• TABLE OF CONTENTS
HIDE
 Front Cover
 Acknowledgement
 Table of Contents
 Preface
 Executive summary
 Introduction
 Environmental and anthropogenic...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 The state of coral reef ecosystems...
 National summary
 Back Matter
 Back Cover






Title: The state of coral reef ecosystems of the United States and Pacific freely associated states : 2005
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Permanent Link: http://ufdc.ufl.edu/CA01300958/00001
 Material Information
Title: The state of coral reef ecosystems of the United States and Pacific freely associated states : 2005
Physical Description: Book
Language: English
Creator: National Oceanic and Atmospheric Administration
Publisher: National Oceanic and Atmospheric Administration
Place of Publication: Silver Spring, Md.
 Record Information
Bibliographic ID: CA01300958
Volume ID: VID00001
Source Institution: University of the Virgin Islands
Holding Location: University of the Virgin Islands
Rights Management: All rights reserved by the source institution and holding location.

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Table of Contents
    Front Cover
        Front Cover
    Acknowledgement
        Page i
        Page ii
    Table of Contents
        Page iii
        Page iv
        Page v
        Page vi
    Preface
        Page vii
        Page viii
        Page ix
    Executive summary
        Page 1
        Page 2
    Introduction
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
    Environmental and anthropogenic threats to coral reef systems
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
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    The state of coral reef ecosystems of the U.S. Virgin Islands
        Page 45
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        Page 90
    The state of coral reef ecosystems of Puerto Rico
        Page 91
        Page 92
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    The state of coral reef ecosystems of Navassa
        Page 135
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    The state of coral reef ecosystems of Florida
        Page 150
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    The state of coral reef ecosystems of the Flower Garden Banks and other banks
        Page 201
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    The state of coral reef ecosystems of the main Hawaiian islands
        Page 222
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    The state of coral reef ecosystems of the northwestern Hawaiian islands
        Page 270
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    The state of coral reef ecosystems of American Samoa
        Page 312
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    The state of coral reef ecosystems of the Pacific remote island areas
        Page 338
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    The state of coral reef ecosystems of the Marshall Islands
        Page 373
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    The state of coral reef ecosystems of the Federated States of Micronesia
        Page 387
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    The state of coral reef ecosystems of the Commonwealth of the Northern Mariana Islands
        Page 399
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    The state of coral reef ecosystems of Guam
        Page 442
        Page 443
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    The state of coral reef ecosystems of the Republic of Palau
        Page 488
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    National summary
        Page 508
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    Back Matter
        Page 523
    Back Cover
        Page 524
Full Text



Th Stt of Coa ReEcsytm


of th Unte Sttsad aii
Freely A oa States: 2005-




































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I Iol











Prdue by the Natina Ocai n topei d iitain
in~~ coprto wit patnr fro Feerl Stte Teriora and
Co mn welt Agnis n*h PcfcFel sscae tts






STATIONS:

station for the entire document:

'addell, J.E. (ed.), 2005. The State of Coral Reef Ecosystems of the United States and Pacific Freely Associ-
ed States: 2005. NOAA Technical Memorandum NOS NCCOS 11. NOAA/NCCOS Center for Coastal Monitor-
g and Assessment's Biogeography Team. Silver Spring, MD. 522 pp.


station for an individual chapter (example of Main Hawaiian Islands chapter):

iedlander, A.M., G. Aeby, E. Brown, A. Clark, S. Coles, S. Dollar, C. Hunter, P. Jokiel, J. Smith, B. Walsh, I.
illiams, and W. Wiltse. 2005. The State of Coral Reef Ecosystems of the Main Hawaiian Islands. pp. 222-269.
: J. Waddell (ed.), The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated
:ates: 2005. NOAA Technical Memorandum NOS NCCOS 11. NOAA/NCCOS Center for Coastal Monitoring
id Assessment's Biogeography Team. Silver Spring, MD. 522 pp.



CKNOWLEDGEMENTS:

ie production of this report would not have been possible without the participation of the people recognized
flow. Their efforts to help compile, format, edit, and review the document are very much appreciated. Particular
cognition goes to Julie Kellner, who formatted the figures and tables in the document; Aurelie Shapiro, who
eated the maps; Lynn Dancy, who editied the text for grammatical accuracy and consistency; and Kevin Mc-
ahon, who formatted the references.

iditional editorial and support staff include:
in Borowik, Kenneth Buja, Chris Caldow, Russell Callender, John Christensen, Sarah Davidson, Katherine
schelbach, Lilli Ferguson, Tracy Gill, Christine Harvey, Jamison Higgins, Tom Hourigan, Christopher Jeffrey,
atthew Kendall, Laura Letson, Kevin McMahon, Mark Monaco, Connie Moy, David Moe Nelson, Simon Pitt-
an, and Shauna Slingsby.



DR MORE INFORMATION:

)r more information about this report or to request a copy, please contact NCCOS CCMA's Biogeography Team
301-713-3028 or visit http://biogeo.nos.noaa.gov.



ISCLAIMER:

iis publication does not constitute an endorsement of any commercial product or intend to be an opinion be-
)nd scientific or other results obtained by the National Oceanic and Atmospheric Administration (NOAA). No
ference shall be made to NOAA, or this publication furnished by NOAA, in any advertising or sales promotion
iich would indicate or imply that NOAA recommends or endorses any proprietary product mentioned herein, or
iich has as its purpose an interest to cause directly or indirectly the advertised product to be used or purchased
,cause of this publication.






i\/r nhntn hv Mlilps Anc1irsnn Analytical I ahnratnrips nf Hawaii






The State of Coral Reef Ecosystems

of the United States and Pacific

Freely Associated States: 2005


Report coordinators/ editors by jurisdiction:
U.S. Virgin Islands-Christopher Jeffrey
Puerto Rico-Jorge (Reni) Garcia-Sais
Navassa-Margaret W. Miller
Florida-Kacky Andrews, Larry Nail, Christopher Jeffrey and Simon Pittman
Flower Garden Banks-Emma Hickerson
Main Hawaiian Islands-Alan Friedlander and Athline Clark
Northwestern Hawaiian Islands-Alan Friedlander and Russell Brainard
American Samoa-Christopher Hawkins
Pacific Remote Island Areas-Russell Brainard and Jim Maragos
Republic of the Marshall Islands-Shauna Slingsby
Federated States of Micronesia-Shauna Slingsby
Commonwealth of the Northern Mariana Islands-John Starmer and Erica Cochrane
Guam-Trina Leberer and Valerie Porter
Republic of Palau-Yimnang Golbuu

Additional contributors:
Andy Bruckner, John Christensen, Elizabeth Fairey, Kelly Gleason, Michelle Harmon,
Christine Harvey, Scott Heron, Tom Hourigan, Christopher Jeffrey, Julie Kellner, Ruth Kelty,
Gang Liu, Mark Monaco, Joel Murray, Simon Pittman, Steve Rohmann, Aurelie Shapiro,
Shauna Slingsby, Dana Topousis, Shay Viehman, Jenny Waddell, Lani Watson.




NOAA Technical Memorandum NOS NCCOS 11
May 2005








United States National Oceanic and
Department of Atmospheric Administration National Ocean Service
Commerce

Carlos M. Gutierrez Vice Admiral Conrad C. Richard W. Spinrad, Ph.D.
Secretary Lautenbacher, Jr. USN (Ret.) Assistant Administrator
Administrator






TABLE OF CONTENTS


Introductory Information
Citations i
Acknowledgements i
Table of Contents iii-vi
Preface vii-ix

Chapter 1: Executive Summary 1


Chapter 2: Introduction 3


Chapter 3: Environmental and Anthropogenic Threats to Coral Reef Ecosystems 12
Andy Bruckner, Ken Buja, Liz Fairey, Kelly Gleason, Michelle Harmon, Scott Heron, Tom Hourigan,
Chris Jeffrey, Julie Kellner, Ruth Kelty, Bob Leeworthy, Gang Liu, Simon Pittman, Aurelie Shapiro,
Al Strong, Jenny Waddell, and Peter Wiley.

Climate Change and Coral Bleaching 13
SDiseases 16
Tropical Storms 17
Coastal Development and Runoff 19
SCoastal Pollution 21
STourism and Recreation 22
SFishing 23
STrade in Coral and Live Reef Species 26
Ships, Boats and Groundings 27
SMarine Debris 29
Aquatic Invasive Species 30
Security Training Activities 32
Offshore Oil and Gas Exploration 33
SOther 35


Chapter 4: The State of Coral Reef Ecosystems of the U.S. Virgin Islands 45
Christopher F G. Jeffrey, Ursula Anlauf, James Beets, Sheri Caseau, William Coles, Alan M. Friedlander,
Steve Herzlieb, Zandy Hillis-Starr, Matthew Kendall, Violeta Mayor, Jeffrey Miller, Richard Nemeth,
Caroline Rogers, and Wesley Toller.

Environmental and Anthropogenic Stressors 47
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 57
Water Quality 59
Benthic Habitats 62
Associated Biological Communities 73
Current Conservation Management Activities 80
Overall Conclusions and Recommendations 83


Chapter 5: The State of Coral Reef Ecosystems of Puerto Rico 91
Jorge (Reni) Garcia-Sais, Richard Appeldoorn, Andy Bruckner, Chris Caldow, John D. Christensen,
Craig Lilyestrom, Mark E. Monaco, Jorge Sabater, Ernest Williams, and Ernesto Diaz.

Environmental and Anthropogenic Stressors 94
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 105
Water Quality 108
Benthic Habitats 109
Associated Biological Communities 120
Current Conservation Management Activities 126
Overall Conclusions and Recommendations 127






Chapter 6: The State of Coral Reef Ecosystems of Navassa 135
Margaret Miller, Joseph Schwagerl, David McClellan, Mark Vermeij, Dana Williams.

Environmental and Anthropogenic Stressors 136
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 140
Water Quality 141
Benthic Habitats 141
Associated Biological Communities 144
Current Conservation Management Activities 148
Overall Conclusions and Recommendations 148


Chapter 7: The State of Coral Reef Ecosystems of Florida 150
Katherine Andrews, Larry Nall, Chris Jeffrey, and Simon Pittman, eds.

Environmental and Anthropogenic Stressors 153
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 165
Water Quality 165
Benthic Habitats 170
Associated Biological Communities 177
Current Conservation Management Activities 186
Overall Conclusions and Recommendations 192


Chapter 8: The State of Coral Reef Ecosystems of the Flower Garden Banks and Other Banks
of the Northwestern Gulf of Mexico 201
Emma L. Hickerson and G.P Schmahl.

Environmental and Anthropogenic Stressors 204
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 209
Water Quality 211
Benthic Habitats 212
Associated Biological Communities 216
Current Conservation Management Activities 218
Overall Conclusions and Recommendations 219


Chapter 9: The State of Coral Reef Ecosystems of the Main Hawaiian Islands 222
Alan Friedlander, Greta Aeby, Eric Brown, Athline Clark, Steve Coles, Steve Dollar,
Cindy Hunter, Paul Jokiel, Jennifer Smith, Bill Walsh, Ivor Williams, and Wendy Wiltse.

Environmental and Anthropogenic Stressors 224
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 243
Water Quality 245
Benthic Habitats 247
Associated Biological Communities 253
Current Conservation Management Activities 259
Overall Conclusions and Recommendations 262


Chapter 10: The State of Coral Reef Ecosystems of the Northwestern Hawaiian Islands 270
Alan Friedlander, Greta Aeby, Russell Brainard, Athline Clark, Edward DeMartini, Scott Godwin,
Jean Kenyon, Randy Kosaki, Jim Maragos, and Peter Vroom.

Environmental and Anthropogenic Stressors 272
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 282
Water Quality 284
Benthic Habitats 288
Associated Biological Communities 297
Current Conservation Management Activities 306
Overall Conclusions and Recommendations 307



































)f the Marshall Island!



ndition


veraii Lonclusions ana Kecommenaations ,0






Chapter 16: The State of Coral Reef Ecosystems of Guam 442
Val Porter, Trina Leberer, Mike Gawel, Jay Gutierrez, David Burdick, Victor Torres, and Evangeline Lujan.

Environmental and Anthropogenic Stressors 445
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 458
Water Quality 461
Benthic Habitats 462
Associated Biological Communities 468
Current Conservation Management Activities 476
Overall Conclusions and Recommendations 481


Chapter 17: The State of Coral Reef Ecosystems of the Republic of Palau 488
Yimnang Golbuu, Andrew Bauman, Jason Kuartei, and Steven Victor.

Environmental and Anthropogenic Stressors 490
Coral Reef Ecosystem Data Gathering Activities and Resource Condition 496
Water Quality 497
Benthic Habitats 498
Associated Biological Communities 501
Current Conservation Management Activities 503
Overall Conclusions and Recommendations 505


Chapter 18: National Summary 508






PREFACE


The purpose of this report is to provide an assessment of the current condition of coral reef ecosystems in U.S.
jurisdictions, including the U.S. Virgin Islands, Puerto Rico, Navassa, Florida, Flower Garden Banks and other
banks of the Gulf of Mexico, Hawaii, the Northwestern Hawaiian Islands, American Samoa, the Pacific Remote
Island Areas, Guam, and the Commonwealth of the Northern Mariana Islands (CNMI). The report also provides
an examination of coral reefs in the Pacific Freely Associated States (FAS), including the Republic of the Marshall
Islands, Federated States of Micronesia, and Republic of Palau. The report focuses primarily on shallow-water
portions of these states and territories, from the shoreline to the maximum depth at which sunlight-dependent
corals can survive. Coral communities occurring in deep and cold waters are the subject of a complementary
report currently under development.

This report is the second in a series of national coral reef ecosystem status reports. The initial report, The State
of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2002 (Turgeon et al., 2002),
is similar to this report in that it incorporates the work of many scientists and managers from across the world.
The first report provided a broad introduction to and a preliminary look at the status of coral reef ecosystems and
was based primarily on qualitative information from the contributing authors. The initial report also included a
considerable amount of background information that is not included in this report.

The lead entity coordinating the development of this report was the National Oceanic and Atmospheric Adminis-
tration's (NOAA) Center for Coastal Monitoring and Assessment's Biogeography Team (CCMA-BT), which is part
of the National Centers for Coastal Ocean Science. CCMA-BT scientists are responsible for three main tasks
related to coral reef ecosystem conservation: 1) administration of a Federal grant program that supports selected
monitoring efforts in U.S. jurisdictions and the FAS; 2) collection of standardized monitoring data in several U.S.
jurisdictions through a well-established scientific field program; and 3) systematic production of benthic (sea
floor) habitat maps depicting the spatial extent of the primary habitats comprising U.S. coral reef ecosystems.
CCMA-BT was assisted in this reporting effort by NOAA Fisheries' Office of Habitat Conservation and NOAA's
Coral Reef Conservation Program.

This report differs from the 2002 status report in several ways. The current report is based primarily on the
analysis of monitoring data collected by scientists rather than qualitative assessments of ecosystem conditions.
It utilizes the most recent monitoring data from all available sources, including but not limited to the activities sup-
ported by the grant program mentioned above. This report also includes a mapping component, which provides
an analysis of the spatial extent of coral reef ecosystem habitats within each jurisdiction based on the estimated
area in nearshore waters to 20 meters of water depth. It is critical to keep in mind that the term 'coral reef eco-
systems' includes not only the coral reefs themselves, but also the associated habitats that are functional compo-
nents of the ecosystem, such as mangroves, seagrass and macroalgae beds, and unconsolidated sediments.

Because the chapters reflect the hard work and dedication of writing teams from each jurisdiction, the teams
should be cited as primary authors for the jurisdictional chapters of this report. Over 160 individuals from 14
jurisdictions contributed to this report, providing not only their time, attention, and hard work, but also in many
cases, unpublished data that would otherwise not be available to the public at this time. The writing teams were
assembled by each jurisdiction's report coordinators, who deserve praise for undertaking the daunting task of
identifying and coordinating writing teams, arranging meetings, assigning tasks, assembling data sets, filling
information gaps, and responding to requests from the report editor. The report would not be possible without
their coordination efforts.

To assist in the challenging task of assimilating data and study results from 14 jurisdictions spanning 16 time
zones, CCMA-BT scientists held two regional workshops in the spring of 2003-one in Saipan, CNMI and one
in San Juan, Puerto Rico. Coordinators and authors from each of the jurisdictions attended the meetings and
helped develop a report outline that would provide a common structure to guide chapter development. The
coordinators, many of whom are the designated point of contact for all coral reef activities in their area, then
assembled a writing team of coral reef ecosystem experts from academic, non-governmental, state, territorial,
and Federal organizations. These teams were tasked with compiling an inventory of current monitoring efforts in
their jurisdiction to determine which data sets should be used to assess ecosystem status within the established
reporting structure. Subsequently, each team summarized the available data and provided a quantitative assess-






ment of the condition of the ecosystem based on three broad themes: water quality, benthic habitats, and associ-
ated biological communities. When considered altogether, these themes provide a basis for assessing overall
condition and diagnosing potential contributing factors to threatened and impacted ecosystems.

Ongoing agency efforts to assess and monitor elements of coral reef ecosystems form the basis for this report.
However, it is important to realize that monitoring data are rarely collected in the same way or with the same
frequency. Indeed, methods differ considerably among jurisdictions. These differences preclude the comparison
of data or important metrics across jurisdictions in the National Summary section of this report. Instead, conclu-
sions drawn across jurisdictions are limited to whether a particular attribute is being measured and whether these
measurements result in data that are sufficiently robust to illuminate trends or patterns. Therefore, the condition
of coral reef ecosystems within each jurisdiction is evaluated independently and is not comparable to other juris-
dictions. Unless all of the jurisdictions implement a standard protocol, it is unlikely that interjurisdictional compari-
sons can ever be made with any scientific rigor. A few agencies have already initiated a standard complement
of monitoring activities across multiple jurisdictions in an attempt to address this problem. If met with success,
these integrated programs may aid coral resource managers throughout the U.S. and FAS in the development of
a common set of diagnostic tools to help affect positive change in coral reef ecosystems.

This report is structured to provide information according to the primary threats, topics, and goals outlined in the
National Coral Reef Action Strategy (NCRAS; NOAA, 2002) and other guidance documents developed by the
U.S. Coral Reef Task Force (USCRTF) and its member organizations. Following the Executive Summary, which
distills general conclusions from the entire document, an introductory chapter provides background information
about the distribution of coral reef ecosystems in the U.S. and FAS, the different types of reefs that occur in these
areas, and an estimate of the potential extent of coral reef ecosystems (including reefs, seagrass and macroal-
gae beds, sand patches, etc.) for each jurisdiction. The third chapter summarizes the current understanding of
the 13 key natural and anthropogenic threats to coral reef ecosystems that were identified in the NCRAS. An ad-
ditional 'other' threat category was included to allow writing teams to characterize threats that may be important
or unique to a specific jurisdiction, but do not appear on the NCRAS list of key threats.

Chapters 4 through 17 comprise the heart of this report. In these chapters, the local writing teams character-
ized the current understanding of the condition of the coral reef ecosystems in their jurisdictions. Writing teams
were asked to: 1) describe the geographical distribution of reefs and provide salient background information; 2)
discuss how each of the key threats has manifested in their area; 3) describe existing monitoring programs and
identify specific data sets upon which their assessments are based; 4) present methods, results, and discussion
for each monitoring data set, organized around the three primary themes of water quality, benthic habitats, and
associated biological communities; 5) introduce the conservation and management actions currently being un-
dertaken to respond to issues of concern; and 6) provide an overall summary of the status of each jurisdiction's
coral reef ecosystems and priority recommendations for future research and management alternatives.

Finally, the National Summary chapter synthesizes and integrates the results and conclusions from each of the
preceding chapters to present broad-scale conclusions from a national perspective. The structure of the National
Summary chapter reframes the results of the jurisdiction chapters in the context of the goals identified in the
NCRAS. Grouping the information in this way clearly demonstrates how the report conclusions can help measure
progress towards overarching NCRAS goals and provide a means to evaluate the effectiveness of management
actions.

This report represents an evolving effort to determine the condition of coral reef ecosystems at both local and
national scales. To do this, scientists must ask the right questions, and then design effective studies to gather
data with sufficient frequency to confidently answer those questions. This report serves as a vehicle for the dis-
semination of information about data collection activities in the U.S. and FAS. As more monitoring data are col-
lected and analyzed, scientists will be better equipped to present time series information and provide condition
reports that address all aspects of these complex and dynamic ecosystems.

Another objective of this report is to increase the participation of scientists and managers at all levels in synthe-
sizing all available information to provide the most robust, integrated assessments possible. Data collection and
integrated reporting of information are crucial to management efforts that strive to protect and conserve coral
reefs, their associated habitats, and the organisms that depend on them. It is hoped that, through this and future






porting efforts, gaps in the current state of knowledge about U.S. coral reef ecosystems will be identified and
ed, and that the availability of up-to-date, accurate, comprehensive scientific information will enable manag-
s to slow or even halt the general decline in coral reef ecosystem health that has become evident in the last
,veral decades.




The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

EXECUTIVE SUMMARY

For over three decades, scientists have been documenting the decline of coral reef ecosystems, amid increas-
ing recognition of their value in supporting high biological diversity and their many benefits to human society.
Coral reef ecosystems are recognized for their benefits on many levels, such as supporting economies by
nurturing fisheries and providing for recreational and tourism opportunities, providing substances useful for
medical purposes, performing essential ecosystem services that protect against coastal erosion, and provid-
ing a diversity of other, more intangible contributions to many cultures. In the past decade, the increased
awareness regarding coral reefs has prompted action by governmental and non-governmental organizations,
including increased funding from the U.S. Congress for conservation of these important ecosystems and
creation of the U.S. Coral Reef Task Force (USCRTF) to coordinate activities and implement conservation
measures [Presidential Executive Order 13089].

Numerous partnerships forged among Federal agencies and state, local, non-governmental, academic and
private partners support activities that range from basic science to systematic monitoring of ecosystem com-
ponents and are conducted by government agencies, non-governmental organizations, universities, and the
private sector. This report shares the results of many of these efforts in the framework of a broad assessment
of the condition of coral reef ecosystems across 14 U.S. jurisdictions and Pacific Freely Associated States.
This report relies heavily on quantitative, spatially-explicit data that has been collected in the recent past and
comparisons with historical data, where possible. The success of this effort can be attributed to the dedication
of over 160 report contributors who comprised the expert writing teams for each jurisdiction. The content of the
report chapters are the result of their considerable collaborative efforts.

The writing teams, which were organized by jurisdiction and comprised of experts from numerous research
and management institutions, were provided a basic chapter outline and a length limit, but the content of each
chapter was left entirely to their discretion. Each jurisdictional chapter in the report is structured to: 1) describe
how each of the primary threats identified in the National Coral Reef Action Strategy (NCRAS) has manifested
in the jurisdiction; 2) introduce ongoing monitoring and assessment activities relative to three major categories
of inquiry water quality, benthic habitats, and associated biological communities and provide summary
results in a data-rich format; and 3) highlight recent management activities that promote conservation of coral
reef ecosystems.

