyclic sea-level fluctuations as well
as alternating wet (interglacial) and dry (glacial maximum)

climatic periods throughout the Pleistocene have greatly
influenced the distribution of species and the composition of
Caribbean biotas. During glacial periods, grasslands, savannas,
and scrub forests predominated in the lowlands of Puerto Rico and
the emergent bank to the east (Pregill & Olson 1981). All the
islands on the Puerto Rico Bank were connected to each other by
dry land at such times, facilitating the exchange of species
especially those adapted to the widespread xeric conditions.

The Puerto Rico Bank became fragmented by rising sea levels
about 7,000 years ago, at which time the Virgin Islands were
isolated from each other and from Puerto Rico (Heatwole & Mac
Kenzie 1967). The rise in sea level was accompanied by a return
to more mesic conditions, which led to the formation of isolated
relict populations of such formerly widespread species as the
iguana Cyclura pinguis on Anegada and the endemic toad
Peltophyrne lemur in xeric areas of Puerto Rico and possibly
Virgin Gorda (Pregill & Olson 1981). Many land vertebrates
became extinct in the Greater Antilles at this time, presumably
as a result of climate and habitat changes in conjunction with
the shrinking areas of islands. The ranges of various vegetation
types would also have been affected, with an increase in mesic
formations at the expense of xeric types.

The Pleistocene glaciations caused sea-surface temperatures
to drop by at least 3-4 degrees Celsius (Pregill & Olson 1981).
This cooling of the Caribbean and the accompanying lowering of
sea level would have resulted in a contraction of the coral reef
communities. Seventeen thousand years ago the shoreline was at
least 120 m lower, and the broad flat island shelves of today
were dry land. The new shorelines consisted of the very steep
slopes that surround most islands today at 120 m depths. The
area suitable for reefs during these glacial periods was probably
10% or less of that available today (Colin 1978).

Pregill & Olson (1981) conclude that because the Wisconsinan
glaciation had such a profound effect on the distribution of West
Indian biota, it can be assumed that earlier glacial/interglacial
cycles in the Pleistocene had similarly drastic effects.
Repeated events of faunal and floral isolation, speciation, and
extinction would have occurred, with the result that relict
distributions would be superimposed on one another through time,
complicating many aspects of Antillean biogeography. Even
farther back in time, the exceedingly complex tectonic history of
the Greater Antilles adds yet another layer of intricacy to
distributional patterns. These authors therefore caution that
any attempt to interpret the biogeography of individual organisms
or the species composition of islands without accounting for
their history is bound to have limited success. This warning is
mainly directed towards the many ecological studies inspired by
MacArthur & Wilson's (1967) Theory of Island Biogeography, which
attempts to explain the numbers of species on islands in terms of

immigration/extinction equilibria. Recent summaries of
biogeographic theory, including island biogeography can be found
in Brown & Gibson (1983), Pielou (1979), Williamson (1981), and
Endler (1982).

RECENT GEOLOGICAL RESEARCH IN ST. JOHN

Rankin (1984, 1985a, 1985b, and work in progress) has
recently conducted geological fieldwork in St. John in connection
with a U.S. Geological Survey project to produce geologic
quadrangle maps of the USVI at a scale of 1:24,000. Chemical
analyses have been performed on rock samples from St. John to
characterize the igneous activity that formed the rocks and to
determine the tectonic setting in which they formed. Rankin's
field work confirms the stratigraphic succession up through the
Tutu formation and the general distribution of units as worked
out by Donnelly (1966), but he has found that a considerable
revision of the geological contacts and structural trends
portrayed by Donelly may be necessary.

Results of the geochemical analyses on St. John done as part
of a mineral resource appraisal of the USVI by the U.S.
Geological Survey (Tucker et al. 1985) indicate an unusual
concentration of heavy metals in some rocks. Concentrations of
copper, lead, iron, tin, barium, zinc, and precious metals were
locally anomalously high. This finding has raised questions
about the concentrations of these potentially toxic substances in
runoff and their possible effects on marine systems. Trace
metals enter surface waters as a result of bedrock weathering.
They are toxic to marine organisms above a certain level although
some are essential for metabolic processes at lower
concentrations. The natural erosion of rocks would not be
expected to adversely affect marine organisms. However, man's
activities could accelerate rates of erosion and/or add
pollutants to runoff. Ramos-Perez et al. (1988) analyzed
saltwater, stream water and marine sediments from Fish, Greater
Lameshur, Bordeaux, Coral and Reef Bays for iron, manganese,
magnesium, copper, chromium, nickel and zinc. The objective of
their baseline study was to determine the contribution of heavy
metals to near-shore marine ecosystems. All water and sediment
samples showed low metal concentrations typical of unpolluted
environments.