Due to the wide variety of monitoring and assessment techniques employed by each jurisdiction, as well as
the variations in spatial and temporal resolution of the data being collected, it is necessary to evaluate each
jurisdiction independently over time and resist the temptation to compare jurisdictions. Only data collection
efforts that employ consistent methods across jurisdictions will allow for the comparison of data values; such
regional efforts are underway and are beginning to yield results. At this point, however, the limited ability to
make cross-jurisdictional data comparisons restricted the authors of the National Summary chapter to conclu-
sions that are primarily qualitative. Still, useful conclusions can be drawn with regard to variables being moni-
tored, data gaps that exist, general trends in the condition of resources, and national-level progress toward
conservation activities.

Ultimately, the goal of this report is to answer the difficult but vital question: what is the condition of U.S. coral
reef ecosystems? Coral reef ecosystems clearly are beset by a wide array of significant threats, and while
managers and scientists may be able to demonstrate improvements in some aspects of an ecosystem, de-
terioration in other aspects may yield an overall conclusion of 'no change' or decline. A valid response to the
above question is that it is too soon to tell, not because deterioration or recovery is in an early stage, but be-
cause the necessary long-term datasets that quantify such conditions have not been amassed. Few monitor-
ing programs have been in place for longer than a decade, and many have been initiated only within the past
two to five years. Some of these monitoring programs are still in their infancy and have not collected enough
data to provide conclusive results. With continued support of these critical monitoring activities, however,
trends may become more apparent over time.

Major conclusions of this report related to the threats and stressors impacting coral reef ecosystems indicate
that some appear to be intensifying while others are decreasing in intensity. Climate change was identified by
11 of the 14 jurisdictions (78%) as being a moderate (6) or high (5) threat to coral reef ecosystem resources.




The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

Climate change, whether due to natural variability or human activity, is central to several of the threats impact-
ing coral reef ecosystems. Potential impacts from climate change on coral reef ecosystems include modifica-
tion of water chemistry and sea level rise that may affect coral growth, the greater incidence and prevalence of
coral bleaching associated with increased sea surface temperatures, and the increased intensity and frequen-
cy of storm events. Coastal development was cited as a moderate (2) or high (8) threat in 10 of the jurisdic-
tions. Coastal development and population growth, whether permanent or temporary (such as in the case of
tourism), are correlated with the intensification of several threats because development frequently translates
to increases in pollution entering the marine environment; sedimentation from construction, agriculture, and
road-building activities; and physical damage from recreational users through trampling, vessel groundings,
or the use of anchors in fragile habitats. Another urgent threat, which was cited as a moderate (6) or high
(8) threat by all of the 14 jurisdictions, stems from fishing. Changes in the populations of marine organisms,
and fish in particular, can have far-reaching cascade effects throughout the ecosystem. For example, the re-
moval of herbivorous fish may precipitate changes in benthic communities by favoring algal species that can
outcompete corals following a release of predation pressure. The removal of top level predators may have a
cascading effect on the entire ecosystem by reducing overall ecosystem productivity and upsetting the bal-
ance of energy flow throughout the system with unknown consequences.

Improvements in the status of some threats have also been documented. One positive development has been
the removal of over 400 tons of marine debris, largely nets and fishing line, from the shallow reefs of the North-
western Hawaiian Islands. In addition, many jurisdictions continue to install mooring buoys to help minimize
anchor damage while facilitating access for recreational activities. Management of the trade in aquarium fish
has resulted in more protection for some U.S. coastal areas, and implementation of the provisions of the Con-
vention on International Trade in Endangered Species of Wild Flora and Fauna and the Federal Endangered
Species Act extend protection to coral species, largely prohibiting their sale or exportation. In addition, nine
grounded, rusted-out fishing vessels were removed from a reef flat in Pago Pago Harbor, American Samoa.
These and other important improvements are detailed in the jurisdictional chapters.

Other important conclusions can be drawn in relation to advancements in management and conservation sci-
ence. Major highlights include the progress made in the development of tools that scientists and coastal man-
agers use to measure the condition of the resources. Digital map products of nearshore (< 30 m) coral reef
habitats now exist for most jurisdictions and are being used to structure monitoring programs, inform manage-
ment decisions, and build capacity in current and future coastal managers. Complementary multibeam maps
of mid-depth (>20 m) environments are also being developed, and products are becoming increasingly avail-
able. Techniques to investigate genetic linkages among populations and identify and track the spread of coral
disease are becoming more sophisticated and more widely disseminated. Other research is being conducted
to determine optimal restoration techniques and calculate resource damages, which enables natural resource
trustees to seek compensation for injured coral reef ecosystems and devote those funds toward restoration
and monitoring activities. Advances in satellite observing systems and the deployment of additional buoys that
monitor oceanographic conditions continue to improve the ability to characterize coral reef ecosystems.

As implementation of the NCRAS continues, it is crucial that existing gaps, especially in the shortage of trained
personnel and infrastructure, be filled with additional resources. Without the availability of reliable, consistent
data collected at sufficient spatial and temporal resolutions, answering management questions and evaluating
management effectiveness cannot be confidently achieved.

This report represents the second in an ongoing series of reports that integrate the wealth of quantitative and
qualitative information on the condition of U.S. coral reef ecosystems that has emerged since the inception
of the USCRTF. Future reporting efforts will continue to document progress toward the goals outlined by the
USCRTF and in the NCRAS and contribute to a broader understanding of U.S. coral reef ecosystems.




The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

INTRODUCTION

Much of the vast ocean realm that covers the planet is composed of very deep water. Thousands of meters be-
low the surface, the bottom of the ocean lies in complete darkness and is sparsely populated. However, where
the seafloor slopes up toward the continental shelf and the flanks of oceanic islands, marine life becomes
more concentrated due to the greater availability of sunlight and nutrients from upwelled water and terrestrial
inputs. Tropical nearshore areas are comprised of a variety of habitats which are frequently classified accord-
ing to their dominant substrate, geological, and biological features. While some sandy or rocky substrates are
sparsely colonized or devoid of life, others provide habitat for seagrasses and other plant and algal communi-
ties. In some hardbottom areas where conditions are right, the seafloor is colonized by a variety of tiny colonial
invertebrates known generally as corals. Over millions of years, these tiny organisms have created enormous
underwater structures that provide a foundation for an elaborate community of creatures that together consti-
tute one of the most amazing and diverse ecosystems on the planet. An oasis in a vast ocean, coral reefs at-
tract and concentrate a breathtaking
assemblage of colorful and fanciful
organisms that challenge the limits of
the imagination (Figure 2.1). Scien-
tists estimate that this highly complex
interdependent system supports over
one million species, with potentially
millions more yet to be described. In
addition to their importance for bio-
diversity, coral reef ecosystems are
important for human communities as
well, by performing essential ecosys-
tem services; supporting major fish-
ery resources; providing educational,
social, recreational, cultural, and
medicinal opportunities; and gener-
ating economic benefits for millions
of people, especially through coastal
tourism. Figure 2.1. Coral reefs provide the structure that attracts and concentrates a colorful
assortment of interesting organisms. Photo: NOAA NOS.
The vibrant underwater world of coral reefs comprises less than 1% of the surface of the planet, primarily due
to the narrow physiological tolerances of hermatypic, or reef-building, corals. Nearly all coral reefs are found
throughout tropical and subtropical oceans between 300S and 300N latitude, primarily in waters less than 30
m deep (Huston, 1985; Grigg and Epp, 1989). Their distribution is influenced by nutrient availability, salinity,
light, substrate, sediment type, temperature, and exposure to wave action (Lalli and Parsons, 1995; Hoegh-
Guldberg, 1999; Szmant, 2002; Leichter et al., 2003; Wolanski et al., 2003). Seawater temperatures in coral
ecosystems generally range between 180-290C (Glynn, 1996; Barnes and Hughes, 1999), although some
corals seem to have adapted to tolerate slightly higher temperatures for short periods of time. Many organ-
isms living in coral reef ecosystems are photosynthetic and are restricted to shallow depths with sufficient light
penetration (Veron, 1986; Barnes, 1987).

Shallow-water coral reef ecosystems under United States jurisdiction occur in the shared waters of the Carib-
bean Basin, Gulf of Mexico, and Atlantic Ocean near the east coast of Florida, and across the Pacific Ocean
on both sides of the equator. The Freely Associated States of the Republic of Palau, the Federated States of
Micronesia (FSM) and the Republic of the Marshall Islands (RMI) are located in the tropical western Pacific
Ocean. Pacific reef systems tend to proliferate on oceanic islands in a number of habitats ranging from off-
shore banks to shallow atoll lagoons. Many Pacific islands formed as a result of volcanic activity beneath the
earth's surface and/or uplift of limestone or sedimentary rock. Movement of the enormous Pacific plate across
tectonic 'hot spots' resulted in the creation of several long island chains which developed complex reef sys-
tems over time. In general, as soon as lava cools and forms stable, hard substrate, corals begin to colonize the
submarine margins of islands as narrow fringing reefs. As the islands age, coral reefs continue to gradually ac-
crete while the central land area slowly erodes and subsides, until, after millions of years, the island itself may




The State of Coral Reef Ecosystems of the United States and j


disappear completely, leaving a necklace of low sand
islets and extensive reefs surrounding a broad lagoon
(Figure 2.2). Cores taken from coral reefs near Bikini
and Enewetak Atolls, Marshall Islands revealed coral
deposits nearly 1.4 km thick, which are believed to be
50-59 million years old (Spalding et al., 2001). Many
stages of island development, from creation by active
volcanoes to submergence beneath the surface, are .
evident in the archipelagos of the Pacific.

In contrast to most Pacific reefs, many reef formations
in the Caribbean Basin have developed in shallow- --
water environments near relatively stable continental
land masses. Coral reef ecosystems near continents
tend to be older than reef systems on many oceanic .
islands, and are often subject to greater terrestrial in-
puts, such as freshwater, sediments and nutrients.
To a large extent, reefs located on broad continental
shelves benefit from their close association with es- -.........
tuaries and mangrove forests which filter out harmful .
nutrients and sediments as well as nurture large ju- ..... ". ...""

In turn, shallow or emergent reefs protect fragile
coastlines by absorbing wave action during storms
and high swells. ::::::..-;......


Coral Reef Ecosystem Components
A coral ecosystem can be considered a mosaic of
habitats defined by substrate, cover, and structural
zones (Figure 2.3). Benthic habitats found in a coral Figure 2.2. A diagram depicting the evolution of a coral ecosys-
ecosystem include unconsolidated sediments (e.g., tem on a volcanic island. As the island ages, wind and rain erode
S the land while reefs along the island perimeter accrete.
sand and mud); mangroves and other emergent
vegetation; submerged vegetation (e.g., seagrass and macroalgae); hermatypic coral reefs and associated
colonized hardbottom habitats (e.g., spur and groove, individual and aggregated patch reefs, and gorgonian-
colonized pavement and bedrock); and uncolonized hardbottom (e.g., reef rubble and uncolonized bedrock).
Typical structural zones include the reef crest, forereef, reef flat, backreef and lagoon (FMRI, 1998; Kendall et
al., 2001; Coyne et al., 2003; NOAA, 2003). While hermatypic coral reefs are important marine habitats, other
habitats, such as bare sand or seagrass, are also important to the overall ecology and function of the eco-
system. Mangrove forests, hardbottom coral habitats, submerged vegetation habitats, and softbottom sand
and mud habitats serve as important spawning and growth areas (Ogden and Ehrlich, 1977; Lindeman, 1986;
Parrish, 1989; Christensen et al., 2003; Kendall et al., 2003; Mumby et al., 2004).


Humans in Coral Reef Ecosystems
For thousands of years, humans have lived in coastal areas adjacent to coral reef ecosystems. Coastal and
island communities regularly harvested marine resources for food, and in some areas, marine resources pro-
vided the primary, if not only, source of protein. In addition to providing basic sustenance to island and coastal
communities, reefs have inspired art and legends and provided humans with natural products, jewelry, phar-
maceuticals, building materials, transportation pathways, and recreational opportunities. Many cultures cite
strong cultural ties to reef ecosystems and resources, and have gone to great lengths to protect the resources
from overexploitation, as evidenced by the elaborate systems of reef tenure and conservation practices de-
vised by some Pacific island communities to regulate the use of marine resources.





The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005


Figure 2.3. Examples of some of the types of benthic habitats found in the shallow-water coral ecosystems of the United States. Left
to right and top to bottom, these are:
1. Mangrove, Salt River, St. Croix, USVI
2. Bare Sand, Midway Atoll, NWHI
3. Macroalgae and sand, Puerto Rico
4. Hardbottom with macroalgae, Kure Atoll, NWHI
5. Thalassia seagrass, St. Croix, USVI
6. Linear reef with live coral, Midway Atoll, NWHI
7. Hardbottom with crustose coraline algae, Lisianski, NWHI
8. Spur and groove, Mona Island, Puerto Rico
9. Uncolonized pavement with sand channels, Mona Island, Puerto Rico
Photos: CCMA-BT.




The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

Reef ecosystems also provide intangible benefits that have inspired a long romance with the coast and draw
millions of people each year to visit or live near coastal areas of the tropics. On the U.S. mainland alone, 10.5
million people live in areas adjacent to coral reef ecosystems, and island populations continue to increase
through population growth and immigration. In fact, population growth and associated development has been
identified as a key threat in American Samoa and several other small island states. Tourism and recreational
activities also temporarily increase the number of people inhabiting coastal areas. A recent report by the U.S.
Commission on Ocean Policy (2004) provides evidence of the increasing importance of tourism and recre-
ation to the economies of coastal communities. Staggering numbers of people visit coastal areas every year
to fish, dive, surf and recreate. As a result, the economic benefits from coastal and ocean resources have
experienced a fundamental shift from a products-based to services-based system, with tourism and recreation
generating more income than mineral and living resource extraction, transportation, and shipbuilding (U.S.
Commission on Ocean Policy, 2004).

Coral reef ecosystems found in the U.S. support millions of dollars worth of goods and services (e.g., com-
mercial and recreational fisheries, tourism, etc.). Recent estimates indicate that activities associated with
Hawaii's coastal ecosystems produce about $364 million for the state's economy every year (Davidson et al.,
2003). Activities associated with Florida's coastal ecosystems contribute an estimated $2.7 billion annually to
its economy (Johns et al., 2001). The intangible values of U.S. coral reef ecosystems-such as aesthetic, eco-
logical, and cultural values-are difficult to quantify and are excluded from these economic value estimates.

All of this attention and interest in coastal areas in general, and coral reef ecosystems in particular, are not
without consequence. Against a background of natural disturbances, increased disturbance from human ac-
tivities reduces the resilience of coral reef ecosystems and can contribute to alarming declines in their overall
health. Key anthropogenic stressors include climate change and bleaching; disease; urban and tourism-re-
lated coastal development; sedimentation; toxic chemical pollution; overfishing; physical damage from ships,
boats, and anchors; invasions of exotic species; and marine debris (Davidson, 2002; Wilkinson, 2002; Gard-
ner et al., 2003; NCRAS, 2002).


Setting
The U.S. is responsible for managing and conserving extensive shallow-water coral reef ecosystems within
its maritime boundaries in cooperation with local governments of various types. U.S. States with coral reef
ecosystems include Florida and Hawaii. U.S. Territories include the U.S. Virgin Islands (USVI), American
Samoa, and Guam. The Commonwealths of Puerto Rico and the Northern Marianas Islands also have coral
reef ecosystems. Navassa Island is an unincorporated U.S. Territory near Haiti. The Flower Garden Banks
lie in Federal waters off the coast of Texas, and some of the banks are managed by the National Oceanic
and Atmospheric Administration's (NOAA) National Marine Sanctuary Program in cooperation with the U.S.
Department of the Interior's Minerals Management Service. The Northwestern Hawaiian Islands (NWHI) are
jointly managed by the U.S. Fish and Wildlife Service (USFWS), the State of Hawaii, and the NWHI Coral Reef
Ecosystem Reserve, but the islands have been proposed as the nation's 13th national marine sanctuary, and
sanctuary designation seems likely in the near future. The Pacific Remote Island Areas (PRIAs) of the Line
and Phoenix Islands and Johnston Atoll are primarily managed by the USFWS as national wildlife refuges,
and jurisdiction over Wake and Johnston Atolls is currently in the process of being transferred from the U.S.
Department of Defense to the USFWS. The Freely Associated States (FAS) of the RMI, FSM, and the Repub-
lic of Palau are sovereign nations that maintain a close economic association with the U.S. and claim similar
maritime boundaries. Coral ecosystems of the U.S. and FAS cover a vast area and are distributed across
large portions of the earth's surface (Figures 2.4 and 2.5).

Using depth curves depicted on nautical charts as a surrogate for the potential distribution and extent of
shallow-water coral ecosystems, Rohmann et al. (in press) estimated that 36,816 km2 of coral ecosystems
may potentially be found in U.S. waters less than 10 fathoms (approximately 18 m) deep, and an estimated
143,058 km2 in waters less than 100 fathoms (approximately 183 m) deep (Table 2.1).
























----------------



















kni
0 250 500



low- Exclusive Economic
Zone(EEZ)


NORTHWESTERN
HAWAIIAN ISLANDS


COMMONWEALTH KU14 NAWAUX
OF THE NORTHERN
MARIANA ISLANDS




GUAM



REPUBLIC
REPUBLIC OF THE
OF PALAU FEDERATED STATES MARSHALL
OF MICRONESIA ISLANDS




AM9WCAN
M04.



(54@
-j4 1
.WK





The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

Table 2.1. The potential area of coral ecosystems within the United States territorial sea and exclusive economic zone.a The area in-
side the 10-fathom (18 m) or 100-fathom (183 m) depth curves was derived from NOAA nautical charts.b Estimates for the RMI, FSM,
and Republic of Palau were derived from Landsat satellite imagery. Source: Rohmann et al., in press.


USVI- 344 2,126


Puerto Rico4

Naassa

Southern Florida'

Flower Gardens NMSS'-

Main Hawaiian Islands

Northwestern Haaiian Islands-

Amencan Samioa'

Pacific Remote Island Areas"


Marshall Islands'


Federated States of Micronesia''

Northern Mariana Islands'"


Guam

PalauIa'


2.302 5.501

3 14

30.801 113,092

0 164

1.231 6.666


1,595


13.771


1 The U.S. territorial sea (and contiguous zone) extends 12 nautical miles from the baseline of each territory or coastal State. The
U.S. exclusive economic zone extends 200 nautical miles from a line coterminous with the seaward boundary (baseline) of each
territory or coastal State.
2 Area estimates from Rohmann et al., in press.
3 The U.S. Virgin Islands includes the islands of St Thomas, St John, and St Croix.
4 Puerto Rico includes the islands of Puerto Rico, Desecheo, Culebra, Vieques, and Mona.
5 Southern Florida extends along the Atlantic Ocean coast of Florida to Jupiter Inlet, Florida and along the Gulf of Mexico coast of
Florida to Tarpon Springs, Florida.
6 The NOAA nautical chart depicts only the 100 fathom depth curve for this location.
7 The Main Hawaiian Islands includes the islands of Hawaii, Maui, Molokai, Lanai, Kahoolawe, Oahu, Kauai, and Niihau.
8 The Northwestern Hawaiian Islands includes the islands and atolls of Nihoa, Necker, French Frigate Shoals, Gardner Pinnacles,
Maro Reef, Laysan, Lisianski, Pearl and Hermes, Midway, and Kure. Numerous shallow-water seamounts, such as St. Rogatein
Bank or Raita Bank, also are located in the NWHI.
9 American Samoa includes the islands of Tutuila, Ofu, Olosega, Tau, Swains, and Rose Atoll.
10 The CNMI includes the islands of Rota, Aguijan, Tinian, Saipan, Farallon de Medinilla, Anatahan, Sarigan, Guguan, Alamagan,
Pagan, Agrihan, Asuncion, Maug, and Farallon de Pajaros.
11 The U.S. Flag Islands include Howland, Baker, and Jarvis Islands, Palmyra, Johnston, and Wake Atolls, and Kingman Reef.
12 Unpublished estimates of potential coral ecosystem area visible in Landsat satellite imagery. Area estimates generally include
seafloor features visible in water 18-27 m (10-15 fathoms) deep. NOAA does not produce nautical charts of these locations.

At this time, nautical charts depicting either depth or extent of shallow-water coral ecosystems for the FAS are
unavailable. However, an analysis of seafloor features visible in Landsat satellite imagery suggests that coral
ecosystems in the FAS may comprise about 30,501 km2 (Table 2.1).

The spatial extent of shallow water coral ecosystems is just one of several variables that differentiate coral reef
ecosystems among U.S. jurisdictions. Perhaps an even more important metric is habitat quality. This metric
can be characterized in a number of ways, but high habitat quality conveys the presence of a rugose and var-
ied assemblage of healthy benthic organisms that provide structure for a robust and diverse assemblage of
organisms within an environment characterized by excellent water quality with low turbidity, limited nutrients,
and few contaminants. Such healthy reef ecosystems tend to support more biomass and a greater number of


13.456

14.517


2.529




f the United States and Pacific Freely Associated States: 2005


species than degraded areas.

Biodiversity, or the number and abundance of species that exist within a region, is another important variable.
Global marine biodiversity is believed to be highest in the western Pacific Ocean, near eastern Indonesia, and
the total number of species tends to decline with distance from this biological hot spot. As a result, among U.S.
and FAS jurisdictions, the Republic of Palau and other western Pacific locations (i.e., Guam, CNMI, FSM and
the Marshall Islands) naturally contain a higher number of species than do locations in the eastern Pacific,
Caribbean, Atlantic or Gulf of Mexico.

The degree of endemism, or the number of species that are found only within a particular location or region,
is another important factor that distinguishes the jurisdictions. Scientists studying remote areas, such as the
NWHI, which have a relatively low overall number of species, have recorded a large number of endemic spe-
cies. Endemic species contribute greatly to the overall diversity of life on the planet and thus constitute an
important conservation priority.

Among other important distinguishing characteristics among the jurisdictions is the actual composition of
the coral and fish communities. Highly disturbed ecosystems often are dominated by species of coral and
macroalgae that are opportunistic and tolerant of negative natural and anthropogenic impacts. Heavily fished
ecosystems often are dominated by small, undesirable food fish not targeted by fishers. For corals, the
prevalence of long-lived versus opportunistic species may provide some indication of the level of disturbance
experienced in a region and thereby the health of the system as a whole.

The prevalence of threats and stressors to coral reef ecosystems also varies among and within jurisdictions.
The NCRAS identified thirteen major threats and stressors to coral reef ecosystems that are introduced in the
following chapter (NCRAS, 2002). Chapter 4 begins a series of fourteen jurisdiction chapters, in which each
jurisdictional writing team provides a condition report according to a standardized structure. Each chapter
begins with a few paragraphs of contextual information and a discussion of how each of the thirteen primary
threats currently affects their jurisdiction. That information is followed by a summary of current monitoring ac-
tivities, and project results which are grouped into the three categories of water quality, benthic habitats, and
associated biological communities. They then discuss current conservation management activities pertinent to
their jurisdiction before providing overall conclusions and recommendations for further action. The final chap-
ter serves as a national-level summary of the preceding information, in addition to providing information about
selected national-level developments that are pertinent to all the jurisdictions.




The State of Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2005

REFERENCES

Barnes, R. 1987. Invertebrate zoology: Fifth Edition. Harcourt Brace Jovanovich, Inc., Orlando, FL. pp. 149-163.

Barnes R. and R. Hughes. 1999. An introduction to marine ecology: Third Edition. Blackwell Science, Inc., Maiden, MA.
pp. 117-141.

Coyne, M.S., T.A. Battista, M. Anderson, J. Waddell, W. Smith, P. Jokiel, M.S. Kendall, and M.E. Monaco. 2003. Benthic
habitats of the main Hawaiian Islands. NOAA Technical Memorandum NOS NCCOS CCMA (On-line). Silver Spring,
Maryland. Available from the internet URL: http://biogeo.nos.noaa.gov/projects/mapping/pacific

Christensen, J.D., C.F.G. Jeffrey, C. Caldow, M.E. Monaco, M.S. Kendall, and R.S. Appeldoorn. 2003. Cross-shelf habitat
utilization patterns of reef fishes in southwestern Puerto Rico. Gulf and Carib Research 14(2): 9-27

Davidson, K., M. Hamnett, and C. Minato. 2003. The first four years: Hawaii coral reef initiative research program (1998-
2002). Social Science Research Institute, University of Hawaii at Manoa. 72 pp

Davidson, M.G. 2002. Protecting coral reefs: the principal national and international legal instruments. Harvard Environ-
mental Law Review 26: 499-546

FMRI (Florida Fish and Wildlife Conservation Commission, Florida Marine Research Institute and National Oceanic and
Atmospheric Administration). 1998. Benthic habitats of the Florida Keys. FMRI Technical Report TR-4. 53 pp.

Gardner, TA., I.M. Cote, J.A. Gill, A.Grant, and A.R. Watkinson. 2003. Long-term region-wide declines in Caribbean cor-
als. Science 301: 958-960

Glynn, P.W. 1996. Coral reef bleaching: facts, hypotheses and implications. Global Change Bio 2: 495-509

Grigg, R.W. and D. Epp. 1989. Critical depth for the survival of coral islands: effects on the Hawaiian Archipelago. Sci-
ence 243: 638-641

Hoegh-Guldberg, O. 1999. Climate change, coral bleaching and the future of the world's coral reefs. Marine and Fresh-
water Research 50: 839-866

Huston, M.A. 1985. Patterns in species diversity on coral reefs. Annual Review of Ecological Systems 6: 149-177.

Johns, G.M., V.R. Leeworthy, F.W. Bell, and M.A. Bonn. 2001. Socioeconomic study of reefs in southeast Florida. Report
by Hazen and Sawyer under contract to Broward County, Florida. 255 pp.

Kendall, M.S., J.D. Christensen, and Z. Hillis-Starr. 2003. Multi-scale data used to analyze the spatial distribution of
French grunts, Haemulon flavolineatum, relative to hard and soft bottom in a benthic landscape. Environmental Biology
of Fishes 66: 19-26

Kendall, M.S., M.E. Monaco, K.R. Buja, J.D. Christensen, C.R Kruer, M. Finkbeiner, and R.A. Warner. 2001. Methods
used to map the benthic habitats of Puerto Rico and the U.S. Virgin Islands. National Oceanic and Atmospheric Admin-
istration Technical Memorandum NOS NCCOS CCMA 152. Silver Spring, Maryland.

Lalli, C.M. and TR.Parsons. 1995. Biological Oceanography: An Introduction. Oxford, UK: Butterworth-Heinemann Ltd.
p 220-233

Leichter, J.J., H.L. Stewart, and S.L. Miller. 2003. Episodic nutrient transport to Florida coral reefs. Limnologic Oceanog-
raphy 48: 1394-1407.

Lindeman, K.C. 1986. Development of larvae of the French grunt, Haemulon flavolineatum, and comparative develop-
ment of twelve western Atlantic species of Haemulon. Bulletin of Marine Science 39: 673-716.

Mumby, P.J., A.J. Edwards, J.E. Arlas-Gonzalez, K.C. Lindeman, P.G. Blackwell, A. Gall, M.I. Gorczynska, A.R. Harborne,
C.L. Pescod, H. Renken, C.C.C. Wabnitz, G. Llewellyn. 2004. Mangroves enhance the biomass of coral reef fish com-
munities in the Caribbean. Nature 427: 533-536.

NCRAS (National Coral Reef Action Strategy). 2002. A National Coral Reef Action Strategy: Report to Congress on
implementation of the Coral Reef Conservation Act of 2002 and the National Action Plan to Conserve Coral Reefs in
2002-2003. NOAA. Silver Spring, Maryland. 120 pp. + appendix.




The State of Coral Reef Ecosystems of the Un


DAA (National Oceanic and Atmospheric Administration). 2003. Atlas of the Shallow-water Benthic Habitats of the
)rthwestern Hawaiian Islands (Draft). 160 pp. Available from the internet URL: http://ccma.nos.noaa.gov/rsd/products.
il#nwhi

)den, J.C., and P.R. Ehrlich. 1977. The behavior of heterotypic resting schools of juvenile grunts (Pomadasyidae).
marine Biology 42: 273-280.

irrish, JD (1989) Fish communities of interacting shallow-water habitats in tropical oceanic regions. Marine Ecology
ogress Series 58: 143-160.

)hmann, S.O., J.J. Hayes, R.C. Newhall, M.E. Monaco, and R.W. Grigg. In press. The Area of Potential Shallow-water
opical and Subtropical Coral Ecosystems in the United States. Coral Reefs.

)alding, M.D., C. Ravilious, and E.P. Green. 2001. World atlas of coral reefs. Prepared at the UNEP World Conservation
)nitoring Centre. University of California Press, Berkeley, California. 424 pp.

:mant, A.M. 2002. Nutrient enrichment on coral reefs: is it a major cause of coral reef decline? Estuaries 25: 743-766

S. Commission on Ocean Policy. 2004. An Ocean Blueprint for the 21st Century: Final Report. Washington, DC.

*ron, J.E.N. 1986. Corals of Australia and the Inso-Pacific. Angus and Robertson, London.

ilkinson, C. (Ed.) 2002. Status of coral reefs of the world: 2002. Australian Institute of Marine Science.

olanski, E., R. Richmond, L. McCook, and H. Sweatman. 2003. Mud, marine snow and coral reefs. Amercan Scientist
:44-51.







Threats and Stressors to U.S. Coral Reef Ecosystems

Andy Bruckner1, Ken Buja2, Liz Fairey1, Kelly Gleason2, Michelle Harmon3, Scott Heron4, Tom Hourigan1, Chris Jeffrey2,
Julie Kellner2, Ruth Kelty3, Bob Leeworthy5, Gang Liu4, Simon Pittman2, Aurelie Shapiro2, Al Strong4, Jenny Waddell2,
Peter Wiley5.

Human activity is commonly identified as a major contributor to the observed global deterioration of coral reef
ecosystem health, with loss of live coral cover, declining species diversity, and reduced abundance reported
in many areas (NOAA, 2002a; Wilkinson, 2002; Turgeon et al., 2002). Degradation in the structure and func-
tioning of coral reef ecosystems results in a concomitant loss in the intrinsic value of the ecological system,
as well as a significant loss in the provision of goods and services for society. Approximately 8% of the global
population live within 100 km of a coral reef (Bryant et al., 1998) and many local communities and national
economies are directly dependent on coral reef ecosystems for tourism revenue, food, and coastal protection
(Spurgeon, 1992). As such, human pressures can be intense, and developing strategies to mitigate stressors
is a complex task.

Shallow-water coral reef ecosystems experience a wide range of physical, biological, and chemical threats
and stressors, which stem from both anthropogenic and natural causes. Threats are defined as environmental
trends with potentially negative impacts. Stressors are defined as factors or processes that harm ecosystem
components, causing lethal or sublethal negative effects. Categories of stressors include chemical (e.g., pol-
lution), physical (e.g., extreme events), and biological (e.g., invasive species) stressors, and the relationship
between key stressors and the threats discussed in this document are listed in Table 3.1. The relative impor-
tance of each threat varies substantially among jurisdictions and individual reefs.

Table 3.1. This table is a crosswalk between the threats identified in "A National Coral Reef Action Strategy" (NOAA, 2002a) and the
stressors identified by the National Science and Technology Council's Committee on Environmental and Natural Resources. Source:
CENR, 2001.


STESOR PLLTIN NVSIE XTEM RSORC CIMT




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


widespread decline. The challenge now is to understand the complex interactions among stressors by refin-
ing existing techniques and developing new multidisciplinary approaches aimed at detailing mechanisms and
predicting effects at multiple spatial and temporal scales.

Determining how humans utilize coral reef ecosystems and estimating the social and economic costs and ben-
efits of those uses are key steps for resource managers. Techniques such as causal chain analysis (e.g., in
Belausteguigoitia, 2004) may provide a useful approach for modeling and communicating the many significant
cause-effect linkages between human systems and coral reef ecosystems.


Climate Change and Coral Bleaching
Climate change refers to any change in climate over time, whether due to natural variability or human activity
(IPCC, 2001). Over the 20th century, mean near-surface air temperature over land and mean sea surface
temperature (SST) increased 0.6 0.2C, with the 1990s being the warmest decade and 1998 being the
warmest year since 1861 when instrumental records began (IPCC, 2001; Figure 3.1).

Most of the observed warming over the last 50 years may have resulted from an increase in concentrations of
greenhouse gases such as carbon dioxide (CO2) and methane (CH4) in the atmosphere (IPCC, 2001; NRC,
2001). The atmospheric concentration of CO2 has increased by 31% since the beginning of the industrial rev-
olution, and represents a level that has not been exceeded in at least the last 420,000 years (Petit et al., 1999),
and probably not exceeded in over 24 million years (Pearson and Palmer, 2000). The rate of increase of CO2
concentration has been about 0.4% per year over the last two decades (IPCC, 2001). Such increases have
been shown to decrease the calcium carbonate (CaCO3) saturation state of seawater and the calcification
rates of corals (Kleypas et al., 1999; Feely et al., 2004). In combination with potentially more frequent bleach-
ing episodes, reduced calcification
could reduce the energy that a coral
would otherwise apply to reproduction Global Surface Mean Temp Anomalies
and thereby impede a reef's ability to
Stre im r ii National Climatic Data Center/NESDIS/NOAA
keep pace with sea level rise (IPCC, 0 . .......
2001) or recover from other potential 0.6 Land and Ocean 1.0
impacts of climate change. 0.
0.3" .IJ^H


Elevated water temperatures cause
corals to bleach, a process that is
characterized by the loss of zooxan-
thellae (a symbiotic alga) from coral
tissues. Increased ultraviolet irradi-
ance, typically from unusually calm,
clear waters, may aggravate the
impact of increased temperatures
(Lesser and Lewis, 1996). Although
corals may recover from brief epi-
sodes of bleaching, if ocean temper-
atures warm too much or remain high
for an extended period, bleached cor-
als often will die. Several correlative
field studies show a close association
between warmer than normal condi-
tions (at least 1 C higher than the an-
nual maximum) and the incidence of
bleaching (Hoegh-Guldberg, 1999).
In 1997-1998, an estimated 16% of
the world's coral reefs were seriously
damaged in a global coral bleach-
ing event associated with high SST


0.0 1.0
-0.3

0.6 1.0
0.3
E 0.0 -0.0
S-0.3 (D
-0.6 -1.0

Sr Land
0.6- 1.0 (

0.0 0.0

-0 6- -1.0

-1 7 -2.0
_0 1900 1920 1940 1960 1980 2000
Year
Figure 3.1. Mean global temperature anomalies over the period 1880-2001. Zero
line represents the long term mean temperature throughout the period, while red and
blue bars indicate annual departures from that mean. Source: NOAAs National Cli-
mactic Data Center.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


which was apparently enhanced by an extreme El Niio event (Wilkinson, 1998). A U.S. Department of State
report to the U.S. Coral Reef Task Force (USCRTF; Pomerance, 1999) concluded that the severity and extent
of the 1998 event cannot be explained by El Niio alone, and that the "...geographic extent, increasing fre-
quency, and regional severity of mass bleaching events are likely a consequence of a steadily rising baseline
of marine temperatures..."

Several bleaching events in Florida, the U.S. Caribbean, and the U.S. Pacific have been associated with el-
evated SST events during the 1980s and 1990s, and especially in 1997-1998. The occurrence of bleaching is
highly variable in both time and space, but generally affects shallow-water reefs with reduced water circulation.
In U.S. waters, substantial bleaching has been observed on shallow reefs off the coasts of Florida, the Com-
monwealth of the Northern Mariana Islands (CNMI), Palmyra Atoll (PRIAs), and portions of the Northwestern
Hawaiian Islands (NWHI), and recent data suggest that elevated SST is still a significant threat to coral reefs
in the U.S. Caribbean (Nemeth and Slakek-Nowlis, 2001). Palau suffered the worst coral bleaching mortality
of any U.S. associated region during the 1997-1998 global bleaching event (Wilkinson, 2000). During a 2002
summertime warm water event in the
higher latitudes of the mid-Pacific,
Midway Atoll (NWHI) experienced
unprecedented bleaching, includ-
ing considerable mortality (Liu et al.,
2004). Mass bleaching episodes are
predicted to reoccur in the future with
M. increasing frequency (IPCC, 2001).

Coral reef ecosystem managers and
stakeholders consistently use one
particular satellite-derived index-the
Degree Heating Week (DHW)-to
gauge accumulated thermal stress on
reef ecosystems. The DHW, which
was developed by scientists in the
Figure 3.2. 2002 Maximum annual DHW values indicate locations that experienced National Oceanic and Atmospheric
significant thermal stress, which has been shown to be highly correlated with coral Administration's (NOAA) Coral Reef
bleaching. Values above 4 represent areas that are likely to experience bleaching, Watch (CRV) Program, represents
while values above 8 represent areas that are likely to experience significant bleach- re
ing with widespread mortality. Source: NOAAs Coral Reef Watch Program. the accumulated temperature stress
for each 50 x 50 km2 pixel during the
preceding 12-week period as compared to the baseline value calculated for that pixel. The unique baseline
value, roughly equal to the expected annual maximum temperature, was empirically determined for each of
the 250 km2 pixels shown in Figure 3.2. To calculate the DHW, temperature deviations (in degrees Celsius)
above this baseline are multiplied by the duration of the elevated temperature event (in weeks). For example,
if there is a sustained SST of 1 VC above the threshold for one week, during a 12-week period, the DHW value
will be one; if SST is 20C above the threshold for three weeks, the DHW value will be six. Figure 3.2 illustrates
the distribution of the maximum DHW values for each pixel for 2002.

In-situ observations show that widespread bleaching is most likely to occur at locations where DHVW4; signifi-
cant bleaching with widespread mortality is expected where the DHW >8. Table 3.2 shows the maximum an-
nual DHW value in the 14 U.S. jurisdictions with coral reefs for 2001-2003. The DHW values are color-coded
to reflect the intensity of accumulated thermal stress [Blue, DHW=0; Green, DHW <4; Orange, 45 DHW <8;
Red, DHW >8]. If a thermal stress event spans two calendar years (e.g., November-January), then the maxi-
mum DHW for each of those years may occur during that single event. This is most likely to occur at reefs
located near the equator. Such occurrences are shown in Table 3.2 as DHW values enclosed in a grey box.
The CRW Program utilizes satellite and in situ tools for near real-time, hindcast, and long-term monitoring,
modeling, and reporting of environmental conditions that affect domestic and foreign coral reef ecosystems.
A full list of the CRW Program's operational products can be found on-line at http://coralreefwatch.noaa.gov
(Accessed 2/16/05).




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


Table 3.2. Maximum annual DHWs for each of the 14 jurisdictions for 2001-2003. The DHW values are color-coded to reflect the
intensity of accumulated thermal stress [Blue, DHW=0; Green, DHW<4; Orange, 4DHW<8; Red, DHW>8]. If a thermal stress event
spans two calendar years (e.g., November-January), then the maximum DHW for each of those years may occur during that single
event. Such occurrences are shown by enclosing the DHWvalues in a grey box.


JURISDICTION I LOCATION


2001


2002


2003 JURISDICTION LOCATION


9 Y ~ 9 Y ~ h E


USVI


Puerto Rico0 0 i

Navassa

Florida
Flower Garden
Banks


Hawaii


Northwestern
Hawaiian
Islands


American
Samoa


USPRIAs


Hawaii


0


0


Oahu


Kauai

Nihoa
French
Frigate
Shoals


Maro Reef

Lisianski


Midway


Kure


Tutuila


Rose Atoll


Johnson


Palmyra IW


0


0


USPRIAs
(cont.)


Marshall
Islands


Federated
States of
Micronesia


CNMI


Guam


Kingman


Baker

Wake


Jarvis

Howland


Bikini


Majuro


Yap

Chuuk


Pohnpei

Kosrae

Asuncion


2001 2002 2003


0

0


0


Agrihan 40 0 0

Pagan 0 0j


Saipan


Palau 0 0 L


Kwajalein 0 0 0




tes and Freely Associated States: 2005


Diseases in Coral Reef Ecosystems
In the past two decades, there has been a worldwide increase in the reporting of diseases affecting marir
organisms, with the Caribbean Basin emerging as a hot spot (Harvell et al., 1999). The first documented cor
reef epizootic was the mass mortality of the keystone herbivore, Diadema antillarum, which was caused by
unknown waterborne pathogen (Figure 3.3). This disease spread throughout the Caribbean between 19(
and 1983, moving with Caribbean oceanic currents and causing the loss of up to 90-95% of the Diadema po
ulation (Lessios et al., 1984). Mass
mortalities of Diadema have contrib-
uted to phase-shifts from coral- to
algal-dominated reefs in many lo-
cations, and the recovery of urchin
populations has been slow. Another
Caribbean-wide epizootic observed
during the 1980s was attributed to a
fungal infection in Thalassia testudi-
num seagrasses. In Florida Bay, an
estimated 4,000 ha of seagrasses
were lost and severe declines were
observed across an additional 23,000
ha (Roblee et al., 1991). During one
of the best documented of coral dis-
ease outbreaks which occurred in
the 1980s, two of the dominant reef-
building coral species on shallow
western Atlantic reefs (Acropora pal- Figure 3.3. Coral disease and mortality from numerous pathogens have been i
ported with increased frequency since the 1970s. Disease in other ecosystem orge
mata and A. cervicornis) were virtual- isms can also result in cascading effects that can disrupt the entire system. Scienti,
ly eradicated by white-band disease believe that -90% of the Caribbean population of Diadema antillarum, an imports
(Aronson and Precht, 2001). The fre- herbivore, was killed by disease in the late 1980s, and the subsequent reduction
grazing pressure allowed for algal overgrowth on many reefs. Populations are beg
quency and severity of outbreaks of ning to rebound as shown in this photo taken in St. Croix in October 2004. Photo:
common as well as newly emerging Clark.
diseases may increase with changing
environmental conditions such as a rise in SST and anthropogenic impacts that: 1) increase the prevalen(
and virulence of pathogens; 2) facilitate invasions of new pathogens from terrestrial or aerial sources; and
reduce host resistance and resilience, thereby facilitating pathogen transmission and infection (Sutherland
al., 2004).

Since the early 1990s, scientists have documented a rapid emergence of diseases among corals, with i
creases in the number of diseases reported, coral species affected, geographic extent, prevalence and i




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

and forereef habitats, is the leading cause of the decline in coral cover in the Caribbean reported during the
1980s and 1990s (Richardson and Aronson, 2002). Coring studies from Belize and other locations revealed
that mass mortalities at this scale had not occurred in at least the previous 3,000-4,000 years (Aronson et al.,
2004). More recently, Montastraea annularis complex populations are experiencing significant declines as
a result of multiple diseases including black-band disease, yellow-band disease, and white plague (Santavy
et al., 1999; Kuta and Richardson, 2002; Gill-Agudelo and Garzon-Ferriera, 2001; Richardson and Aronson,
2002; Bruckner and Bruckner, 2003, 2004).

Understanding the relationships between coral health and environmental parameters is of key importance in
the study of coral disease (Harvell et al., 1999; Green and Bruckner, 2000; Kuta and Richardson, 2002). En-
vironmental stressors, including those associated with degraded water quality and climate change, are often
cited as potential factors causing coral mortality, yet rarely have studies adequately identified causal linkages
to specific environmental stressors (Woodley et al., 2003). In addition, human activity may enhance the
global transport of pathogens, such as Aspergillus sydowii (a fungus of terrestrial origin) that causes infection
and mortality in sea fans and other gorgonians, and is postulated to have entered the marine environment
via terrestrial runoff or clouds of dust from West Africa (Harvell et al., 1999; Richardson and Aronson, 2002).
White pox, a disease only known to affect Acropora palmata in Florida, is caused by a common fecal entero-
bacterium Serratia marcescens, which may enter the marine environment via sewage discharge (Patterson
et al., 2002). Other diseases are thought to be caused by known microorganisms that have changed hosts
or exhibited increased virulence in response to environmental stresses and reduced resistance of the host
coral (Santavy and Peters, 1997; Harvell et al., 1999; Sutherland et al., 2004). At least four coral diseases
(black-band disease, white plague, dark-spots disease, and Aspergillosis) are associated with high water
temperatures (Kuta and Richardson, 1996; Bruckner et al., 1997; Richardson et al., 1998; Gill-Agudelo and
Garzon-Ferriera, 2001; Alker et al., 2001). Nutrient input, sedimentation, and runoff have also been implicated
as potential contributing factors in the initiation and elevated virulence of a disease, although few quantitative
data have been published (Bruckner et al., 1997; Harvell et al., 1999; Kim and Harvell, 2001; Richardson and
Aronson, 2002).

It appears that the ability of corals and other organisms to withstand infection has been compromised by
climate change, eutrophication, sedimentation (Rogers, 1990), and other human-induced ecosystem pertur-
bations (Knowlton, 2001). The vulnerability of tropical coral reef ecosystems is related to the fact that many
warm water corals grow slowly and persist only within a narrow range of light, temperature, dissolved oxygen
and salinity fluctuations, and, in an evolutionary sense, they are thought to have a limited ability to recover
from disease (Knowlton, 2001). However, the relative importance of anthropogenic influences is still unclear,
especially since disease outbreaks are being reported with increasing frequency on reefs that exist in areas
relatively far from the direct effects of human activity (Bruckner and Bruckner, 2004).

A decline in the health of many coral reefs worldwide has created an urgent need for multidisciplinary studies
of coral health and disease, with emphases on coral physiology, biology, and disease etiology, including mech-
anisms of resistance and susceptibility to disease, factors affecting the transmission, spread and virulence of
pathogens, and relationships between environmental factors and disease. By better understanding causative
agents and factors responsible for the emergence and proliferation of diseases, scientists will be able to con-
tribute to the development of strategies that can be used by resource managers to mitigate disease impacts.