CONCLUSIONS

The National Park Service has the responsibility and the
obligation to manage Virgin Islands National Park not only as a
national park but as a biosphere reserve. The UNESCO Action Plan
for Biosphere Reserves (1984) lists several objectives including
establishment of research facilities and a research program;
establishment of monitoring procedures; compilation of baseline
inventories; preparation of a history of research; development of
a training/education program; and preparation of a management
plan specifiying steps to be taken in fulfilling biosphere
reserve functions.

The National Park Service has made a strong commitment to
the biosphere reserve program, particularly in its support of
research by VIRMC. However, much remains to be done in
developing a fully functioning biosphere reserve. The challenge
is to take the information which is available and apply it to
achieve a successful balancing of the increasing pressures from
the island's rapid development and tourism on the natural
resources with conservation of these resources. Local residents
must be integrated into management of the biosphere reserve. The
General Management Plan for VINP should be revised or updated to
identify a core area for the biosphere reserve which would
include the Lameshur and Reef Bay watersheds. This core area
would be strictly protected. Awareness of park regulations and
environmental concerns should be increased through continuation
of the series of seminars at the Virgin Islands Biosphere Reserve
Center which began in 1987 and through further development of
interpretive programs (snorkel trips, evening programs).
Additional fliers on protection of natural resources, such as the
new one on anchoring and snorkeling, could be developed.
Research by VIRMC has focused attention on the degradation of
marine and terrestrial ecosystems in VINP. VIRMC and VINP should
work more effectively with the Coastal Zone Management Division
to help control development of St. John and its consequences for
the marine and terrestrial systems, for example in developing
comprehensive land and water use plans which are still lacking
for the islands.

7 6

WATER MANAGEMENT PLAN-BASE MAP

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U.S. VIRGIN ISLANDS
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APPENDIX III

TECHNICAL PUBLICATION SERIES

Beets, J., Lewand, L. & Zullo, E. 1986. Marine community
descriptions and maps of bays within the Virgin Islands National
Park/Biosphere Reserve. Report #2.

Beet, J. & Lewand, L. 1986. Collection of common organisms within
the Virgin Islands National Park/Biosphere Reserve. Report #3.

Anderson, M., Lund, H., Gladfelter, E. & Davis, M. 1986.
Ecological community type maps and biological descriptions for
Buck Island Reef National Monument and proposed marine park sites
in the British Virgin Islands. Report #4.

Lund, H., Anderson, M., Gladfelter, E. & Davis, M. 1986. Trends
in recreational boating in the British 'Virgin Islands: A
preliminary assessment of impact from human activities on
anchorages and development of a monitoring program for safe
anchorages. Report #5.

Davis, M., Gladfelter, E., Lund, H. & Anderson, M. 1986.
Geographic range and research plan for monitoring white band
disease. Report #6.

Gladfelter, E., Anderson, M., Lund, H. & Davis, M. 1986. Marine
ecosystems of the Lesser Antilles: Identification of
representative sites. Report #7.

Boulon, R. 1986. Map of fishery habitats within the Virgin
Islands Biosphere Reserve. Report #8.

3oulon, R. 1986. Fisheries habitat of the Virgin Islands region
of ecological importance to the fishery resources of the Virgin
Islands Biosphere Reserve. Report #9.

Dammann, A. 1986. Assessment of fish and shellfish stocks
produced in the biosphere reserve. Report #10.

Boulon, R. & Clavijo, I. 1986. Utilization of the Virgin Islands
Biosphere Reserve by artisanal fishermen. Report #11.

Koester, S. 1986. Socioeconomic and cultural role of fishing and
shellfishing in the Virgin Islands Biosphere Reserve. Report #12.

Boulon, R., Beets, J. & Zullo, S. 1986. Long-term monitoring of
fisheries in the Virgin Islands Biosphere Reserve. Report #13.

Goodwin, M. 1986. Characterization of Lesser Antillean fisheries.
Report #14.

Putney, A. 1987. Data synthesis and development of a basis for
zoning of the Virgin Islands Biosphere Reserve. Report #15.

Putney, A. 1987. Conceptual framework for the management of the
Virgin Islands Biosphere Reserve. Report #16.

Rogers, C. & Zullo, E. 1987. Initiation of a long-term monitoring
program for coral reefs in the Virgin Islands National Park.
Report #17.