Tropical Storms
Most coral reef environments are found in tropical climates and periodically experience cyclonic storm events.
Cyclonic storms are an important process in the structure and dynamics of coral reef ecosystems (Hughes and
Connell, 1999). They are classified as "pulse disturbances" since they are typically intense and of relatively
short duration, yet are a powerful mechanism for change and can dramatically disrupt ecosystems, com-
munities, population structure, resource availability, and the physical environment (Pickett and White, 1985).
Coral reefs, however, are often located in dynamic regions of the ocean and have clearly shown resilience to
historical bouts of disturbance. In fact, such disturbances are thought to maintain high species diversity, par-
ticularly when the disturbance alters the structure of the reef by opening up bare substratum, thereby creating


F




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


space available for the settlement of new coral recruits (planulae). The influence of disturbance in community
structure and dynamics has been illustrated by the intermediate disturbance hypothesis, which states that the
highest number of species in a community will occur at intermediate levels (frequency and size) of natural
disturbance. Lower diversity will exist where disturbances are either very large or very small, or very frequent
or very infrequent (Connell, 1978, 1979). The size of the new space also influences the type of recruitment.
Small patches are usually colonized by the nearest dominant species, while larger areas provide an opportu-
nity for less dominant species to establish. Interestingly, many Caribbean corals release planulae in late sum-
mer/early fall, which coincides with the hurricane season in the Atlantic, and this may enhance recolonization
(Rogers, 1993).

The effect of storms is strongly dependent on the ecology and geology of a specific area and the characteris-
tics of the storm. For instance, a wide range of reef-specific variables influence the magnitude of the impact
including spatial location, community structure, coral age, size, morphology, and reef depth. Variables associ-
ated with the storm itself include the path of the storm and its strength (measures of wind velocity and wave
height), and heavy rain can cause excessive runoff as well as localized decreases in salinity which have been
linked to a reduction in the planulae production (Figure 3.4; Jokiel, 1985). Some species of corals exhibit
a growth form that is more robust to
storm energy than others (e.g., boul-
der shapes). In contrast, corals with
fragile skeletons and typically those
with branching morphology will be
more easily damaged by extreme
wave action. In the Caribbean, Acro-
pora palmata and Acropora cervi-
cornis are very susceptible to storm
damage (Brown, 1997). Breakages
may be advantageous to these spe-
cies since they produce relatively
few larvae and instead are thought
to rely primarily on asexual reproduc-
tion through fragmentation to pro-
duce new colonies (Bak and Engel,
1979; Hughes, 1985). Furthermore,
delayed mortality from outbreaks of
disease among injured corals, bio-
erosion of damaged skeleton, and Figure 3.4. Hurricane Georges, a category 3-4 storm hit the USVI, Puerto Rico,
Sre portion s and the Florida Keys in September, 1998. Damage included the physical breakage
altered predator-prey relationships of corals and a massive pulse of sediment and nutrients that were discharged into
may occur for years after a hurricane nearshore waters. Georges was one of four hurricanes in progress in the Caribbean
has struck (Knowlton et al., 1990). at the time. Photo: NASA and NOAA, http://rsd.gsfc.nasa.gov/rsd/images/Georges.
html, Accessed 2/10/05.
Age is another factor that influences the ability of a coral colony to withstand the mechanical stresses of large
storms. As corals grow, they become more vulnerable to breakage and dislocation (Brown, 1997). The ma-
jority of wave impacts occur in the shallowest (0-20 m) depth range, so corals at greater depths are generally
less directly impacted. Deeper corals, however, can be significantly damaged indirectly by large blocks that
tumble down from shallower waters (Brown, 1997). Damage to corals can indirectly impact other reef-associ-
ated organisms through the reduction of coral cover and topographic complexity which influence biological
interactions such as predation, succession, and competition. As coral cover is reduced, the refuge function for
many fish and invertebrates is diminished. Also the removal of organisms from substrate via scouring reduces
the abundance of food available for some species. In addition, increases in turbidity and sedimentation that
often accompany storms can affect the emergent community by impairing photosynthesis and feeding, and
limiting sexual reproduction (Kojis and Quinn, 1985).

The direct effects of cyclones on fish are size-specific. Lassig (1983) noted that during the final stages of Cy-
clone Peter, many fish that were normally associated closely to the benthos were found in the water column




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


and some had fresh wounds. This suggests that fish try to weather the storm in the water column, where they
are less likely to be injured. It was also noted that after the storm, overall fish abundance decreased signifi-
cantly, with juveniles sustaining higher mortality than adults due to strong storm-driven currents.

To understand how a cyclonic storm affects a reef requires examination of the recovery patterns and process-
es. Detailed comparative investigations of pre- and post-hurricane coral reef ecosystems that include vari-
ables such as amount of coral, number of species, settlement characteristics and growth rates, and nutrient
cycling may provide valuable insights. Multiple year trends using continuous monitoring data, however, are
likely to provide the most accurate assessment of both short- and longer-term impact and recovery (Hughes
and Connell, 1999). The trajectory and rate of recovery will be influenced by a number of interacting factors
including the rates of recruitment, species involved, and sequence of colonization (Brown, 1997). Research
also suggests that anthropogenic impacts can interfere with the recovery process. Finally, separating storm
effects from those caused by direct human activity and phenomena such as coral bleaching and competition
with algae, is problematic due to the level of degradation of some reef systems (Brown, 1997).

The terms "hurricane" and "typhoon" are regionally specific names for a strong tropical cyclone. This report
follows the geographically-specific naming convention recognized by NOAA (i.e., NOAA Research's Hurricane
Research Division, http://www.aoml.noaa.gov/hrd/tcfaq/Al.html, Accessed 01/07/05) whereby the term "hur-
ricane" applies to the North Atlantic Ocean, Northeast Pacific Ocean east of the dateline, and South Pacific
Ocean east of 160E; "typhoon" applies to the Northwest Pacific Ocean west of the dateline; "cyclone" applies
to the Southwest Pacific Ocean west of 160E and Southeast Indian Ocean east of 90E. The characteristics
of storm and hurricane categories are given in Table 3.3.

Table 3.3. The Saffir-Simpson scale for tropical storm and hurricane classification and associated storm characteristics provide a
consistent way to characterize major storm events. na=not applicable. Source: NOAA National Hurricane Center.



Tropical Depression na 20-34 kts 23-39 mph 1007 mb na
Tropical Storm na 35-64 kts 39-74 mph 1006-1000 mb na
Hurricane 1 65- 82 kts 74- 95 mph 980-999 mb minimal
Hurricane 2 83-95 kts 96-110 mph 965-979 mb moderate
Hurricane 3 96-113 kts 111-130 mph 945-964 mb extensive
Hurricane 4 114-135 kts 131-155 mph 920-944 mb extreme
Hurricane 5 >135 kts >155 mph 919 mb catastrophic


Coastal Development and Runoff
In the past several decades, there has been a well-documented demographic shift toward higher concentra-
tions of human settlement in the coastal zones of many countries including the U.S. (Culliton et al., 1990;
Figure 3.5). More than half of the U.S. population now lives in coastal counties, a trend that is expected to
continue to increase (Pew Oceans Commission, 2003; Cicin-Sain et al., 1999). This trend has increased the
frequency and magnitude of impacts from activities such as the construction of residential developments,
hotels and resorts, recreational facilities, and infrastructure such as roads and wastewater treatment plants
(WVVTPs).

Terriginous sediments in runoff from construction sites and roads are often a major threat to nearshore areas.
Dredging of nearshore sediments for marina facilities, ship access and navigation, beach nourishment, and
building materials can introduce significant quantities of particulate matter into the water column. While strong
currents tend to dissipate some of the added sediments, nearshore areas with gentle slopes and low flushing
rates tend to accumulate sediments, which can have detrimental effects on sessile invertebrates like corals
(Rogers, 1990). Physical smothering may be the most obvious effect of sedimentation. Although most cor-
als have some ability to rid themselves of foreign particles, the removal of sediments requires the diversion
of energy from vital activities such as reproduction and feeding. The negative effects of the accumulation of




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


,*~









Guam and s.6-11.e%
Southern 11.63-20.8%
Islands of .20.8%
CNMI
Southern
o United States





Main Hawaiian
Islands





American Puerto Rico and the
Samoa U.S. Virgin Islands
Figure 3.5. Coastal population change between 1990 and 2000 and associated development pressure pose a significant threat to
coral reef ecosystems, particularly in island jurisdictions with limited land area. Maps not drawn to scale. Maps: K. Buja. Data: U.S.
Census, 1990, 2000; Secretariat of the Pacific Community, http://www.spc.org.nc/prism, Accessed 2/15/05.

sediments on corals can be exacerbated by wave action that repeatedly resuspends sediments into the water
column (Rogers, 1990). Increased turbidity in the water column, whether episodic or chronic, reduces light
availability for photosynthesis and growth. Increases in nearshore sediment loads have been shown to affect
morphology of corals and gorgonians as well as inhibit the development and recruitment of coral larvae (Rog-
ers, 1990). Coral species react differently to this stressor, and coral reefs in waters experiencing increased
turbidity may exhibit a shift in community composition toward greater dominance of corals that are more toler-
ant of lower light levels and better adapted to remove sediments.

Alteration of watersheds and associated changes in vegetative cover often decrease the ability of the land to
absorb rainfall, which flows through streams and channels, carrying sediments and pollutants into nearshore
areas. Generally, runoff from developed watersheds carries higher sediment loads than from undeveloped
areas, and this is more pronounced in areas where the topography is characterized by steep slopes. Removal
of mangrove forests that normally trap sediments may allow a greater proportion of terriginous sediments to
reach reef ares.

In addition to sediments, runoff from developed watersheds tends to have higher concentrations of waste
products. Increased freshwater inputs are actually considered pollutants as they can decrease the salinity
levels in some nearshore areas. Other contaminants derived from human use of nearshore areas include oil
leaking from vehicles, pesticides and lawn fertilizers applied to yards, parks and golf courses, chemicals in
asphalt that wash off roads, excrement from livestock and domesticated animals, and litter.




stems of the United States and Freely Associated States: 2005


The development of infrastructure is also a major concern. In many areas, coastal development often oc-
curs without a commensurate improvement in the wastewater infrastructure, and existing systems cannot
adequately accommodate the added burden. As a result, untreated or partially-treated sewage overflows into
nearshore areas. Outside of urban areas, many homeowners are not able to access VVVTPs and often must
rely on septic tanks, which are subject to corrosion and leakage. The hard-to-detect leaks often allow untreat-
ed sewage to seep into groundwater and nearshore waters. A recent report (Carter and Burgess, Inc., 2002)
assessing the sustainability of tourism in Hawaii noted that many of the island's municipal wastewater systems
are nearing capacity. While most new developments have private WVVWTPs to satisfy permit conditions, many
residents still rely on private systems, such as septic tanks, which are in various stages of disrepair. Though
they considered myriad aspects of tourism, the authors of the study contend that such nonpoint source pollu-
tion is "one of Hawaii's greatest environmental threats" (Carter and Burgess, Inc., 2002).

Other infrastructural issues include the problems of adequate waste disposal and the construction of docks
and piers that can result in habitat loss. In summary, coastal development presents a wide range of chal-
lenges for coastal areas, especially in terms of the number and scale of construction projects, capabilities of
infrastructure, intensity and type of land use, and increases in sedimentation and pollution levels.


Coastal Pollution
Worldwide, the threat to coral reef ecosystems from pollution is surpassed in severity only by coral bleach-
ing and fishing (Spalding et al., 2001). Model estimates indicate 22% of the world's coral reef ecosystems
are threatened by land-based pollution and soil erosion (Bryant et al., 1998). Pollution often desensitizes
the ecosystem, so that it becomes more susceptible to other stressors such as climate change, disease, and
invasive species. The primary stressors from land-based sources are nutrient and chemical pollution from
fertilizers, herbicides, pesticides, human-derived sewage, and increased amounts of sediment from coastal
development and storm water runoff. Other pollutants, such as heavy metals and oil, can also be prominent
at specific locations.

This section focuses on point source pollution. Point sources of pollution originate from confined or discrete
conveyances, such as a pipe, tunnel, ditch, channel, well, or fissure. Examples of point source pollution in-
clude sewage outfalls, factory wastewater, and dumping of chemicals. Household chemicals and untreated
industrial wastewater may also be discharged into the domestic wastewater stream. Finally, short outfalls
contribute to the pollution of nearshore waters. Other point sources include vessels without holding tanks that
discharge their wastes in marinas and nearshore coastal areas. Dredging for shipping lanes, marinas, and
coastal construction projects resuspends sediments that increase turbidity and decrease coral reef ecosys-
tem productivity. Industrial point sources include manufacturing operations, effluent discharges, accidental
oil spills and the release of contaminants discharged as a byproduct of oil-drilling (e.g., toxic poly-aromatic
hydrocarbons (PAHs), benzene, ethylbenzene, xylene) and heavy metals, such as lead, copper, nickel and
mercury.

Direct impacts of pollutants include reduced recruitment, loss of biodiversity, altered species composition (a
shift from predominantly phototrophic to heterotropic fauna), and shallower depth distribution limits. Sew-
age pollution causes nutrient enrichment around population centers, treatment facilities, and sewage outfalls.
Increased nutrient concentrations promote increased algal and bacterial growth, can degrade seagrass and
coral reef ecosystems, and ultimately may decrease fisheries production. Sediments smother benthic organ-
isms, which can become diseased when exposed to dredged sediments contaminated with toxic heavy metals
and organic pollutants. Toxic chemicals can decrease coral reef ecosystem productivity and biodiversity and
increase human health risks through food contamination.

Management actions by NOAA to address water quality concerns are taken in partnership with the Environ-
mental Protection Agency (EPA), the Department of Agriculture, and local or state governments. Research is
needed to understand how coral reef ecosystems respond to poor water quality, and to provide mangers with
tools to detect, assess, and remedy negative impacts from pollution. Therefore, the sources of the substances
that adversely affect water quality must be identified, and relevant policies and control strategies for limiting
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The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


programs should be integrated into modeling efforts that quantify the relative amounts of natural and anthropo-
genic inputs to ecosystems Additionally, monitoring results should be used to develop models and indicators
that assess threats or identify stressors causing coral reef ecosystem decline.


Tourism and Recreation
Tourism and recreation are by far the fastest growing sector of coastal area economies. This growth is predict-
ed to continue as incomes rise, more Americans retire, leisure time expands and accessibility to the coasts and
oceans increases (U.S. Commission on Ocean Policy, 2004). Coral reefs, in particular, have a major economic
value. Cesar et al. (2002) calculated that the greatest contribution to the annual value of coral reefs in Hawaii
is tourism and recreation, which brings in $304 million per year. Coastal tourism contributes $9.9 billion to the
Californian economy annually and is considered the largest sector of the "ocean industry" compared with $6
billion/year for ports, $860 million/year for offshore oil and gas development, and $550 million/year for fisheries
and mariculture (Wilson and Wheeler, 1997; Cicin-Sain and Knecht, 2000). Travel and tourism are estimated
to have provided $746 billion annually to the U.S. gross domestic product (GDP), making travel and tourism
the second largest contributor to GDP (Houston, 1995). Tourism is particularly significant in many Caribbean
and Pacific islands surrounded by coral reef ecosystems. In the Florida Keys alone, over four million tourists
purchase about $1.2 billion in services annually. Over three million tourists visit at least one of Hawaii's coral
reef sites per year, and approximately 90% of new economic development in Guam and the CNMI is related
to coastal tourism (NOAA, 1997). The vast demand for tourism and recreational services associated with
coral reefs generates considerable income for many local communities. Those who engage in reef-related
recreational activities purchase goods and services, such as charter boats and diving trips via dive centers. In
addition, they spend money on lodging, travel, food and beverages, etc. English et al. (1996) estimate an an-
nual economic impact of $1.2 billion in visitor spending in the Florida Keys which results in a total sales impact
of $1.3 billion, $506 million in income, and over 33,000 jobs. Leeworthy and Wiley (1997) estimate an annual
economic impact of $94.3 million in resident spending in the Florida Keys, resulting in a total sales impact of
$105.6 million and supporting over 2,400 jobs. Cesar et al. (2002) estimated that recreational use values in
Hawaii represent 85% of annual benefits accrued from coral reefs (the others being amenity/property values,
biodiversity, fisheries, and educational spillover), which amount to $304.16 million/year. In southeast Florida,
the annual use value accrued from coral reefs is estimated at $229.3 million (Johns et al., 2003).

Human uses of coral reefs are both direct and indirect, with recreation and tourism among the most promi-
nent uses. Recreational activities on U.S. coral reefs include snorkeling, scuba diving, boating, fishing, and
shell-collecting. The intensity of each activity varies widely from region to region, but can be considerable
in some areas. In southeast Florida, residents and visitors spent 28 million person-days using artificial and
natural reefs during a 12 month pe-
riod (June 2000 to May 2001) and
4.94 million person-days snorkeling
and scuba diving (Johns et al., 2003;
Figure 3.6). Water-based activities
such as scuba diving are increasing
in popularity, and over 3 million peo-
ple are currently certified to dive in
the U.S. Scientific studies have now
shown that divers and snorkelers can
have a significant negative impact on
coral reefs in terms of physical dam-
age and a concomitant reduction in
their aesthetic appeal (Hawkins and
Roberts, 1993; Hawkins et al., 1999;
Rouphael and Inglis, 2001). For ex-
ample, a snorkeling trail created in
the Virgin Islands National Park's
Trunk Bay in the 1960s had deterio- Figure 3.6. Some reef areas in the Florida Keys may have hundreds of visitors per
rated substantially when observed in day. Photo: Bill Harrigan.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


1986 with visitor numbers estimated at over 170,000 per year. Only 10 of 50 tagged Elkhorn coral colonies re-
mained undisturbed during a seven-month period of observation (Rogers et al., 1988). Plathong et al. (2000)
examined the effects of snorkelers using self-guided interpretative trails around a reef within the Great Barrier
Reef Marine Park, Australia and found that despite comparatively low levels of use (approximately 15 snorkel-
ers per trail per week), snorkelers caused significant damage to corals along the trails. Hawkins et al. (1999)
examined the impacts of diving on a reef off the Caribbean island of Bonaire and concluded that impacts would
be minimized by maintaining a site carrying capacity of between 4,000 and 6,000 dives per year. In contrast,
Rouphael and Inglis (2002) suggested that management actions should focus on identifying and mitigating the
causes of damaging behavior rather than setting numerical limits to site use.

Concern has also been directed at the activity of fish-feeding. Feeding fishes negatively impacts both fishes
and habitat in several ways including: (1) fish consume food that is very different to their normal diets; 2) the
concentration of fish at feeding stations disrupts normal distribution/abundance patterns; (3) fish behavior
changes with some individuals or aggregations exhibiting abnormal aggression; and (4) inputs of nutrients
and incidental damage to benthic structure can result in an increase of macroalgae (Perrine, 1989; Alevizon,
2004).

In addition to these direct threats, indirect threats can be equally, if not more devastating to coral reefs. In-
direct threats include development of hotels and resorts, construction of the infrastructure needed to support
such resorts, seafood consumption, beach replenishment, construction of airports and marinas, as well as the
operation of cruise ships. The impacts resulting from these activities include increased sedimentation, nutri-
ent enrichment, pollution, exploitation of endangered species, and increased litter and waste (UNEP, 2002).
Mitigation of the impacts of tourism often involves education and raising awareness with the goal of behavioral
change (UNEP, 2002). In Hawaii, a strategy for both defining a carrying capacity and influencing visitor be-
havior through education has been implemented. Oahu's Hanauma Bay Nature Preserve in Hawaii has an
estimated three million visitors annually and 13,000 per day in the high season. Impacts at Hanauma Bay,
including widespread trampling of reefs and resuspension of sediments, fish-feeding, littering, and other pollu-
tion, prompted a management strategy to limit visitor numbers (NOAA CSC, 2004). Determining the carrying
capacity for this area was critical to its long-term sustainability and was supported by the development of an
education center aimed at influencing visitor behavior (Cesar et al., 2002).

Clearly, tourism is a major source of economic welfare and livelihood for many coastal communities. Unfor-
tunately, detrimental side effects and physical damage often result from direct visitor activity and the devel-
opment of facilities to support tourism. Without long-term planning for tourist activities at these fragile sites,
both resource and revenues are at risk. Sites such as Hanauma Bay Nature Preserve have had to make
operational adjustments and offer education and instruction to visitors. Managers are increasingly challenged
to develop strategies that mitigate unsustainable usage, while continuing to support the tourism industry.


Fishing
Coral reefs and associated habitats support important commercial and recreational fisheries. Over 4,000 spe-
cies of fishes (>25% of all marine fishes) inhabit shallow coral reefs (Spalding et al., 2001), along with a large
number of marine plants and invertebrates many of which are exploited for human use. Coral reef fisher-
ies support and sustain communities by providing food and sources of income. Fishing also plays a central
social and cultural role in many island communities. Coral reef fisheries are generally small-scale, but coral
reef fishers exploit hundreds of species of fishes and invertebrates using a wide variety of fishing gear. In a
number of U.S. reef areas, recreational fishery catch now equals or exceeds the commercial catch. The rich
biodiversity of coral reefs also supports a valuable marine aquarium industry, especially in Hawaii and Florida,
and provides materials for a range of natural products developed by the biotechnology and pharmaceutical
industries.

Unfortunately, these fishery resources and the ecosystems that support them are under increasing threat
from overfishing and fishery-associated impacts on habitats and ecosystems. Fishery-related impacts include:
1) direct overexploitation of fish, invertebrates, and algae for food and the aquarium trade; 2) removal of a




tes and Freely Associated States: 2005


species or group of species which can impact multiple trophic levels; 3) by-catch and mortality of non-targ
species; and 4) physical impacts to reef environments associated with fishing techniques, fishing gear, ar
anchoring of fishing vessels.

Overfishing
Overfishing, along with pollution and global climate change, is generally considered to be one of the grec
est threats to the health of coral reefs. It is also the most widespread threat, estimated to be of medium
high threat to over 35% of the world's reefs (Bryant et al., 1998). In many cases, significant depletion of re
resources (especially large fishes and sea turtles) had already occurred before 1900 (Jackson et al., 200
Pandolfi et al., 2003). Since then, increases in coastal population, improved fishing technology, and ove
capitalization of fishing fleets driven by demand from rapidly growing export markets have greatly accelerate
resource depletion. Many reef fishes have relatively slow growth rates, late maturity, and irregular recruitme
- characteristics that make overexploitation more likely. The trend is for high-value or vulnerable resource<
- generally large predators such as groupers, jacks and sharks to be removed first, and then target specih
further down the food chain are subsequently fished (Pauly et al., 1998).

Overfishing has been identified as a major concern in all U.S. states and territories with coral reefs and h<
been identified by the USCRTF as a priority reason for the development of local action strategies. In most ca
es, the large number of species in these multi-gear, small-scale fisheries has made it impractical to condu
standard stock assessments for more than a fraction of the species (see Table 3.4), and such data-intensiv
single-species approaches have been criticized as unrealistic for most reef fish systems (Sale, 2002). The
is evidence of serial depletion of reef resources in Florida and around all populated U.S. islands. In Haws
long-term catch rates suggest that stocks of nearshore fishes have declined by nearly 80% between 1900 ar
the mid-1980s (Shomura, 1987). Catch per unit effort (CPUE) of reef fishes in Guam fell by more than 50
between 1985 and 2000 (Birkeland et al., 2000), while the CPUE fell 70% in the American Samoan reef fisher
accompanied by a shift in species composition, over a period of 15 years between 1979 and 1994 (Birkelan
1997). The Nassau grouper fishery, the highest value commercial fishery in Puerto Rico and the U.S. Virg
Islands (USVI), collapsed in the 1980s due to overexploitation of spawning aggregation sites and the special
was identified as a candidate to be listed as threatened or endangered under the Federal Endangered Specii
Act (16 U.S.C. 460 et seq.) in 1991. In the Florida Keys, the nation's most extensive and long-term reef fi,
monitoring program has revealed that 77% of the 35 individual stocks that could be analyzed in Biscayne B&
are overfished (Ault et al., 2001).

Table 3.4. Overfished Coral Reef Species in Federal Fishery Management Plans (FMPs). Source: 2003 Status of U.S. Fisheri
Report (NOAA, http://www.nmfs.noaa.gov/sfa/reports/html, Accessed 2/14/05) and Western Pacific Coral Reef Ecosystem Fishe
Management Plan (NOAA, http://www.wpcouncil.org/coralreef.htm, Accessed 2/14/05).





South Atlantic2 62 8 12 42
Gulf of Mexico2 44 5 4 35
Caribbean2 154 3 1 150
Western Pacific3 28 0 0 28
Total 422 16 16 389
Notes:
1 Overfished analysis includes only stocks in Federal waters-most reefs and fishing pressure occur in state and territorial waters.
2 Excludes coral species for which the fishery is closed.
3 From the Bottomfish, Precious Coral and Crustacean FMPs only-does not include the hundreds of species covered by the new
Coral Reef Ecosystem FMP.

Because of long-term trends in the exploitation of mixed reef fisheries, there are few places that maintain rel
tively intact fish populations to serve as experimental controls. The Northwestern Hawaiian Islands (NWF
and some of the uninhabited U.S. Pacific Remote Island Areas probably represent the closest approximate
to unexploited coral reef ecosystems in U.S. waters. The average fish biomass in the NWHI is 2.6 timi




The State of Coral Reef Ecosystems of


eater than in the Main Hawaiian Islands (MHI). More than 54% of the total fish biomass in the NWHI is
imposedd of apex predators, compared to less than 3% in the MHI. These differences have been attributed
overfishing in the MHI (Friedlander and DeMartini, 2002).

ecosystem Shifts
iere is increasing evidence that overfishing on reefs results not just in shifts in fish size, abundance, and
)ecies composition, but that it is also a major driver altering the ecological balance and contributing to the
,gradation of coral reef ecosystems (Bellwood et al., 2004). In particular, overfishing of herbivorous fishes
is been linked to phase-shifts from high-diversity coral-dominated systems to low-productivity algal-dominat-
I communities (Hughes, 1994). U.S. reefs, especially in the Atlantic, are increasingly facing coral declines,
ough uncertainty remains about the processes and links to fishing levels, especially in the Pacific (Jennings
id Polunin, 1997). Herbivores comprise a significant component of the catch in the MHI, Guam, CNMI,
id American Samoa. Parrotfishes and surgeonfishes are increasingly important in Puerto Rico and in St.
roix, where they represent the predominant catch. In nearly all areas except Florida, declines in the abun-
ince of these species have been observed. There is also evidence that heavy fishing pressure on certain
vertebrate-feeding fishes has played a key role in outbreaks of crown-of-thorns (COTS) starfish, snails, and
wrbivorous sea urchins (Hay, 1984; McClanahan, 2000; Dulvy et al., 2004). There is no clear evidence of the
:tent to which this has been an important factor in bioerosion on U.S. reefs, nor is there a clear understanding
the ecosystem effects due to the removal of top predators. Overfishing can also compound the impact of
her threats. For example, overfishing of herbivorous fishes and enhanced nutrient flows to reefs may lead to
ef overgrowth by macroalgae. Likewise, reefs devoid of herbivores may be less likely to recover from coral
teaching events (Westmacott et al., 2000).

pacts from Fishing Gear
number of protected species, such as hawksbill and green sea turtles as well as a number of seabird species
e untargeted victims of fishing activity and are especially vulnerable to longline fishing and shrimp trawling.
aps and gill nets also result in mortality of non-target species.

iysical damage to the benthos from certain fishing techniques is well-documented. Traps set for fishes or
bsters can cause physical damage to corals, gorgonians, and sponges. They may also result in by-catch
id "ghost fishing" if they are lost or not regularly checked. Trap fisheries are most common in Florida (lobster
id stone crab) and the U.S. Caribbean (fish and lobster), and are generally less prevalent in the U.S. Pacific.
irge gill and trammel nets have also been identified as a growing concern, particularly in St. Croix (USVI)
id Hawaii. Large gill nets are set on reefs and their lead-lines can cause extensive damage when the nets
e hauled into the boats. In addition to legal fishing activities, illegal techniques can cause severe damage
reefs. Use of chlorine bleach has been reported in Hawaii, Guam, and Puerto Rico (USCRTF, 1999), and
additional plant-derived poisons are still used occasionally in the subsistence fishery in American Samoa.
ie use of cyanide for fishing has not been reported on U.S. reefs, although the expansion of the live food
,h trade to the Marshall Islands has raised concerns about its potential use there. Blast fishing, probably the
ost destructive technique, has rarely been reported on U.S. reefs.

their indirect impacts to coral reefs associated with fisheries include anchor damage from fishing boats,
iich has been identified as a problem in Florida and the U.S. Caribbean. Trawling damage to coral areas
is been identified as a problem in deeper coral areas in the Gulf of Mexico. It was also a major cause of
destruction of the deep water Oculina coral banks off the east coast of Florida before the development of the
,hing, but trawls can cause tremendous damage when hauled over hard bottoms with coral. Furthermore,
oundings of fishing vessels have had major, albeit localized, impacts on certain reefs.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


Trade in Coral and Live Reef Species
Many coral reef species are harvested domestically and internationally to supply a growing international de-
mand for seafood, aquarium pets, live food fish, construction materials, jewelry, pharmaceuticals, traditional
medicines and other products. In many locations, collection is occurring at unsustainable levels, and overhar-
vesting may lead to reductions in the abundance and biomass of target species, shifts in species composition,
and large-scale ecosystem shifts including population explosions of non-target species or the replacement of
thriving, coral-dominated systems with low-productivity algal reefs (Hughes, 1994; McClanahan, 1995; Jen-
nings and Polunin, 1996). In addition to overfishing, there is widespread use of destructive techniques such as
cyanide poisoning of fishes and coral colony breakage. Cyanide is used illegally in Southeast Asia and other
parts of the Indo-Pacific to capture live reef fish for the aquarium trade and live fish markets, and has been
found to: 1) kill many non-target species, 2) cause habitat damage, and 3) pose human health risks (Barber
and Pratt, 1997). High levels of mortality associated with cyanide and inadequate handling and transport
practices pose significant challenges to achieving sustainability. The use of cyanide has not been reported or
observed in the U.S., with the possible exception of limited use in some of the Freely Associated States (e.g.,
Marshall Islands) associated with the live reef fish food trade. In addition, unsafe diving practices resulting
from the collection of corals, sea cucumbers, fish, and other species in deep water are causing a high inci-
dence of illness, paralysis, and even death of collectors in some regions (Johannes and Riepen, 1995; Barber
and Pratt, 1997).

The Marine Aquarium Trade
The marine aquarium trade has an estimated value of $200-300 million per year (Larkin and Degner, 2001).
The global trade in coral has increased by 500% over the last 10 years, with over one million live corals and
1.87 million kg of live rock traded in 2002 (Bruckner, 2003). In addition, an estimated 20-24 million reef fishes
are traded annually, representing 1,450 species in 50 families (Balboa, 2002; Wabnitz et al., 2003). The U.S.
is the world's largest consumer of ornamental coral reef species, importing 60-80% of the live coral, over 50%
of the curio coral, 95% of live rock, and 50-60% of the marine aquarium fishes each year (Wood, 2001; Bruck-
ner, 2003). The most important sources of coral are currently Indonesia, Fiji, and Vietnam (Bruckner, 2001).
Indonesia and the Philippines each supply about 30% of the total global trade in reef fishes, with another 30%
exported from five locations (Brazil, the Maldives, Hawaii, Sri Lanka, and Vietnam); Florida and Puerto Rico
are currently the largest exporters from the wider Caribbean (Wood, 2001; Balboa, 2002).

Although it is illegal to harvest stony corals and live rock in U.S. waters, ornamental reef fishes and many mo-
tile invertebrates are collected in U.S. waters both for domestic use and export. In Florida, 318 marine species
(181 fishes and 137 invertebrates) have been collected for commercial purposes, with a total annual value of
up to $4.2 million. Over 200,000 ornamental reef fishes are landed in Florida each year, with a maximum of
425,781 fishes in 1994 (Larkin, 2003). Annual reported harvest of ornamentals from West Hawaii rose from
90,000 in 1973 to 422,823 in 1995 (Tissot and Hallacher, 1999).

The Live Reef Food Fish Trade
Groupers, humphead wrasse, coral trout, and other large fishes that use coral reefs are harvested live to
supply restaurants in Hong Kong. Exports increased rapidly during the 1990s and peaked at 32,000 metric
tons (mt) in 1997, with a slight decline between 1998 and 2000 due to the Asian economic crisis (Lau and
Parry-Jones, 1999). More recently an estimated 22,000 to 28,000 mt of live reef fishes have been imported by
Hong Kong, China, Taiwan, and other Asian markets, with Hong Kong imports comprising 65-80% of the total
regional trade (Graham et al., 2001). In addition to widespread use of cyanide to capture the fish live, fishers
target spawning aggregations and have been reported to eliminate entire breeding populations relatively rap-
idly (Lau and Parry-Jones, 1999). In addition to concerns regarding the use of destructive fishing techniques,
most of these species are vulnerable to heavy fishing pressure due to their longevity, late sexual maturation,
aggregation spawning, and sex change habits (Sadovy et al., 2004).

Curios and Jewelry Trade
Coral reef species harvested for curios and jewelry include mollusk shells; stony coral skeletons; and black,
pink, gold, bamboo, and other precious corals (Figure 3.7). Of these species, only stony corals, black coral,
and giant clams are internationally regulated through the Convention on International Trade in Endangered
Species of Wild Fauna and Flora (CITES). International trade in shells involves as many as 5,000 species




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


of an unknown volume primarily sup-
plied by the Philippines, Indonesia,
Thailand, Singapore, Taiwan, Mexico,
India, Africa, and Haiti (Wells, 1989).
Shells are used for construction ma-
terials; shell craft; mother of pearl and
other collectors items; as well as ad-
ditives to floor tiles, toothpaste, pot-
tery, and poultry feed (Marshall et al.,
2001). The volume of trade in coral
skeletons has varied over the years,
with the Philippines being the major
supplier in the 1970s and 1980s; ex-
ports from the Philippines were pro-
hibited in the late 1980s, with a tem-
porary lifting of trade bans in 1992
during which over three million kg
were exported. Fiji and Vietnam are Figure 3.7. The shells of reef organisms are often sold at curio shops, such as this
currently the major source countries one in Palau. Although many of the shells were probably imported from Southeast
for coral skeleton (Bruckner, 2001). Asia, some local collection is thought to occur as well. Photo: J. Waddell.

International trade in black coral, according to the CITES trade database, has averaged 430,000 items per
year since 1983, with the maximum trade in 1994, and 320,000 items traded in 1998 (CITES Trade Database,
http://www.cites.org/eng/resources/trade.shtml, Accessed 02/16/05). The world's largest supplier of worked
black coral is Taiwan (>90% of the total), with most reported to be harvested in the Philippines. Commercial
harvest occurs in U.S. waters in Hawaii, with annual landings averaging 1,014 kg/year; about 90% of this is
for domestic use.

International Protection
CITES is an international agreement among the governments of 165 countries to protect wildlife by ensuring
that international trade does not threaten the survival of a species in the wild. CITES regulates international
trade in wildlife according to three levels, or appendices, of threat. Species listed in Appendix I, which includes
marine turtles and most whales, are believed to be threatened with extinction and thus, commercial trade of
these species is generally prohibited. Most species are listed in Appendix II which includes organisms that
are not presently threatened or endangered, but may become so if trade is not regulated. These species can
still be commercially traded with export permits which require the exporting country to ensure that the species
was legally harvested and its export will not be detrimental. Coral reef species currently listed in Appendix II
include about 2,000 species of stony corals (including all scleractinian corals), black coral, giant clams, queen
conch, and seahorses. Trade of Appendix III species requires an export permit ensuring that the organism
was harvested legally and prepared and shipped so as to minimize damage, injury or cruel treatment.


Ships, Boats, and Groundings
Of all physical damage caused to coral reefs by human activity, ship groundings and the impacts of boats and
anchors are perhaps the most destructive. The U.S. Coast Guard (USCG) reports that over 2,100 grounding
accidents are reported annually, with about 440 vessels sinking each year. In addition, over 800 abandoned
barges litter the inland and coastal waters of the U.S., many still loaded with hazardous cargo (Helton, 2003).
As recreational and commercial boating traffic increases in nearshore ocean waters, these shipwrecks pose
a threat to coral reef habitat. When anchors, especially the enormous anchors of cruise ships, are carelessly
dropped and dragged on fragile reef, hundreds of meters of habitat can be destroyed. Recent studies dem-
onstrate the extensive impacts of groundings when hazardous cargo is released. However, once cargo and
fuel are spilled, the vessel may continue to cause repeated physical damage to the reef due to movement by
wind and waves. Furthermore, abandoned barges can often become illegal dump sites for other hazardous
materials, trap wildlife, and become public safety hazards (Helton and Zelo, 2003).




The State of Coral Reef Ecosystems of the United States and A


Initially many considered the impacts of grounded vessels to be significant only at a local level, but the wide-
spread effects of these events have recently been the subject of closer examination (Precht et al., 2001;
Ebersole, 2001). Damage resulting from ship groundings often continues well beyond the initial event of
impact as a result of slow recovery and fragmentation of keystone species essential to reef structure and
function. In particular, spur and groove reefs do not seem to recover their diverse fish assemblages following
a ship grounding incident (Ebersole, 2001). The potential threats of grounded vessels became the subject
of increased political attention in 1999 when nine vessels were cleaned, cut apart, and removed from a reef
in Pago Pago, American Samoa and the grounding sites were restored by the USCG, NOAA, and American
Samoan government. The increasing frequency of vessel groundings in coral reef environments led to the de-
velopment of the National Action Plan to Conserve Coral Reefs (USCRTF, 2000) which recognizes the impact
of grounded vessels to coral reefs and their associated habitats (Helton and Zelo, 2003). In response, NOAA
initiated the Abandoned Vessel Project, which seeks to increase awareness of abandoned vessels, particular-
ly where they occur in coral reef systems, as well as provide the technical assistance necessary to remove the
vessels (NOAA OR&R, http://response.restoration.noaa.gov/dac.vessels/overview.html, Accessed 6/2/04).

A study conducted on the site of the 1984 grounding of the M/V Wellwood in the Florida Keys National Marine
Sanctuary suggested that damaged spur and groove habitat will take decades to recover without substantial
restoration efforts (Smith et al., 1998). A reduction of topographic complexity also influences local hydrody-
namics and the structure of reef fish and invertebrate communities (Miller et al., 1993; Szmant, 1997).

The damage caused to a coral reef habitat by boat anchors is an additional threat resulting from frequent boat
traffic. A study conducted in a 220 ha area of coral reef in Fort Jefferson National Monument, Dry Tortugas,




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


ment in the area of the grounding and may consequently hinder recovery of the community (Negri et al.,
2002).

With boat traffic rapidly growing, it is crucial to better understand the ecological implications of vessel ground-
ings and anchor damage, and to take steps to limit or prevent damage through education and guidance sup-
ported by strong legislation. Severe physical damage to coral reefs by vessels requires a rapid response and
carefully designed methods of removal and restoration to limit the extent of the impact (NOAA, 2002b).


Marine Debris
Globally, marine debris presents a continuous threat to the marine environment. Marine debris adversely im-
pacts marine life through the destruction of essential habitat as well as entanglement and ingestion by marine
organisms and seabirds. Typically, the majority of marine debris comes from land-based sources, particularly
urban centers, but a significant proportion comes from ships.

All U.S. jurisdictions with coral reefs
participate in the International Coast-
al Cleanup to remove marine debris
from their shorelines and nearshore
waters. Additional community-based
cleanup efforts have been conducted
at many locations, including South
Point and Kahoolawe in Hawaii. Typ-
ical debris collected from the shore-
lines includes beverage cans and
bottles, cigarettes, disposable light-
ers, plastic utensils, food wrappers,
and fishing line (Figure 3.9). Under-
water cleanup conducted by snor-
kelers and divers have found similar
materials beneath the surface.

The most notable impacts of ma-
rine debris on coral reef ecosystems
come from derelict fishing gear in-
cluding nets, fishing line, and traps.
Prior to the 1950s, fishing gear was
composed of natural fibers, such as
cotton and linen, and was susceptible
to environmental degradation. Since
the 1950s, fishing gear has primarily
been constructed with synthetic ma-
terials, such as nylon and polyethyl-
ene, which is less susceptible to en-
vironmental degradation. Synthetic Figure 3.9. Tons of marine debris wash up on the shores of the NWHI every year.
Though NOAAs Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Di-
nets and fishing line can persist in the vision has removed 401,055 kg of debris from the shallow waters of the NWHI since
ocean for decades and can be trans- 2001, resource limitations prevent debris removal on land. Photo: S. Hoist.
ported for thousands of kilometers.

The NWHI has been a focal point for the removal of abandoned fishing gear comprised of conglomerates of
netting and fishing line that roll across coral reef habitats, crushing corals and dislodging sessile organisms
(Figure 3.10). Fishing gear frequently becomes snagged on corals and continues to trap fish ("ghost fishing")
and endangered monk seals and sea turtles (Boland and Donohue, 2003; Donohue et al., 2001; Henderson,
2001; Balazs, 1985). Since 2001, NOAA's Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Di-
vision (PIFSC-CRED) has led a large-scale interagency partnership to study and remove derelict fishing gear
from the NWHI. NOAA collaborates with the State of Hawaii, City and County of Honolulu, U.S. Fish and Wild-




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


life Service (USFWS), USCG, U.S. 0 i
Navy, University of Hawaii, Hawaii
Sea Grant, Hawaii Metals and Re-
cycling, Honolulu Waste Disposal,
and other partners from local agen-
cies, businesses, and non-govern-
mental organizations. From 2001 to
2004, this large-scale effort removed
401,055 kg of fishing gear from these
remote islands and atolls (R. Brain-
ard, pers. comm.). Types of fishing
gear removed included monofilament
gillnet, seine net, and trawl nets, the
majority of which was thought to
have originated from fisheries oper-
ating around the continental shelves
of the North Pacific Rim which are
located thousands of kilometers from
the NWHI.

Derelict fishing gear has also been a
concern in other U.S. coral reef eco-
systems. Chiappone et al. (2002)
surveyed the Florida Keys for fish-
ing gear and other marine debris
and concluded that lobster trap de-
bris was often found in offshore and
mid-channel patch reefs, while hook
and line gear was more common in
shallow and deep forereef areas.
Since 1994, the FKNMS, The Nature
Conservancy, The Bacardi Founda-
tion, and local dive operators have Figure 3.10. A tangle of abandoned fishing gear removed from Pearl and Hermes
supported an annual effort to clean Atoll in the NWHI by a team of divers from PIFSC-CRED and the Joint Institute for
Marine and Atmospheric Research (JIMAR). The net had to be freed from the reef,
the reefs around the Florida Keys. In lifted to the surface, and towed to shallow water before debris team members could
2002, divers removed over 1,800 kg cut it into smaller pieces and remove it. Photo: A. Hall.
of marine debris including fishing line
from the Keys. In 2003 and 2004, Amigos de Amona, Inc. and other partners removed 3,235 kg of marine
debris from the islands in Puerto Rico's Mona Channel. The debris consisted of fishing gear (48%), plastics
(13%), glass (14%), metal (8%), and miscellaneous items such as refrigerator doors, rubber shoes, packing
and insulation materials, and washing machines (17%; Amigos de Amona, Inc., 2004).


Aquatic Invasive Species
Aquatic invasive species are aquatic organisms that have been introduced, either intentionally or unintention-
ally, into new ecosystems which result in harmful ecological, economic, and human health impacts (USDA,
http://www.invasivespecies.gov, Accessed 2/11/05). Aquatic invasive species have been reported in all U.S.
reions and probably exist in every region of the world. Invasive species are generally second only to habitat
destruction in causing declines in biodiversity and are thought to impact nearly half of the species currently
listed as threatened or endangered under the Federal Endangered Species Act (Wilcove et al., 1998).

The impacts are not only ecological. Damages to infrastructure, such as clogged intake pipes, and environ-
mental losses due to terrestrial and aquatic invasive species cost over $120 billion per year in the U.S. alone
(Pimentel et al., in press). The cumulative effects and costs of aquatic invasive species are difficult to quan-
tify, but evidence clearly indicates that the impacts will continue to increase. In fact, the frequency of aquatic






invasions has increased exponentially since the late 1700s and shows no signs of diminishing (Ruiz et al.,
2000).

Although there have not been many studies that focus specifically on the impacts of aquatic invasive spe-
cies on shallow-water coral reef ecosystems as a whole, there have been a handful of smaller studies. In
Hawaii, it has been determined that the number of marine and estuarine invasive species is approximately
343, including 287 invertebrates, 24 algae, 20 fish, and 12 flowering plants (Bishop Museum, http://www2.
bishopmuseum.org/HBS/invertguide, Accessed 02/14/05). Pearl Harbor alone contains more than 100 inva-
sive species. Additionally, some of Hawaii's worst invaders have been intentionally introduced, such as algal
species, Kappaphycus alvarezii and K. striatum, which smothered large tracts of coral reefs in Kaneohe Bay,
thus diminishing the ecological and economic value of the area (Carlton, 2001).

Shallow-water coral reef ecosystems are particularly sensitive to a number of non-native species introduction
pathways, including ships (due to ballast water discharges and hull fouling), aquaculture of non-native spe-
cies, releases by aquarium hobbyists, and marine debris.

Introductions from Ballast Water
By 1996, 80% of all commercial goods were being transported aboard ocean-going vessels (NRC, 1996).
That percentage is likely to increase as global trade increases. In addition to greater movement of goods
across the world's oceans, the speed and size of ships have greatly increased, resulting in faster voyages
and larger volumes of ballast water. Because most marine species have planktonic stages as part of their life
cycle, they are subject to entrainment during the uptake and discharge of ballast water. Furthermore, because
voyage times have greatly decreased, the chances of survival are greater. Ballast tanks have been shown
to carry bacteria, protests, dinoflagellates, diatoms, zooplankton, algae, benthic invertebrates (e.g., mollusks,
corals, sea anemones, and crustaceans), and fish (LaVoie et al., 1999; NRC, 1996).