Knausenberger, W., Matuszak, J. & Thomas, T. 1987. Herbarium of
the Virgin Islands National Park: Consolidation and curation of a
reference collection. Report #18A.

Matuszak, J., Craft, E. & Knausenberger, W. 1987. Establishment
and soil characterization of long-term forest monitoring plots in
the Virgin Islands Biosphere Reserve. Report #18B.

Tyson, G. 1987. Historic land use in the Reef Bay, Fish Bay and
Hawksnest Bay watersheds, St. John, U.S. Virgin Islands, 1718-
1950. Report #19.

Hubbard, D. 1987. A general review of sedimentation as it relates
to environmental stress in the Virgin Islands Biosphere Reserve
and the Eastern Caribbean in general. Report #20.

Hubbard, D., Stump, J. & Carter, B. 1987. Sedimentation and reef
development in Hawksnest, Fish and Reef Bays, St. John, U.S.
Virgin Islands. Report #21.

Boulon, R. 1987. A basis for long-term monitoring of fish and
shellfish species in the Virgin Islands National Park. Report
#22.

Nichols, M. & Brush, G. 1988. Man's long-term impact on
sedimentation: Evidence from salt pond deposits. Report #23.

Rogers, C., McLain, L. & Zullo, E. 1988. Recreational uses of
marine resources in the Virgin Islands National Park and
Biosphere Reserve: Trends and consequences. Report #24.

Williams, S. 1988. Assessment of anchor damage and carrying
capacity of seagrass beds in Francis and Maho Bays for green sea
turtles. Report #25.

Tobias, W., Telemaque, E. & Davis, M. 1988. Buck Island fish and
shellfish populations. Report #26.

Earhart, J., Reilly, A. & Davis, M. 1988. Initial inventory of
three permanent forest plots in the Virgin Islands National Park.
Report #27.

Ramos-Perez, C. & Gines-Sanchez, C. 1988. Geochemistry of St.
John and influence on marine systems. Report #28.

Rogers, C. & Teytaud, R. 1988. Synthesis of selected resource
management information for Virgin Islands National Park and
Biosphere Reserve. Report #29.

APPENDIX IV

REFERENCES CITED

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Adey, W. 1977. Field guidebook to the reefs and reef communities
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Miami.

American Ornithologists Union 1983. Check-list of North American
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Anderson, M., et al 1986. Ecological community type maps and
biological community descriptions for Buck Island Reef National
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Anonymous 1979. An approach for developing, assessing, and
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Askins, R., Ewert, D. & Norton, R. 1987. Effect of habitat
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Beard, J. 1944. Climax vegetation in tropical America. Ecology
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Beard, J. 1945. Forestry in the Leeward Islands: The British
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Beard, J. 1949. Natural vegetation of the Windward and Leeward
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Beard, J. 1955. The classification of tropical American
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Beatty, H. 1944. Fauna of St. Croix, V. I. Jour. Agri. Univ. of

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Beets, J. & Lewand, L. 1986. Collection of common organisms
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Beets, J., Lewand, L. & Zullo, E. 1986. Marine community
descriptions and maps of bays within the Virgin Islands National
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Birkeland, C., Reimer, A. & Young, J. 1976. Survey of marine
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108

INDEX

Agricultural practices 6, Christopher Columbus 6
9, 23, 53, 54, 62, 64, 65 Ciguatera 37
Algal plains 18, 36, 42 Cinnamon Bay 21, 27, 29,
Amphibians 55, 56 49
Anchor damage 14-15, 23-26 Coastal Zone Management
Anegada 3, 48, 55, 60, 71, Division 65, 76
74 Cocoloba beach 30
Anegada Passage 3 Conch 10, 13, 32, 33, 35-
Aquatic fauna 55-56 40, 42-46
Arawaks 6 Coral Bay 15, 27, 38, 56,
Archaeology 6 75
Bank pavement 25, 36, 38 Coral bleaching 28-29
Barracudas 17, 37 Coral diseases 27-28
Beaches 13, 18-20, 23, 26, Black Band Disease 28
29-30, 32, 61 White Band Disease 27-28
Beachrock 18 Coral growth 9, 22-23
Benthic microalgae 10 Coral recruitment 10
Biogeography 54, 73-75 Coral reefs 8-13, 15, 17,
Birds 15, 18, 20, 55-60, 20-21, 25, 26-29, 34, 46,
64 62, 74
Bridled Quail Dove 58, Artificial reefs 35
59 Bank reefs 10, 38
Brown Pelican 15, 57 Barrier reefs 10
Lesser Antillean Bull- Fringing reefs 10, 17,
finch 57 35
Roseate Tern 57 Patch reefs 10, 15, 22,
Stolid Flycatcher 57 38
Black sea urchins (Diadema Corals
antillarum) 14, 29, 42 Black coral (Anti-
Blue-green algae 28 patharians) 31
British Virgin Islands Brain coral (Diploria
(see BVI) spp.) 10, 28
Brown Bay 27 Elkhorn coral (Acropora
Buck Island 9, 19, 28-30, palmata) 10, 23, 25, 27,
32, 43, 53, 71 28
Buck Island Reef National Fire coral (Millepora
Monument (see BUIS) spp.) 10, 29
Buffer strips 64 Lettuce coral (Agaricia
BUIS 1, 28, 43 spp.) 10, 13
Burros 61 Crabs 17, 20, 42, 55, 56
BVI 9, 28, 32, 33, 35, 37, Ghost Crabs 20
44, 48, 52, 55, 60 Hermit Crabs 42
Calcareous algae 13 Land Crabs 20
Halimeda 13 Cruz Bay 5, 8, 21, 53
Penicillus 13 Cuba 72
Cambridge Ornithological Damselfish 10
expedition 60 Danish Colonization 6
Caneel Bay 8, 15 DCCA (see DPNR)
Caiibbean Plate 71-73 Dept. Conservation &
Cayman Trough 72 Cultural Affairs (see
Chocolate Hole 8, 13 DPNR)