Releases by Aquarium Hobbyists
Although there are relatively few documented marine fish invasions, 94 of the 241 documented invasions
involved tropical marine species. Additionally, a link has been identified between invasions and marine aquar-
ium imports. Such findings highlight the susceptibility of warm water coral reef ecosystems to intentional intro-
ductions by hobbyists and the need for public education. For example, a species of lionfish (Pterois volitans)
common to the Indo-Pacific regions that was thought to have been introduced from a home aquarium in 1992
has established viable populations all
along the southeastern coast of the
U.S., with juveniles recently found as
far north as Long Island (Figure 3.11;
Whitfield et al., 2002).

Introductions from Marine Debris
The amount of marine debris gen-
erated as waste from society has
increased at a rapid rate in recent
years (Silvia-lniguez and Fischer,
2003; Moore, 2003). For instance,
the amount of marine debris in the
waters around Great Britain doubled
from 1994 to 1998 (Barnes, 2002).
Much of the debris is fisheries re-
lated, comprised mostly of netting.
Floating material provides habitat for
many organisms and can result in the
transportation of species into new ar-
eas, often many thousands of kilome- Figure 3.11. The Red Lionfish, Pterois volitans, is native to the Indo-Pacific but has
ters from their existing species range established viable populations along the southeastern coast of the U.S. This fish was
(Barnes and Fraser, 2003). Problems photographed off the coast of Beaufort Inlet, North Carolina in about 40m of water.
Photo: P. Whitfield.
occur when newly arrived alien spe-




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


cies successfully colonize and overwhelm local marine ecosystems. Barnes (2002) found that marine debris
was typically colonized by bryozoans, barnacles, polychaetes, hydroids and mollusks.


Security Training Activities
U.S. military installations near coral reefs include operations in Hawaii (Hickam Air Force Base, Pearl Harbor,
and Kaneohe Bay); Johnston Atoll (PRIAs); Wake Atoll (PRIAs); Kwajelein Atoll (Republic of the Marshall Is-
lands); Guam; CNMI; Key West and Panama City, Florida; Puerto Rico; USVI; Cuba; and Diego Garcia in the
Indian Ocean. Military bases and associated activities including exercises, training, and operational proce-
dures (i.e., construction, dredging, and sewage discharge) have the potential for adverse ecological impacts
on coral reefs such as excessive noise, explosives and munitions disposal, oil and fuel spillage, wreckage and
debris, breakage of reef structure, and non-native species introductions from ship bilge water or aircraft cargo
(Coral Reef Conservation Guide for the Military, https://www.denix.osd.mil/denix/Public/ES-Programs/Conser-
vation/Legacy/Coral/coral.html, Accessed 12/6/04).

In recent years, the military has decommissioned several properties and transferred management responsi-
bility to other agencies. In June 1997, the U.S. Navy officially turned over the management of Midway Atoll
(NWHI) to the USFWS for use as a national wildlife refuge. Parts of the island required major remediation
to mitigate contamination by lead-based paints, asbestos, fuels and chemicals, but the refuge soon offered
fishing, diving, and eco-tour opportunities. When the military decommissioned Kaho'olawe, a former naval
bombing range in the MHI, they established a framework for cleanup that included government-appropriated
funds and a transfer of the island to a native Hawaiian organization with a state-appointed council to oversee
the cleanup process. In June 1995, an evaluation of the nearshore coral reef resources of Kaho'olawe docu-
mented the continued presence of metal debris, but reported that relatively few pieces of ordnance were found
despite many years of bombing exercises on the island (Naughton, 1995). The 10-year, $460 million cleanup
on Kaho'olawe ended November 11, 2003. At that time, the Navy ceased active remediation and access
control was returned to the State of Hawaii. The Navy continued surface clearance as a further risk reduc-
tion measure until April 2004 when final demobilization occurred. At that point, full-time management of the
island shifted to the state. In May 2003, the U.S. Navy ceased military training on the eastern side of Vieques
Island, Puerto Rico and transferred management of all remaining Navy property on Vieques, including the
bombing training range on the easternmost parcel, to the USFWS. According to the statute governing such
transfers, the property can only be used as a wilderness area. Vieques and the surrounding waters have been
proposed by the U.S. Environmental Protection Agency (EPA) for listing on the National Priorities List, which
EPA uses to determine which uncontrolled waste sites warrant further investigation. As such, the Navy, EPA,
and Puerto Rico Environmental Quality Board will work cooperatively on conducting investigations required
by the Comprehensive Environmental Response, Compensation and Liability Act (42 U.S.C. 9601 et seq.).
The investigation may conclude the need for the Navy to complete hazardous substances remediation and/or
munitions clearance in some areas. Baseline assessments of 24 permanent coral reef monitoring sites at
Vieques Island were commissioned by the U.S. Navy and completed in 2001-2002 in an effort to comply with
Executive Order 13089 and the U.S. Department of Defense (DoD) Initiative for Coral Reef Protection at the
Roosevelt Roads Naval Station in Puerto Rico (Deslares et al., 2004).

According to the DoD Coral Reef Implementation Plan (2000), U.S. military services (i.e., the Air Force, Army,
Navy, and Marine Corps) "generally avoid coral reef areas in their normal operations except for some mission-
essential ashore and afloat activities." DoD policy is to avoid adversely impacting coral reefs during military
operations and ensure safe and environmentally responsible action in and around coral reef ecosystems, to
the maximum extent practicable. However, exceptions to this policy can be made during wars; national emer-
gencies; and threats to national security, human health, and the safety of vessels, aircraft, and platforms (Ex-
ecutive Order 13089, 1998). DoD has implemented a number of actions to comply with natural resource and
environmental protection laws, and has developed programs to protect and enhance coral reef ecosystems.
These efforts include developing geographic information system (GIS) planning tools, coral surveys to evalu-
ate impacts from bombing exercises, assessments to determine the impact of amphibious training exercises
on reef ecosystems, pollution and oil spill prevention programs, and invasive species management and effec-
tive land management programs (Defense Environmental Network and Information Exchange, https://www.
denix.osd.mil, Accessed 2/14/05).




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


Oil and Gas in Coral Reef Ecosystems
The introduction of oil and other hydrocarbons into the marine environment can have serious consequences for
coral reef ecosystems. Whether from chronic or episodic oil spills or from activities related to the exploration,
production or transport of energy resources, oil can impact reefs through physical breakage, sedimentation
and smothering, toxic contamination by heavy metals, and by inhibition of growth and recruitment. Sources of
oil entering the marine environment vary. Summary information for North America is provided in Figure 3.12.

Once introduced, oil tends to persist OffshoreOil and Gas
Development (including
in sheltered tropical coastal envi- Dp.mest)
ronments. Because of the difficulty Recreatonal
of navigation in shallow-water coral Marine Vese, Atmoseic Fallout
2% from Human Aolvities
reef environments, cleanup follow- a/
ing a spill is often extremely diffi- Marine Transporttion
cult. Booms and skimmers can be
used in lagoon areas when the oil is
on the surface, but these responses
become less useful over time as the
oil combines with mineral particles X I : i dsntaaste
in the water and sinks or is churned andRunoff
into the water column during inclem-
ent weather. The use of dispersants Natural Seepae
is often discouraged in shallow-water
areas because they cause the oil to
sink to the bottom where it comes
into contact with sensitive reef habi-
tats. Reduced water circulation in
Figure 3.12. Sources of oil entering the marine environment of North America.
nearshore areas hinders natural dis- Source: Minerals Management Service, 2002.
sipation by currents. When spills oc-
cur in shallow-water coral reef ecosystems, the best option may be to let natural processes handle the task of
removing oil from the fine sediments of mangrove forests, seagrass meadows, and complex reef frameworks
(Corredor et al., 1990; Guzman et al., 1994). Oil spill recovery in shallow-water reef ecosystems can require
decades. Five years after a major oil spill on a Panamanian reef (April 1986), scientists found that surviving
colonies of the four most massive species of reef-building corals were still experiencing extensive, chronic ef-
fects on vital processes (Guzman et al., 1994).

Several studies have been undertaken to determine the impact of oil on the physiology of coral reef organisms
(reviews in Shigenaka, 2001). Laboratory experiments have demonstrated that exposure of coral species to
oil can result in decreased growth, reproduction, and colonization capacity, as well as other negative effects on
feeding, behavior, and mucous cell function (IPIECA, 1992). Afield study in the Gulf of Eilat, Red Sea demon-
strated that repeated discharges of oil onto a coral reef caused many changes to the reef system as a whole,
and in particular damaged the reproductive system of scleractinian corals (Rinkevich and Loya, 1979).

In southern Florida, Dustan et al. (1991) evaluated the impacts of drilling wells on reef building corals, gor-
gonians, sea grasses, macroalgae, and reef fishes. Primary impacts included physical destruction by drilling
machinery and the accumulation of drilling debris, although no organisms appeared to be damaged by drill-
ing fluids or cuttings. The results implied that exploratory drilling, in light of present technology and stringent
dumping regulations, may be achieved without leaving lasting impacts; however, no conclusions could be
drawn from this study relative to the drilling production wells (Dustan et al., 1991).

In the North Sea, Olsgard and Gray (1995) assessed the spatial and temporal effects of production discharges
on benthic fauna along contamination gradients. Results suggested that discharges reduced abundance of
benthic fauna, many of which were key prey species for bottom-living fish. The fauna that became established
in the contaminated sediments was considered less valuable as a food source for fish populations.

In addition to spills, exploration for offshore oil and gas reserves has the potential to have major impacts on
marine ecosystems. Petroleum resources are difficult to find, and the process of locating, recovering, trans-




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

ferring and transporting them can pose a significant potential hazard to species living in the surrounding area.
In the early stages, exploration for oil and gas involves seismic testing which involves emitting loud booming
Shock waves in order to determine what lies under the seafloor. The impacts of seismic testing on marine or-
Sganisms are not well understood (The Ocean Conservancy, 2003). Once oil and gas reserves are located, en-
ergy exploration and production requires platform installation; dredging; drilling; the discharge of liquid, solid,
and gaseous wastes and drill cuttings; noise and light pollution; and polluted air emissions. These impacts,
in addition to the physical effects related to the movement of ships and equipment, can all present significant
threats to the environment where the activity is taking place (http://earthsci.org/energy/gasexpl/exproil.html,
Accessed 6/25/04).

The primary drilling areas in the U.S. Exclusive Economic Zone that occur near reef ecosystems are in the
Gulf of Mexico, where major development has resulted in the installation of 6,500 production platforms and
over a 160,900 km of pipelines and other infrastructure. Numerous wells, platforms and pipelines surround
the Flower Garden Banks National Marine Sanctuary (FGBNMS) in the northwestern Gulf of Mexico (see
Chapter 8), and one oil production platform even lies within the boundaries of the sanctuary, less than 1.6 km
from the East Flower Garden coral cap. Fortunately, FGBNMS managers report that no major spills or impacts
have occurred to date within sanctuary waters.

Because oil and gas development is such a major activity on the outer continental shelf in the Gulf of Mexico,
the U.S. Department of the Interior's Minerals Management Service (MMS) has supported mapping and study
programs of the Flower Garden Banks since the early 1970s to determine how to mitigate environmental
impacts of oil and gas exploration. Information from these studies has supported MMS's belief that lease
stipulations can minimize the potential impact of discharged contaminants to reef communities in the area.
One such important stipulation requires shunting of drill cuttings so that they are deposited within 10 m of the
bottom and not further up in the water column (MMS, http://www.mms.gov/eppd/compliance/13089/banks.
htm, accessed 6/25/04).

Furthermore, removal of the enormous platforms, which weigh thousands of tons, is nearly impossible without
the use of explosive materials. Gitschlag and Herczeg (1994) conducted one of the few known observations
of fish mortality following such explosive activity. They reported that one event killed as many as 51,000 fish
(larvae and juveniles were not counted). Removal of structures may also decrease the availability of habitat
for fish that utilize the sites as artificial reefs (Patin, 2004, http://www.offshore-environment.com/abandon-
ment.html, accessed 6/24/04).
























I




ie United States and Freely Associated States: 2005


Other Threats
Crown-of-Thorns Starfish Outbreaks
The COTS (Acanthaster plan)
is a species of echinoderm found
throughout the Indo-Pacific region
(Figure 3.13). COTS feeds on sev-
eral common species of hard coral,
particularly Acropora spp., showing
a clear preference for tabular forms
and those corals that are least well
defended (De'ath and Moran, 1998;
Pratchett, 2001). They reproduce
sexually with synchronized release
of gametes and have a remarkable
ability to regenerate damaged parts.
COTS is preyed upon by several
species of fish including triggerfish
(Balistidae), and pufferfish (Tetradon-
tidae), and a few large crustaceans
and mollusks. At relatively low den-
sities, the starfish are considered to
play an important role in maintain- Figure 3.13. A closeup of a crown-of-thorns starfish, Acanthaster planci, on a reef
ing high diversity on coral reefs (Ar- in the PRIAs. Photo: J. Maragos.
onson and Precht, 1995). At many
locations, however, populations periodically increase to levels that result in the degradation of coral reefs.
Aggregations of hundreds of thousands of individuals have been reported across the Indo-Pacific, including
Australia's Great Barrier Reef, Fiji, Micronesia, American Samoa, the Cook Islands, the Society Islands, the
Ryukyu Islands (Japan), Hawaii, Malaysia, the Maldives, and the Red Sea. The rate of recovery after a ma-
jor outbreak is highly variable, with full recovery estimated to take decades or even many hundreds of years
(Sano, 2000; Lourey, 2000).

A number of environmental factors have been considered causative in COTS outbreaks, including hurricanes,
nutrient input, and overfishing (Birkeland, 1982; Ormond et al., 1991). The level of impact from human activity
is still unclear since outbreaks have also been reported in remote areas with very little human activity. Nev-
ertheless, stressors generated through human activity are likely to influence the trajectory and rate of post-
outbreak recovery.

Outbreaks of other echinoderms, such as spiny sea urchins (Echinoidea), can also adversely impact coral reef
ecosystems through excessive erosion of coral substratum, removal of newly settled corals, and intense her-
bivory (Sammarco, 1982; Carreiro-Silva and McClanahan, 2001). Damage to coral reefs due to high density
populations (12-100 urchins/m2) of urchins have been occasionally reported in U.S. waters including Hawaii,
USVI, and the Marshall Islands.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


Earthquakes and Volcanoes
Many islands in the Pacific and Carib-
bean were formed and transformed
through tectonic and volcanic activity.
In fact, coral reef atolls are formed
through the erosion and subsidence
of volcanoes and the subsequent
gradual upward growth of coral reefs
(Darwin, 1842). Volcanic eruptions
can have important direct and indi-
rect consequences for coral reefs.
The eruption of Mt. Pagan, CNMI in
1981 resulted in extensive damage to
coral communities due to scouring by
lava and smothering by volcanic ash,
although observation of new coral
recruits indicated recovery occur-
ring within two years of the eruption
(Eldredge and Kropp, 1985). Simi-
larly, rapid recovery was observed Figure 3.14. In the past few years, eruptions of the volcanic island Anatahan in CNMI
after high coral mortality as a result have deposited tons of ash on nearby reefs and temporarily closed international air-
ports in Saipan and Guam. The latest major cluster of eruptions occurred in April
of burial by ash after the 1994 erup- 2005. Photo: NASA, MODIS sensor.
tion of Rabaul Caldera in Papua New
Guinea (Maniwavie et al., 2001). Major eruptions can also impact coral reefs many thousands of kilometers
away through a complex sequence of events (Figure 3.14). For example, the 1991 eruption of Mount Pinatubo
in the Philippines led to a short-term atmospheric cooling throughout the Middle East during the winter of 1992.
This abnormal cooling resulted in deep vertical mixing in the Gulf of Eilat and excessive nutrient upwelling,
which in turn, triggered algal blooms causing widespread coral death (Genin et al., 1995). However, cooled
larva flow can also create new habitat suitable for the settlement and growth of corals and other organisms.

In 1993, an earthquake measuring 8.2 on the Richter scale caused collapse of some coral reefs around Guam
and also destroyed some large coral colonies that had formed on unstable substrata (Birkeland, 1997). Earth-
quakes that uplift some areas while subsiding others, or even triggering catastrophic sedimentary events, are
thought to be important factors in the present spatial patterns of fringing reefs in the Gulf of Aqaba, Red Sea
(Shaked et al., 2004). In the Hawaiian archipelago, a high frequency of deep earthquakes combined with
submergence and rising sea-level may explain the absence of coral reefs in some locations around the island
of Hawaii.

Cable-laying Operations
There has been a rapid increase in the need for submarine cables, particularly fiber optic cables, to support
the telecommunications industry. Cable-laying operations and the movements of unsecured cables have
been found to disrupt and destabilize benthic structure (Sultzman, 2002). The impact of laying a cable on
benthic habitats will depend on the location of landing points, route chosen, and installation process. In some
instances, sand channels through reefs have been used, but damage has occurred where cables have been
laid directly over corals. Coral transplants and artificial reef modules have been used to replace lost hard
coral, yet little is known about the effectiveness of these methods. Furthermore, few restoration efforts have
considered damage to non-scleractinian components of the biota.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005


REFERENCES

Alevizon, W 2004. Divers feeding fishes: a continuing issue in MPA management. MPA News 6 (5). Available from the internet
URL: http://depts.washington.edu/mpanews/MPA58.htm.

Alker, A.P., G.W Smith and K. Kim. 2001. Characterization of Aspergillus sydowii (Thom et Church), a fungal pathogen of
Caribbean sea fan corals. Hydrobiologia 460: 105-111.

Amigos de Amon6, Inc. 2004. Final Report fo the Mona Channel Marine Debris Removal, Puerto Rico. Prepared for the
National Fish and Wildlife Foundation and funded through the NOAA Coral Reef Conservation Program in 2002. Cabo Rojo,
Puerto Rico.

Aronson, R.B., I.G. Macintyre, C.M. Wapnick and WM. O'Neill. 2004. Phase Shifts, Alternate States, and the Unprecedented
Convergence of Two Reef Systems. Ecology 85 (7): 1876-1891.

Aronson, R.B. and WF. Precht. 1995. Landscape patterns of reef coral diversity: a test of the intermediate disturbance hypoth-
esis. Journal of Experimental Marine Biology and Ecology 192: 1-14.

Aronson, R.B. and WF. Precht. 2001. White-band disease and the changing face of Caribbean coral reefs, Hydrobiologia 460
(1-3): 25-38.

Ault, J.S., S.G. Smith, G.A. Meester, J. Juo and J.A. Bohnsack. 2001. Site Characterization for Biscayne National Park: As-
sessment of Fisheries Resources and Habitats. NOAA Technical Memorandum NMFS-SEFSC-468. 156 pp.

Bak, R.P.M. and M.S. Engel. 1979. Distribution, abundance and survival of juvenile hermatypic corals (Scleractinia) and the
importance of life history strategies in the parent coral community. Marine Biology 54: 341-352.

Balazs, G.H. 1985. Impact of ocean debris on marine turtle in the Hawaiian Islands. pp. 387-429. In: R.S. Shomura and H.O.
Yoshida (eds.) In: Proceedings of the Workshop on the Fate and Impact of Marine Debris. NOAA Technical Memorandum
NOAA-TM-NMFS-SWFC-45.

Balboa, C.M. 2002. The United States consumption of ornamental fish: a preliminary analysis of import data. Final report for
NOAA grant NFFKP300-2-00010. World Resources Institute, Washington, DC. 46 pp.

Barber, C.V. and V.R. Pratt. 1997. Sullied Seas: Strategies for Combatting Cyanide Fishing in Southeast Asia and Beyond,
World Resources Institute. 57 pp.

Barnes, D.K.A. 2002. Invasions by marine life on plastic debris. Nature 416 (6883): 808-809.

Barnes, D.K.A. and K.P.P. Fraser. 2003. Rafting by five phyla on man-made flotsam in the Southern Ocean. Marine Ecology
Progress Series 262: 289-291.

Belausteguigoitia, J.C 2004. Causal chain analysis and root causes: The GIWA approach. AMBIO 33 (1-2): 7-12.

Bellwood, D.R., TP. Hughes, C. Folke and M. Nystrom. 2004. Confronting the coral reef crisis. Nature 429: 827-833.

Birkeland, C.E. 1982. Terrestrial runoff as a cause of outbreaks of Acanthaster planci (Echinodermata: Asteroidea). Marine
Biology 69: 175-185.

Birkeland, C.E. (ed.) 1997. Life and death of coral reefs. Chapman and Hall, New York. 536 pp.

Birkeland, C.E. 1997. Status of coral reefs in the Marianas. pp. 91-100. In: R.W Grigg and C. Birkeland (eds.) Status of Coral
Reefs in the Pacific. Sea Grant College Program, University of Hawaii.

Birkeland, C.E., P. Craig, G. Davis, A. Edward, Y Golbuu, J. Higgins, J. Gutierrez, N. Idechong, J. Maragos, K. Miller, G. Paulay,
R. Richmond, A. Tafileichig and D. Turgeon. 2000. Status of coral reefs of American Samoa and Micronesia: U.S.-affiliated and
Freely Associated islands of the Pacific. pp. 199-217. In: C. Wilkinson (ed.) Status of Coral Reefs of the World 2000. Australian
Institute of Marine Science.

Boland, R.C. and M.J. Donohue. 2003. Marine debris accumulation in the nearshore marine habitat of the endangered Hawai-
ian monk seal, Monachus schauinslandi 1999-2001. Marine Pollution Bulletin 46 (11): 1385-1394.

Brainard, Russell. Pacific Islands Fisheries Science Center, Coral Reef Ecosystem Division. Honolulu, HI. Personal commu-
nication.

Brown, B.E. 1997. Disturbances to reefs in recent times. pp. 354-379. In: C. Birkeland (ed.) Life and Death of Coral Reefs.
Kluwer Academic Publishers.

Bruckner, A.W 2001. Tracking the Trade in Ornamental Coral Reef Organisms: The Importance of CITES and its limitations.
Aquarium Sciences and Conservation 3 (1-3): 79-94.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

Bruckner, A.W. 2003. Sustainable Management Guidelines for Stony Coral Fisheries. pp. 167-184. In: Cato, J. and C. Brown
(eds.) Marine ornamental species collection, culture and conservation. Blackwell Scientific. Iowa State University Press,
Iowa.

Bruckner, A.W 2004. Coral health and mortality: Recognizing the signs of coral diseases and predators. Undersea Journal.
First quarter 2004.

BrucknerA.W, R.J. Bruckner and E.H. Williams. 1997. Spread of a black-band disease epizootic through the coral reef system
in St. Ann's Bay, Jamaica. Bulletin of Marine Science 61: 918-928.

Bruckner, A.W and R.J. Bruckner. 2003. Condition of coral reefs off less developed coastlines of Curagao (stony corals and
algae). Atoll Research Bulletin 496: 370-393.