109

Dept. Planning & Natural Greater Antilles 3, 58,
Resources (see DPNR) 72-74
Development vi, vii, 2, 9, Groundwater 54, 63, 64
13, 18, 21-23, 36, 38, 49, Grunts 10, 13, 36
62, 64-66, 68, 76 Haulover Bay 31
DPNR 8, 63, 64 Hawksnest Bay 10, 13, 19,
Dredging 20, 38, 68 21, 22, 25, 36, 39-41
Dry Tortugas 41, 42, 46 Hawksnest Reef 25
Eagle rays 39 Hawksnest watershed 13,
Echinoids 42 49, 52, 54
Lytechinus 42 Herbivory 14, 18
Tripneustes 42 History 1, 6-7
Ecosystems vi, vii, 1, 2, History of research
9, 10, 13, 15-17, 21, 34, Fauna 54-61
43, 46, 53, 67, 68, 75, 76 Flora 48-49
Endangered and threatened Marine systems 8-9
species 29-31, 34, 50, 55, Hurricane Hole 38
57, 60, 61, 68 Hurricanes (see Storms)
Erosion 53, 61, 62-65, 68, Intertidal zone 20
69, 75 Jacks 37
Europa Bay (see Lameshur Johnson's Reef 37
Bay) Jost van Dyke 48
Fireworm 29 Jumbie Bay 27
Fish Bay 13, 22, 27, 36, Juvenile fish 9, 15, 36
39, 49, 56 Lameshur Bay 8, 19, 29,
Fish community structure 35-37, 55
35-36 Europa Bay 42
Fish grazing 9-10, 15 Greater Lameshur Bay 8,
Fish poisoning (see 18, 42, 75
Ciguatera) Little Lameshur Bay 8
Fish stocks 32, 34-35, 38, Land birds 58-59
44 Land use 18, 23, 52, 54,
Fish traps 32-34, 37, 38, 62-64, 67
42, 43, 45 Leinster Bay 10, 25
Fisheries 32-45 Leinster Hill 69
Cultural role 32-34 Lesser Antilles vi, vii,
Socioeconomic role 32-34 3, 32, 43, 48, 52, 58, 72-
Fisheries landings 36-37 73
Fisheries regulations 40, Lesser Antillean fisheries
43, 44-45 32, 43-44
Fishery exports 38 Lobsters 8, 15, 32-33, 35-
Francis Bay 14, 17, 25, 41 43, 45
Freshwater habitats 56 Spiny lobster (Panulirus
Geochemistry 75 argus) 35, 40, 41, 42
Geology 69-73 Spotted lobster (Pan-
Gill nets 34 ulirus guttatus) 40
Goatfish 36 Long-term monitoring 32,
Gorgonians 28 36, 49
Great Cruz Bay 21 Mackerel 36
Great Thatch Island 38 Maho Bay 14, 15, 25, 69