Bruckner, A.W and R.J. Bruckner 2004. Impact of yellow-band disease (YBD) on Montastraea annularis (species complex)
populations on remote reefs off Mona Island, Puerto Rico. Abstract. In: Proceedings of the 10th International Coral Reef Sym-
posium.

Bryant, D., L. Burke, J. McManus and M. Spaulding. 1998. Reefs at Risk: A Map-Based Indicator of Threats to the World's
Coral Reefs. World Resources Institute Report. 56 pp

Bythell, J., M. Barer, R. Cooney, J. Guest, A. O'donnell, O. Pantos and M. Le Tissier. 2002. Histopathological methods for the
investigation of microbial communities associated with disease lesions in reef corals. Letters in Applied Microbiology 34 (5):
359-364.

Carlton, J.T 2001. Introduced species in U.S. Coastal waters: Environmental Impacts and Management Priorities. Pew Oceans
Commission, Arlington, FL.

Carreiro-Silva, M. and TR. McClanahan. 2001 Echinoid bioerosion and herbivory on Kenyan coral reefs: the role of protection
from fishing. Journal of Experimental Marine Biology and Ecology 262 (2): 133-153.

Carter and Burgess, Inc., 2002. Planning for Sustainable Tourism in Hawaii. Part 1: Infrastructure and Environmental Overview
Study. Prepared for the State of Hawaii Department of Business, Economic Development and Tourism. Honolulu. 141 pp.

Cesar, H., P. van Beukering, S. Pintz and J. Dierking. 2002. Economic Valuation of the Coral Reefs of Hawaii: Final Report (FY
2001-2002). Hawaii Coral Reef Initiative Research Program. University of Hawaii, Honolulu.

Chiappone, M., A. White, D.W Swanson and S.L. Miller. 2002. Occurrence and Biological Impacts of Fishing Gear and Other
Marine Debris in the Florida Keys. Marine Pollution Bulletin 44: 597-604.

Cicin-Sain B., R.W Knecht and N. Foster (eds.). 1999. Trends and Future Challenges for U.S. National Ocean and Coastal
Policy. National Oceanic and Atmospheric Administration, Silver Spring, Maryland. 142 pp.

Cicin-Sain, B and R.W Knecht. 2000. The Future of U.S. Ocean Policy: Choices for the New Century. Island Press, Washing-
ton, DC. 416 pp.

CENR (Committee on Environment and Natural Resources, Subcommittee on Ecological Systems). 2001. Ecological Fore-
casting: Agenda for the Future. Washington, DC. 12 pp.

Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199: 1302-1310.

Connell, J.H. 1979. Tropical rain forests and coral reefs as open non-equilibrium systems. pp. 141-163. In: R.M. Anderson,
B.D. Turner and L.R. Taylor (eds.) Population Dynamics, Symposium of British Ecological Society. Blackwell Science Publica-
tions, Oxford, England.

Corredor, J.E., J.M. Morell and C.E. del Castillo, 1990. Persistence of spilled crude oil in a tropical intertidal environment. Ma-
rine Pollution Bulletin 21: 385-388.

Culliton, TJ., M.A. Warren, TR. Goodspeed, D.G. Remer, C.M. Blackwell and J.J. McDonough, II. 1990. 50 years of popula-
tion change along the Nation's coasts, 1960-2010. National Oceanic and Atmospheric Administration Coastal Trends Series.
Rockville, MD. 41 pp.

Darwin, C. 1842. The Structure and Distribution of Coral Reefs. Being the First Part of the Geology of the Voyage of the
'Beagle.' Smith, Elder and Co., London.

Davis, G.E. 1977. Anchor Damage to a Coral Reef on the Coast of Florida. Biological Conservation 11: 29-34.

De'ath G. and P.J. Moran. 1998. Factors affecting the behaviour of crown-of-thorns starfish (Acanthasterplanci) on the Great
Barrier Reef 2: Feeding preferences. Journal of Experimental Marine Biology and Ecology 220 (1): 107-126.

Deslarez, K.J.P., R. Nawojchik, D.J. Evans, C.J. McGarrity and P. Gehring. 2004. The condition of fringing reefs off former mili-
tary bombing ranges at Isla de Culebra and Isla de Vieques, Puerto Rico. Report prepared forthe U.S. Department of Defense,
U.S. Navy. 24 pp.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

Donohue, M.J., R.C. Boland, C.M. Sramek and G.A. Antonelis. 2001. Derelict Fishing Gear in the Northwest Hawaiian Islands:
Diving Surveys and Debris Removal in 1999 Confirm Threat to Coral Reef Ecosystems. Marine Pollution Bulletin 42 (12):
1301-1312.

Dulvy, N.K., R.P. Freckleton and N.V.C. Polunin. 2004. Coral reef cascades and the indirect effects of predator removal by
exploitation. Ecology Letters 7: 410-416.

Dustan, P., B.H. Lidz and E.A. Shinn, 1991. Impact of Exploratory Wells, Offshore Florida: A Biological Assessment. Bulletin of
Marine Science 48 (1): 94-124.

Ebersole, J.P. 2001. Recovery of Fish Assemblages From Ship Groundings On Coral Reefs in the Florida Keys National Ma-
rine Sanctuary. Bulletin of Marine Science 69 (2): 655-671.

Eldredge, L.G. and R.K. Kropp. 1985. Volcanic ashfall effects on intertidal and shallow water coral reef zones at Pagan, Mari-
ana Islands. pp. 4: 195-200. In: Proceedings of the 5th International Coral Reef Congress.

English, D., W.A. Kriesel, V.R. Leeworthy and P.C. Wiley. 1996. Economic Contribution of Recreating Visitors to the Florida
Keys/Key West. Athens, GA: USDA Forest Service, Southern Research Station, Outdoor Recreation and Wilderness Assess-
ment Group. University of Georgia, Department of Agricultural and Applied Economics, Athens, GA; and National Oceanic and
Atmospheric Administration, Strategic Environmental Assessments Division, Silver Spring, MD. 22 pp.

Feely, R.A., C.L. Sabine, K. Lee, W Berelson, J. Kleypas, V.J. Fabry and F.J. Millero. 2004. Impact of anthropogenic CO2 on
the CaCO3 system in the oceans. Science 305 (5682): 362-366.

Friedlander, A.M. and E.E. DeMartini. 2002. Contrasts in density, size, and biomass of reef fishes between the northwester and
the main Hawaiian islands: the effects of fishing down apex predators. Marine Ecology Progress Series 230: 253-264.

Genin A., B. Lazar and S. Brenner. 1995. Atmospheric cooling, unusual vertical mixing and coral mortality following the eruption
of Mt. Pinatubo. Nature 377: 507-510.

Gill-Agudelo, D. and J. Garz6n-Ferreira. 2001. Spatial and seasonal variations of dark-spots disease in coral communities of
the Santa Marta area (Columbian Caribbean). Bulletin of Marine Science 69: 619-630.

Gitschlag, G.R. and B.A. Herczeg. 1994. Sea turtle observations at explosive removals of energy structures. Marine Fisheries
Review 56 (2): 1-8.

Graham, T 2001. A collaborative strategy to address the live reef food fish trade. Asia-Pacific Coastal Marine Program, Report
0101. The Nature Conservancy, Honolulu.

Green, E.P. and A.W Bruckner. 2000. The significance of coral disease epizootiology for coral reef conservation. Biological
Conservation 96 (3): 347-361.

Guzman, H.M., K.A. Burns and J.B.C. Jackson, 1994. Injury, regeneration and growth of Caribbean reef corals after a major
oil spill in Panama. Marine Ecology Progress Series 10: 231-241.

Harvell, C.D., K. Kim, J.M. Burkholder, R.R. Colwell, P.R. Epstein, D.J. Grimes, E.E. Hofmann, E.K. Lipp, A.D.M.E. Osterhaus,
R.M. Overstreet, J.W Porter, G.W Smith and G.R. Vasta. 1999. Emerging Marine Diseases--Climate Links and Anthropogenic
Factors. Science 285:1505-1510.

Hawkins, J.P. and C.M. Roberts. 1993. Effects of Recreational scuba diving on coral reefs: trampling on reef-flat communities.
Journal of Applied Ecology. 30: 25-30.

Hawkins, J.P., C.M. Roberts, T Van'T Hof, K. De Meyer, J. Tratalos and C. Aldam. 1999. Effects of Recreational Scuba Diving
on Caribbean Coral and Fish Communities. Conservation Biology 13 (4): 888-897.

Hay, M.E. 1984. Patterns of fish and urchins grazing on Caribbean coral reefs: are previous results typical? Ecology 65: 446-
454.

Helton, D. 2003. Wreck Removal: A Federal Perspective. In: National Salvage Conference Proceedings. Available from the
internet URL: http://response.restoration.noaa.gov/dac/vessels/docs/Heltonsalvage2003.pdf.

Helton, D. and I. Zelo. 2003. Developing Information and Support Necessary to Prioritize and Support Removal of Abandoned
Vessels Impacting Coral Resources. In: Proceedings of the 13th Biennial Coastal Zone Conference.

Henderson, J.R. 2001. A Pre- and Post- MARPOL Annex V Summary of Hawaiian Monk Seal Entanglement and Marine De-
bris Accumulation in the Northwestern Hawaiian Islands, 1982-1998. Marine Pollution Bulletin 42 (7): 584-589.

Hoegh-Guldberg, O. 1999. Climate change, coral bleaching and the future of the world's coral reefs. Marine and Freshwater
Research 50: 839-866.

Houston, J.R. 1995. The Economic Value of Beaches. Coastal Engineering Research Center. Volume CERC-95-4.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

Hughes, TP. 1985. Life histories and population dynamics of early successional corals. pp. 101-106. In: Proceedings of the 5th
International Coral Reef Symposium.

Hughes, TP. 1994. Catastrophes, phase-shifts, and large-scale degradation of a Caribbean coral reef. Science 265: 1547-
1551

Hughes TP. and J.H. Connell. 1999. Multiple stressors on coral reefs: A long-term perspective. Limnology and Oceanography
44: 932-940.

Intergovernmental Panel on Climate Change. 2001. Climate Change 2001: The Scientific Basis, Contribution of Working
Group I to the Third Assessment Report of the Intergovernmental Panel on Climate Change (IPCC). J.T Houghton, Y Ding,
D.J. Griggs, M. Noguer, P.J. van der Linden and D. Xiaosu (eds.). Cambridge University Press. 944 pp.

International Petroleum Industry Environmental Conservation Association (IPIECA). 2000. Biological Impacts of Oil Pollution:
Coral Reefs. Available from the internet URL: http://www.ipieca.org.

IPIECA (International Pertoleum Industry Environmental Conservation Association). 1992. Biological Impacts of Oil Pollution:
Coral Reefs. International Pertoleum Industry Environmental Conservation Association. IPIECA Report Series, Volume 3.
London, England. 17 pp.

Jackson, J.B.C., M.X. Kirby, WH. Berger, K.A. Bjorndal, L.W Botsford, B.J. Bourque, R.H. Bradbury, R. Cooke, J. Erlandson,
J.A. Estes, TP. Hughes, S. Kidwell, C.B. Lange, H.S. Lenihan, J.M. Pandolfi, C.H. Peterson, R.S. Steneck, M.J. Tegner and
R.R. Warner. 2001. Historical overfishing and the recent collapse of coastal ecosystems. Science 293: 629-638.

Jennings, S. and N.V.C Polunin. 1996. Impacts of fishing on tropical reef ecosystems. Ambio 25: 44-49.

Jennings, S. and N.V.C. Polunin. 1997. Impacts of predator depletion by fishing on the biomass and diversity of non-target reef
fish communities. Coral Reefs 16: 71-82

Johannes, R.E. and M. Riepen. 1995. Environmental, economic and social implications of the live reef fish trade in Asia and
the Western Pacific. Report to the Nature Conservancy and the South Pacific Commission. 83 pp.

Johns, G.M., V.R. Leeworthy, F.W Bell and M.A. Bonn. 2003. Socioeconomic study of reefs in southeast Florida October 19,
2001 as revised April 18, 2003. National Oceanic and Atmospheric Administration, National Ocean Service. Silver Spring, MD.
255 pp.

Jokiel, P.L. 1985. Lunar periodicity of planulae release in the reef coral Pocillopora damicornis in relation to various environ-
mental factors. pp. (4): 289-293. In: Proceedings of the 5th International Coral Reef Symposium.

Kim, K. and C.D. Harvell. 2001. Aspergillosis of Sea Fan Corals: Disease Dynamics in the Florida Keys. pp. 813-824. In: Porter,
J.W and K.G. Porter [eds.]. The Everglades, Florida Bay, and Coral Reefs of the Florida Keys: An Ecosystem Sourcebook.
CRC Press, New York.
Kleypas, J.A., R.W Buddemeier, D. Archer, J.P. Gattuso, C. Langdon and B.N. Opdyke. 1999. Geochemical consequences of
increased atmospheric carbon dioxide on coral reefs. Science 284: 118-119.

Knowlton, N. 2001. The future of coral reefs. pp. 5419-5425 In: Proceedings of the National Academy of Science 98.

Knowlton, N., J.C. Lang and B.D. Keller. 1990. Case study of natural population collapse: post-hurricane predation on Jamai-
can staghorn corals. Smithsonian Contributions to the Marine Sciences 31: 1-25.

Kojis B.L. and N.J. Quinn. 1985. Puberty in Goniastrea favulus. Age or size limited? pp. (4): 289-283. In: Proceedings of the
5th Internatinal Coral Reef Symposium.

Kuta, K.G. and L.L. Richardson. 1996. Abundance and distribution of black band disease on coral reefs in the northern Florida
Keys. Coral Reefs. 15: 219-223.

Kuta, K.G. and L.L. Richardson. 2002. Ecological aspects of black band disease of corals: relationship between disease inci-
dence and environmental factors. Coral Reefs 21 (4): 393-398.

Larkin, S. 2003. The U.S. Wholesale Marine Ornamental Market: Trade, Landings, and Market Opinions. pp. 77-89. In: Marine
Ornamental Species: Collection, Culture and Conservation. Iowa State Press.

Larkin, S. and R. Degner. 2001. "The U.S. Wholesale Market for Marine Ornamentals." Aquarium Sciences and Conservation
3 (1/3): 13-24.

Lassig, B.R. 1983. The effects of cyclonic storms on coral reef fish assemblages. Environmental Biology of Fishes 9 (1): 55-
63.

Lau, P.P.F. and R. Parry-Jones. 1999. The Hong Kong trade in live reef fish for food. Traffic East Asia and the World Wildlife
Fund for Nature Hong Kong.

LaVoie, D.M., L.D. Smith and G.M. Ruiz. 1999. The potential for intracoastal transfer of nonindigenous species in the ballast
water of ships. Estuarine Coastal and Shelf Science 48: 551-564.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

Leeworthy, V.R. and P.C. Wiley. 1997. A Socioeconomic Analysis of the Recreation Activities of Monroe County Residents in
the Florida Keys/Key West. National Oceanic and AtmosphericAdministration, National Ocean Service, Silver Spring, MD. 49
pp.
Lesser, M.P. and S. Lewis. 1996. Action spectrum for the effects of UV radiation on photosynthesis in the hermatypic coral
Pocillopora damicornis. Marine Ecology Progress Series 134 (1-3): 171-177.

Lessios, H.A., D.R. Robertson and J.D. Cubit. 1984. Spread of Diadema mass mortality throughout the Caribbean. Science
226: 335-337.

Liu, G, A.E. Strong, W Skirving and L.F. Arzayus, 2004. Overview of NOAA Coral Reef Watch Program's Near-Real Time Glob-
al Satellite Coral Bleaching Monitoring Activities. In: Proceedings of the 10th International Coral Reef Symposium, Okinawa.

Lourey, M.L., D.A.J. Ryan and I.R. Miller. 2000. Rates of decline and recovery of coral cover on reefs impacted by, recovering
from and unaffected by crown-of thorns starfish Acanthaster planci: a regional perspective of the Great Barrier Reef. Marine
Ecology Progress Series 196:176-186.

Maniwavie, TJ. Rewald, J. Aitsi, TP. Wagner and P.L. Munday. 2001. Recovery of corals after volcanic eruptions in Papua New
Guinea. Coral Reefs 20 (1): 24.

Marshall, N, SA.H. Milledge and P.S. Afonso. 2001. Trade review. Stormy seas for marine invertebrates. Trade in sea cucum-
bers, seashells and lobsters in Kenya, Tanzania and Mozambique. TRAFFIC East/Southern Africa. Nairobi, Kenya 70 pp.

McClanahan, TR. 1995. A coral reef ecosystem-fisheries model: impacts of fishing intensity and catch selection on reef struc-
ture and processes. Ecological Modelling. 80: 1-19.

McClanahan, TR. 2000. Recovery of a coral reef keystone predator, Balistapus undulatus, in East African marine parks. Bio-
logical Conservation 94:191-198

McManus, J.W, L.A.B. Menez, K.N.K. Reyes, S.G. Vergara and M.C. Ablan. 2000. Coral reef fishing and coral-algal phase
shifts: implications for global reef status. ICES Journal of Marine Science, 57 (3): 572 -578.

Miller, S.L., G.B. McFall and A.W Hulbert, 1993. Guidelines and recommendations for coral reef restoration in the Florida Keys
National Marine Sanctuary. National Undersea Research Center. University of North Carolina at Wilmington. 38 pp.

Minerals Management Service. 2002. OCS Oil Spill Facts. Available from the internet URL: http://www.mms.gov/stats/
PDFs/2002_OilSpillFacts.pdf.

Minerals Management Service. 2004. E.O. 13089-The Flower Garden Banks. Available from the internet URL: http://www.
mms.gov/eppd/compliance/13089/banks.htm.

Moore, C. 2003. Trashed-Across the Pacific Ocean, plastics, plastics, everywhere. Natural History 112 (9): 46-51.

NRC (National Research Council). 1996. Stemming the Tide: Controlling Introductions of Nonindigenous Species by Ships'
Ballast Water. National Academy Press, Washington, DC.

Naughton, J. 1995. Inshore Fisheries and Fishery Habitat, Sea Turtles, and Marine Mammals. In: An Evaluation of the Near-
shore Coral Reef Resources of Kahoolawe, Hawaii. Chapter 4. National Marine Fisheries Service, NOAA. http://cramp.wcc.
hawaii.edu/Study_Sites/Kahoolawe/An_Evaluationof theNearshore_Coral_Reef_Resources ofKahoolawe/CHAP4.asp.

Negri, A.P., L.D. Smith, N.S. Webster and A.J. Heyward. 2002. Understanding ship-grounding impacts on a coral reef: potential
effects of anti-foulant paint contamination on coral recruitment. Marine Pollution Bulletin 44: 111-117.

Nemeth R.S. and J.S. Nowlis. 2001. Monitoring the effects of land development on the near-shore reef environment of St.
Thomas, USVI. Bulletin of Marine Science 69 (2): 759-775.

NMFS (National Marine Fisheries Service). 2004, Annual Report to Congress on the Status of U.S. Fisheries 2003. National
Oceanic and AtmosphericAdministration, NMFS. Silver Spring, MD. 24 pp.

NOAA (National Oceanic and Atmospheric Administration). 1997. NOAA Coral Reef Initiative. Silver Spring, MD. 12 pp.

NOAA (National Oceanic and Atmospheric Administration). 2002a. A National Coral Reef Action Strategy: Report to Congress
on implementation of the Coral Reef Conservation Act of 2002 and the National Action Plan to Conserve Coral Reefs in 2002-
2003. NOAA. Silver Spring, Maryland.

NOAA (National Oceanic and Atmospheric Administration). 2002b. Environmental Assessment: M/V Wellwood Grounding
Site Restoration. Florida Keys National Marine Sanctuary, Monroe County, Florida. NOAA, Marine Sanctuaries Division. Silver
Spring, MD. 10 pp.

NOAA (National Oceanic and Atmospheric Administration). 2004. Coral Reef Initiative. From the internet url: http://www.publi-
caffairs.noaa.gov/cri.pdf.




eely Associated States: 2005


NOAA CSC (National Oceanic and Atmospheric Administration, Coastal Services Center). 2004. Hawaii Balances Use ai
Sustainability of Hanuama Bay. NOAA, Coastal Services Center (CSC). Available from the internet URL: http://www.csc.noe
gov/techniques/recreation/hanauma.html.

NRC (National Research Council). 2001. NRC Committee on the Science of Climate Change, Climate Change Science:
Analysis of Some Key Questions. National Academy Press, Washington, D.C.

Ocean Conservancy. 2003. Offshore Oil and Gas Leasing, Exploration, and Development. Available from the internet UF
http://www.oceanconservancy.org.

Olsgard, F. and J.S. Gray. 1995. A comprehensive analysis of the effects of offshore oil and gas exploration and production i
the benthic communities of the Norwegian continental shelf. Marine Ecology Progress Series 122: 277-306.

Ormond, R.F.G., R. Bradbury, S. Bainbridge, K. Fabricus, J. Keesing, L. De Vantier, P. Medley, and A. Steven. 1991. Test
a model of regulation of Crown-of-Thorns starfish by fish predators. In: R.H. Bradbury (ed.) Acanthaster and the coral reef:
theoretical perspective. Springer-Verlag, Berlin.

Pandolfi, J.M., R.H Bradbury, E. Sala, TP. Hughes, K.A. Bjorndal, R.G. Cooke, D. McArdle, L. McClenachan, M.J.H. Newme
G. Paredes, R.R. Warner and J.B.C. Jackson. 2003. Global trajectories of the long-term decline of coral reef ecosystems. S
ence 301: 955-958.

Patin, S. 2004. Decommissioning, abandonment and removal of obsolete offshore installations. Available from the intern
URL: http://www.offshore-environment.com/abandonment.html.

Patterson, K.L., J.W. Porter, K.B. Ritchie, S.W. Poison, E. Mueller, E.C. Peters, D.L. Santavy and G.W Smith. 2002. The eti,
ogy of white pox, a lethal disease of the Caribbean elkhorn coral, Acropora palmata. pp. 99 (13): 8725-8730. In: Proceedin
of the National Academy of Sciences.

Pauly, D., V. Christensen, J. Dalsgaard, R. Froese and F. Torres Jr. 1998. Fishing down marine food webs. Science 279: 86
863.

Pearson, P.N. and M.R. Palmer, 2000, Atmospheric carbon dioxide concentrations over the past 60 million years. Nature 4C
695.

Perrine, D. 1989. Reef fish feedings: amusement or nuisance? Sea Frontiers 35 (5): 272-279.

Petit, J.R., J. Jouzel, D. Raynaud, N.I. Barkov, J.-M. Barnola, I. Basile, M. Benders, J. Chappellaz, M. Davis, G. Delayque, I
Delmotte, V.M. Kotlyakov, M. Legrand, VY. Lipenkov, C. Lorius, L. P6pin, C. Ritz, E. Saltzman and M. Stievenard. 1999. C
mate and atmospheric history of the past 420,000 years from the Vostok ice core, Antarctica. Nature 399: 429-436.

Pew Oceans Commission. 2003. America's living oceans: charting a course for sea change. Pew Oceans Commission, Arlin
ton, VA. Available from the Internet URL: http://www.pewoceans.org.