110

Mammals 55, 56, 60-61 Puerto Rico Bank 3, 56,
Antillean Hutia 60 59, 60, 71, 74
Bats 60 Puerto Rico Trench 3, 72
Indian mongoose 30, 60- Queen conch (Strombus
61 gigas) (see Conch)
Rats 60 Ram Head Bay 35
White-tailed deer 60 Recreational use 8, 9, 23,
Mangroves 9, 10, 15-16, 25, 33, 42, 66
17, 23, 26, 34, 35, 36, 38, Reef Bay 6, 10, 13, 14,
41, 46, 62, 68 21, 22, 23, 27, 37, 38, 39,
Black iantjrove (Avi- 40, 49, 52, 75, 76
cennia germinans) 15, 26 Reeffish 33, 35, 36, 43,
Red mangrove (Rhizophora 44
mangle) 9, 15, 26 Rendezvous Bay 14
White mangrove (Lagun- Reptiles 55, 56
cularia racemosa) 15 Rocky shores 18-20
Mangrove forests 10, 15-16 Runoff 9, 15, 18, 20, 21,
Basin forests 15 22, 36, 53, 62-65, 75
Fringe forests 15 Sage Mountain 52
Marine algae 18 Salt ponds 15, 17-18, 23,
Marine mammals 31 68
Mary Point 69 Mandal Pond 17, 23
Mary's Creek 11, 13 Southside Pond 17
Mennebeck Bay 27 Sandy Point 29
Migration 9, 35, 36, 37, Sea turtles 1, 8, 9, 14,
40, 46, 75 20, 29-30, 32, 34, 44, 61
Mollusks 20, 42 Green turtles (Chelonia
Mongooses (see Mammals) mydas) 1, 14, 29, 30
Mullets 17 Hawksbill turtles
Natural stresses 27-29 (Eretmochelys imbricata)
Newfound Bay 10, 17 29, 30, 61
Norman Islands 37, 38 Leatherback turtle
Nutrients 9-10, 15, 53, (Demochelys coriacea) 29,
54, 64 30
OECS 44 Seabirds 20, 58
Oil pollution 26, 63 Seagrass 8, 9, 13, 25, 36,
Organization of Eastern 37, 39
Caribbean States (see Manatee grass (Syrin-
OECS) godium filiforme) 13
Overfishing 33, 43 Shoal grass (Halodule
Parrotfish 9, 36 wrightii) 13
Physical setting 3-5 Turtle grass (Thalassia
Rainfall 5 testudinum) 13, 15
Tides 5 Seagrass beds 8, 9, 13-14,
Topography 3 15, 20, 23, 25, 26, 34, 35,
Waves 5 39, 40, 43, 62, 68
Winds 3, 5 Sedimentation 8, 13, 20-
Pillsbury Sound 38, 69 23, 62, 64, 65
Plantation system 6, 13, Shoreline Use Plan 25-26,
22-23, 58, 62, 65 47
Protective legislation 34

111

Snappers 10, 13, 36, 37, VIRMC (see Virgin Islands)
44 Water Birds 58
Soils 6, 15, 18, 20, 22, Watersheds 9, 13, 18, 20,
26, 49, 53-54, 62-65, 67 21, 22, 23, 36, 49, 52, 54,
Soper's Hole 38 59, 62, 63, 64, 68, 76
Squirrelfish 36 Whelks 20, 32, 33, 42, 44,
St. Croix 1, 3, 5, 14, 19, 46
26, 27, 29, 40, 43, 71 Whistling Cay 27, 69
St. John clinic 13, 21, 36 Windswept Bay 27, 42
Storms 5, 8, 13, 17, 18, Windswept Reef 25-26
19, 21, 27, 49, 63, 67
Hurricane David 27
Hurricane Frederic 27
Tropical Storm Klaus 13
Subsistence fishermen 36-
37
Surgeonfish 9
Tektite Project 8, 18, 41
Terrestrial systems 48-53
Flora and vegetation 49-
53
History of research 48-
49
Threadneedle Bay 39, 40
Tourism 6, 23, 33, 34, 38,
65, 66, 76
Triggerfish 36, 39
Trunk Bay 19, 23, 29, 33,
36, 45
Turner Bay 8
U.S. possession 6
UNESCO Action Plan vi, 2,
76
VIBR (see Virgin Islands)
VIERS (see Virgin Islands)
VINP (see Virgin Islands)
Virgin Gorda 48, 74
Virgin Islands
Biosphere Reserve vi, 1,
2, 9, 32, 35, 37, 38, 43,
44
Environmental Research
Station 8, 55
National Park 1, 2, 8,
13, 21, 23, 25, 26, 28, 34,
35, 36, 37, 43, 45, 47, 49,
57, 62, 65, 67, 76
Resource Management
Cooperative vi, vii, 1, 2,
9, 21, 26, 32, 64, 76
Virgin Islands Basin 3, 71

112