Pickett, S.TA. and P.S. White. 1985. Patch dynamics: a synthesis. pp. 371 384. In: S.TA. Pickett and P.S. White (eds.) TI
ecology of natural disturbance and patch dynamics. Academic Press, Orlando.

Pimental, D., R. Zuniga and D. Morrison. In press. Update on the environmental and economic costs associated with alie
invasive species in the United States. Ecological Economics.

Plathong, S., G.J. Inglis and M.E. Huber. 2000. Effects of Self-Guided Snorkeling Trails on Corals in a Tropical Marine Pal
Conservation Biology. 14 (6): 1821-1830.

Pomerance, R. 1999. Coral bleaching, coral mortality, and global climate change, U.S. Bureau of Oceans and Internatior
Environmental and Scientific Affairs. Available from the internet url: http://www.state.gov/www/global/globalissues/cor
reefs/990305_coralreefrpt.html.

Porter, J.W and J.I. Tougas. 2001. Reef ecosystems: threats to their biodiversity. Encyclopedia of Biodiversity 5: 73-95

Pratchett M.S. 2001. Influence of coral symbionts on feeding preferences of crown-of-thorns starfish. Marine Ecology Progre
Series 214: 111-119.

Precht, WF. 1998. The art and science of reef restoration. Geotimes. 43 (1): 16-20.

Precht, WF., R.B. Aronson and D.W Swanson. 2001. Improving Scientific Decision-Making In the Restoration of Ship-Groun
ing Sites on Coral Reefs. Bulletin of Marine Science. 69 (2): 1001-1012.

Richardson, L.L. 1998. Coral diseases: what is really known. Trends in Ecological Evolution 13: 438-443.

Richardson L.L, W.M. Goldberg, R.G. Carlton and J.C. Halas. 1998. Coral disease outbreak in the Florida Keys: Plague tyl
II. Revista de Biologica Tropical 46 (5): 187-198.

Richardson, L.L. and R.A. Aronson. 2002. Infectious diseases of reef corals. In: Proceedings of the 9th International Coral Re
Symposium.




The State of Coral Reef Ecosystems of the United States and Freely Associated States: 2005

Rinkevich, B. and Y Loya, 1979. Laboratory Experiments on the Effects of Crude Oil on the Red Sea Coral Sylophora pistillata.
Marine Pollution Bulletin 10: 328-330.

Roblee, M.B., TR. Barber, P.R. Carlson, M.J. Durako, J.W Fourqurean, L.K. Muehlstein, D. Porter, L.A. Yarboro, L.A. Zieman
and R.T. Zieman. 1991. Mass mortality of the tropical seagrass Thalassia testudinum in Florida Bay (USA). Marine Ecology
Progress Series 71: 297-299.

Rogers, C.S. 1990. Reponses of coral reefs and reef organisms to sedimentation. Marine Ecology Progress Series 62: 185-
202.

Rogers, C.S. 1993. Hurricanes and coral reefs: the intermediate disturbance hypothesis revised. Coral Reefs 12: 127-137.

Rogers, C.S., L.N. McLain and E. Zullo. 1988. Damage to coral reef in the Virgin Islands National Park and Biosphere Reserve
from recreational activities. Proceedings from the 6th International Coral Reef Symposium. 2: 405-410.
Rouphael, A.B. and G.J. Inglis. 2001. Take only photographs and leave only footprints: an experimental study of the impacts of
underwater photographers on coral reef dive sites. Biological Conservation 100: 281-287.

Rouphael A.B. and G.J. Inglis. 2002. Increased spatial and temporal variability in coral damage caused by recreational scuba
diving. Ecological Applications 12(2): 427-440.

Ruiz, G.M., P.W Fofonoff, J.T Carlton, M.J. Wonham and A.H. Hines. 2000. Invasion of coastal marine communities in North
America: apparent patterns, processes, and biases. Annual Review of Ecology and Systematics 31: 481-531.

Sadovy, YJ., TJ. Donaldson, TR. Graham, F. McGilvray, G.J. Muldoon, M.J. Phillips, M.A. Rimmer, A. Smith and B. Yeeting.
2004. While stocks last: The live reef food fish trade. Asian Development Bank, Manila. 147 pp.

Sale, P.F. 2002. The Science we need to develop more effective management. pp. 361-376. In: P.F. Sale (ed.) Coral Reef
Fishes: Dynamics and Diversity in a Complex Ecosystem. Academic Press, San Diego.

Sammarco, P.W 1982. Echinoid grazing as a structuring force in coral communities: whole reef manipulations. Journal of Ex-
perimental Marine Biology and Ecology 61: 31-55.

Sano, M. 2000. Stability of reef fish assemblages: Responses to coral recovery after catastrophic predation by Acanthaster
planci. Marine Ecology Progress Series 198: 121-130.

Santavy, D.L. and E.C. Peters. 1997. Microbial pests: Coral disease in the western Atlantic. pp. 1: 607-612. In: Proceedings of
the 8th International Coral Reef Symposium.

Santavy, D.L., E.C. Peters and C. Quirolo. 1999. Yellow-blotch disease outbreak on reefs of the San Bias Islands, Panama.
Coral Reefs 18 (1): 97.

Schultz, S. 1998. Passenger Ship May Have Destroyed Coral Reef off Mexico. Shipping International 1: 1-2.

Shaked, Y, A. Agnon, B. Lazar, S. Marco, U. Avner and M. Stein. 2004. Large earthquakes kill coral reefs at the north-west Gulf
of Aqaba. Terra Nova 16 (3): 133-138.

Shigenaka, G. 2001. Toxicity of oil to reef-building corals: a spill response perspective. National Oceanic and Atmospheric
Administration Technical Memorandum, National Ocean Service, Office of Research and Restoration 8. Seattle. 87 pp.

Shomura, R.S. 1987. Hawaii's Marine Fisheries Resources: Yesterday (1900) and Today (1986). NMFS Southwest Fisheries
Science CenterAdministrative Report H-87-21. NMFS, SFSC, St. Petersburg, FL. 15 pp.

Silva-lniguez, L. and D.W Fischer. 2003. Quantification and classification of marine litter on the municipal beach of Ensenada,
Baja California, Mexico. Marine Pollution Bulletin 46 (1): 132-138.

Smith, S.H. 1998. Cruise ships: A Serious Threat to Coral Reefs and Associated Organisms. Ocean and Shoreline Manage-
ment. (11): 231-248.

Smith, S.R., D.C. Hellin and S.A. McKenna. 1998. Patterns of Juvenile Coral Abundance, Mortality, and Recruitment at the
M/V Wellwood and M/V Elpis Grounding Sites and Their Comparison to Undisturbed Reefs in the Florida Keys. Final Report
to NOAA Sanctuary and Reserves Division and the National Undersea Research Program. University of North Carolina, Wilm-
ington. 42 pp.

Spaulding, M.D., C. Ravilious and E.P. Green. 2001. World Atlas of Coral Reefs. UNEP World Conservation Monitoring Centre.
University of California Press, Berkley. 424 pp.

Spurgeon, J.P.G. 1992. The Economic Valuation of Coral Reefs. Marine Pollution Bulletin 24 (11): 529-536.

Sultzman, C. 2002. A Professional Jury Report on the Biological Impacts of Submarine Fiber Optic Cables on Shallow Reefs
off Hollywood, Florida. Report to PEER.

Sutherland, K.P., J.W Porter and C. Torres.2004. Disease and immunity in Caribbean and Indo-Pacific zooxanthellate corals.
Marine Ecological Progress Series 266: 265-272.




eely Associated States: 2005


Szmant, A.M. 1997. Nutrient effects on coral reefs: a hypothesis on the importance of topographic and trophic complexity
reef nutrient dynamics. pp. (2): 1527-1532. In: Proceedings of the 8th International Coral Reef Symposium.

Tissot, B.N. and L.E. Hallacher. 1999. Impact of aquarium collectors on reef fishes in Kona, Hawaii. Final report. Departme
of Land and Natural Resources, State of Hawaii, Honolulu. 32 pp.

Turgeon, D.D., R.G. Asch, B.D. Causey, R.E. Dodge, W Jaap, K. Banks, J. Delaney, B.D. Keller, R. Speiler, C.A. Matos, J.
Garcia, E. Diaz, D. Catanzaro, C.S. Rogers, Z. Hillis-Starr, R. Nemeth, M. Taylor, G.P. Schmahl, M.W. Miller, D.A. Gulko, J.
Maragos, A.M. Friedlander, C.L. Hunter, R.S. Brainard, P. Craig, R.H. Richond, G. Davis, J. Starmer, M. Trianni, P. Houk, C.
Birkeland, A. Edward, Y Golbuu, J. Gutierrez, N. Idechong, G. Paulay, A. Tafileichig and N. Vander Velde. 2002. The State
Coral Reef Ecosystems of the United States and Pacific Freely Associated States: 2002. National Oceanic and Atmosphe
Administration/National Ocean Service/National Centers for Coastal Ocean Science, Silver Spring, MD. 265 pp.

UNEP (United Nations Environmental Programme). 2004. Available from the internet URL: http://www.uneptie.org/pc/tourisl
sensitive/coral-threats.htm.

UNEP (United Nations Environmental Programme). 2002. Industry as a partner for sustainable development: tourism. Earl
print: England. 76 pp.

U.S. Census Data. 1990. http://www.census.gov/main/ww/cen1990.html.

U.S. Census Data. 2000. http://www.census.gov/main/ww/cen2000.html.

U.S. Commission on Ocean Policy. 2004. Preliminary Report of the U.S. Commission on Ocean Policy Governor's Draft.

USCRTF (United States Coral Reef Task Force). 1999. Coastal Uses Working Group Summary Report. November 1999. U.
Coral Reef Task Force. Washington, D.C. 170 pp.

USCRTF (United States Coral Reef Task Force). 2000. The National Action Plan to Conserve Coral Reefs. Washington, D.
34 pp.

U.S. Department of Defense. 2000. Coral Reef Protection Implementation Plan. Washington, D.C. 61 pp. plus appendices
Available from the internet URL: https://www.denix.osd.mil/denix/Public/ES-Programs/Conservation/Legacy/Coral-Reef/Ple
coralreef.html.

Wabnitz, C, M. Taylor, E. Green and T Razak. 2003. From Ocean to Aquarium. UNEP-WCMC, Cambribge, U.K. 64 pp.

Wells, S.M. 1989. Impacts of the precious shell harvest and trade: conservation of rare or fragile resources. pp. 443-454.
J.F. Caddy (ed.) Marine Invertebrate Fisheries: their assessment and management. John Wiley and Sons, Inc., New York.

Westmacott, S., K. Teleki, S. Wells and J. West. 2000. Management of bleached and severely damaged coral reefs. IUC
Gland, Switzerland and Cambridge, UK. vii + 36 pp.

Whitfield, P.E., T Gardner, S.P. Vives, M.R. Gilligan, W.R. Courtenay, G.C. Ray and J.A. Hare. 2002. Biological invasion oftl
Indo-Pacific lionfish (Pterois volitans) along the Atlantic coast of North America. Marine Ecology Progress Series 235: 28
297.

Wilcove, D.S., D. Rothstein, J. Dubow and A. Phillips and E. Losos. 1998. Quantifying threats to imperiled species in the Unite
States. BioScience 48: 607-615.

Wilkinson, C. (ed.). 1998. Status of Coral Reefs of the World: 1998. Australian Institute of Marine Science, Townsville, Austral
Available from the internet URL: http://www.aims.gov.au/scr1998.

Wilkinson, Clive (ed.). 2002. Status of Coral Reefs of the World: 2002. Australian Institute of Marine Science, Townsville, AL
tralia. 357 pp. Available from the internet URL: http://www.reefbase.org/pdf/GCRMN/GCRMN2000.pdf.

Wilson, P. and D.P. Wheeler. 1997. California's Ocean Resources: An Agenda for the Future. The Resources Agency of Cc
fornia.

Wood, E.M. 2001. Collection of coral reef fish for aquaria: Global trade, conservation issues and management strategies. M
rine Conservation Society, Ross-on-Wye, UK.

Woodley CM,A.W Bruckner, S.B. Galloway, S.M. McLaughlin, C.A. Downs, J.E. Fauth, E.B. Shotts and K.L. Lidie. 2003. Col
Disease and Health: A National Research Plan. National Oceanic and AtmosphericAdministration, Silver Spring, MD. 72 pr

Zobrist, E.C. 1998. Coral Reef Restoration and Protection from Vessel Groundings. In: GERS Meeting Abstracts.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


The State of Coral Reef Ecosystems of the U.S. Virgin Islands

Christopher F.G. Jeffrey1, Ursula Anlauf', James Beets3, Sheri Caseau4, William Coles2, Alan M. Friedlander1 5, Steve
Herzlieb6, Zandy Hillis-Starr7, Matthew Kendall1, Violeta Mayor2, Jeffrey Miller4, Richard Nemeth6, Caroline Rogers8,
Wesley Toller2

INTRODUCTION AND SETTING

Coral reef ecosystems in the U.S. Virgin Islands (USVI) consist of a mosaic of habitats, namely coral and other
hardbottom areas, seagrasses, and mangroves that house a large diversity of organisms. These biologically
rich ecosystems provide important ecosystem services (e.g., shoreline protection) and support valuable so-
cio-economic activities (e.g., fishing and tourism), but they are also affected directly and indirectly by these
activities. This chapter presents an assessment of the current status of coral reef ecosystems in the USVI. It
provides a comprehensive review of historic and current literature and long-term datasets that describe coral
reef ecosystems of the territory. It also provides data synthesized from current monitoring programs con-
ducted by Federal and territorial organizations.

The USVI comprises three large main islands and several smaller islands (Figure 4.1). St. Croix the largest
island is 207 km2 in size. St. Thomas is the second largest island at 83 km2, and St. John is the third largest
at 52 km2. The geologically dissimilar islands lie between two major island archipelagos: the older Greater
Antilles to the west and the younger Lesser Antilles to the east. St. Thomas and St. John are more similar to
the Lesser Antilles than to Puerto Rico with which they share an extensive shallow water platform (Adey et al.,
1977). St. Croix geologically belongs to the Greater Antilles but is isolated by the Virgin Islands Trough that is
over 4,000 m deep (NOAA National Geophysical Data Center, http://www.ngdc.noaa.gov/mgg/gdas/gd_
sys.html, accessed: 11/2/2004). Managed areas in coastal waters of the three main islands exist to protect,
maintain, or restore natural and cultural resources (Figure 4.1).

Reefs in St. Thomas and St. John generally form fringing, patch, or spur and groove formations that are distrib-
uted patchily around the islands (see Figure 4.20). The eastern and southern shores of St. Croix are protected
by well-developed barrier reef systems with near-emergent reef crests that separate lagoons from off-shore
bank areas (Adey, 1975; Hubbard et al., 1993). Bank reefs and scattered patch reefs occur on geological
features at greater depths offshore. Recently, the National Oceanic and Atmospheric Administration (NOAA)
mapped 485 km2 of benthic habitats in the USVI to a nominal depth of 30 m. Analyses of these maps revealed
that coral reef and hard-bottom habitats comprise 300 km2 (61%), submerged aquatic vegetation covers 161
km2 (33%), and unconsolidated sediments comprise 24 km2 (4%) of shallow water areas (Kendall et al., 2001;
Monaco, 2001; http://biogeo.nos.noaa.gov, accessed 1/19/05).













1 NOAA National Ocean Service, Center for Coastal Monitoring and Assessment, Silver Spring, MD
2 Virgin Islands Department of Planning and Natural Resources, St. Croix, USVI.
3 University of Hawaii, HI
4 National Park Service, St. John, USVI
5 Oceanic Institute, HI
6 University of the Virgin Islands Center for Marine Environmental Studies, St. .Thomas, USVI.
7 National Park Service, St. Croix, USVI
8 US Geological Survey, St. John, USVI.










































Locations Cited in Text
1. Caret Bay
2. Black Point, Brewr's Bay
3. Fat Cay
4. Red Hind Bank
5. Grammanlk Bank
6. Buck land
7. Spnrt Bay
8. Seahorse Cottage Shoal
9. Buner Bay
10. Great Cruz Bay
11. Fish Bay
12 Cocoloba Cay
13. Yawst Point
14. Lameahur BSay
IL. Tktite Ref
16. Coral Bay
17. Mennebeck
18. Haulover Bay
19. Newfound Bay
20. Sprat Hole (off Sprat Ha Beach)
21. Mutton Snapper Reef
22. USVI Rum Factory
23. USVI Rum Factory outfall
24. St. Croix Internatonal Airport
2. Cane Bay
28. Govt Sewage Outfall
27. Hovensa il Refinery
28. Salt River
29. Long Reot


30. LBJ Pump Station Outfall
31. Long ReefEagle Ray
32. Alton Lagoon
33. Great Pond
34. Great Pond Bay
35. Solitude Bay
36. Castle
37. Buck Island
3. GQerson
39. Jacksllsaac Bay
40. Lang Bank
SCity

- V.I Natlona Park
V. I. Coral Reef
National Monument
Buck Island Reef
National Monument
East End Marine Park

- Land


Water <20m

Deep Water


Figure 4.1. A map of the USVI showing managed areas, municipalities, and other locations mentioned in this chapter. Map: A. Sha-
piro.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


ENVIRONMENTAL AND ANTHROPOGENIC STRESSORS

Coral reefs in the USVI face similar pressures as reefs elsewhere in the Caribbean (Rogers and Beets, 2001).
Of the 13 major coral reef stressors identified by the U.S. Coral Reef Task Force, 10 have been identified
as being problematic to reef ecosystems in the territory. These stressors include climate change; diseases;
tropical storms; coastal development and runoff; coastal pollution; tourism and recreation; fishing; and ships,
boats, and groundings. The impacts of these stressors on USVI coral reefs are summarized in this chapter.
Other stressors such as alien species, security activities, and offshore oil activities are not relevant to the
USVI. Stressors are described fully in Chapter 3 of this report.


Climate Change and Coral Bleaching


Climate change refers to the trend of
increasing mean global air tempera-
ture and sea surface temperatures
(SST) within the last century com-
pared with previous estimates. This
warming trend is generally attributed
to the atmospheric accumulation of
greenhouse gases. Bleaching in the
USVI has been reported since 1987
(Figure 4.2). Bleaching was most
severe and had the highest reported
incidence of occurrence during the
Caribbean-wide event of 1998-1999.
According to the U.S. National Park
Service (NPS), the 1998 bleaching
event coincided with the highest re-
corded SSTs in the USVI. Bleach-
ing was less severe in 1999 probably
because water temperatures were
slightly lower (28.80C) during that
year. The 1999 bleaching event did
not result in extensive coral colony
mortality because most colonies re-
covered within six months of being
bleached (Nemeth and Sladek-Now-
lis, 2001; Nemeth et al., 2003c). For
both years, bleaching was most se-
vere in St. Croix, followed by St. John,
and then St. Thomas (Figure 4.2).


Diseases
Several diseases have affected coral
community structure and have de-
graded coral cover (Table 4.1). Be-
tween 1976 and 1989, white band
disease (WBD), bleaching, and hur-
ricanes reduced the cover of elkhorn
coral (Acropora palmata) by as much
as 85% within the Virgin Islands Na-
tional Park (VINP) and the Buck Is-


Date

60
1999
50-

S40-

t 30-
0
20

10-

0
St Croix St John St Thomas
Location
Figure 4.2. Annual trends in coral bleaching in the USVI. Upper panel shows the
number of bleaching reports by year and severity. Arrows indicate the Caribbean-
wide bleaching event of 1998-1999. Source: Reefbase 2003, http://www.reefbase.
org, Accessed: 10/23/2003. Lower panel shows the estimated percent of coral tissues
that bleached in 1998-1999. Bars represent the maximum percent of sampled coral
colonies that bleached by island and year. Source: Rogers and Miller, 2001; Nemeth
et al., 2003c.


land Reef National Monument (BIRNM; Gladfelter et al., 1977; Rogers et al., 1982; Edmunds and Witman,
1991; Bythell et al., 1992; Rogers and Beets, 2001). Between December 1997 and May 2001, 14 species of


5


4

a




*1




o
0


Unknown
-1 Low
Iln High








Table 4.1. Diseases affecting coral reef organisms in the U.S. Caribbean and Florida. Source: Bruckner, 2001.






Irregular lesions. i of anrou-S sizes Fungu.s 10- (IVVell Common sea fan IJnkno'.'n
distributed throughout the sea fan blade i4 .pe,-tilus and Smith IiGog' r-ia entaina\
due to loss of tissue and skeleton Tissue si do 2 00 .I ''enus sea fan IG
Cs irriounding the lesion often becomes flabellumLs and
dark purple and maV ha..e nodules both other branching
of ".hich occur in response to a ariety gorgonians including
of stressors Identification of this disease Fseudobecta i na s ppI
requires confirmation of the presence of
h..hitte fungal filaments
Crescent shaped or circular band of black rCvanobacteria 20 Se -eral soft corals 1 20
filamentousi material separating II ng Sulfide- and 20 hard corals
colored Coral tiss.Ue from white e posed o0-dizng incingi ng boulder star
Coral skeleton bacteria Corals Alhonasili aea
Sulfate- anm 'iaIs couple, i
reducing and s iimmetrical
bacteria brain coral IC'plc'a


black band disease in that it forms a cactus corals
stincti"e band that separates ii.e coral IAll celoph lla spp
sue from bare a..hite skeleton blushing star coral.
the common seafan
i'.iona renlahna l
ring daylight the filaments spread out C anobacteria 6 siQosa C ralans unkno ,n
e a net in a diffuse fashion o er II e Al ani1lula.s Al
sue and bare skeleton at night the band ca eiosa Poties
mns a compact balled-up mat at the astredces and
erface bet', een Ii e tissue and e posed Siideashea iadijans
eleton
)ral tissue peels or sloughs off from coral I_lnkno. n .Elkhorn and staghorn .
eleton in a uniform band from the base corals i 4clopora spp
the colon., up yardss second form
JE1--II i e, hibits a transient zone bet ,-een
'parently healthy tissue and e posed
.-.I.-..-.it th m t .-.-. ^ .-.f 1.I-. -..- h...t




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Table 4.1 (con't.). Diseases affecting coral reef organisms in the U.S. Caribbean and Florida. Source: Bruckner, 2001.





infected colonies. marcescens palmata)
Pale, circular blotches of translucent tissue Unknown 3 Boulder star corals < 1
or a narrow band of pale tissue at the (Montastraea spp.)
colony margin surrounded by normal, fully and the brain coral
pigmented tissue. Infected tissue dies, and (Colpophyllia natans)
exposed skeleton is colonized by algae.


hard coral in the VINP were infected
60 Annualmean with the white plague type II, a newly

S--- SEofmean identified disease (Miller et al., 2003;
So Weil and Smith, 2003). Miller et al.
(2003) observed a new incidence of
S40 -- -- white plague type II every month, al-

1 though the monthly frequency of in-
30 ._ __ fections decreased during the study
S-- (Figure 4.3). A disease-causing fun-
S20- ------ gus, Aspergillis sydowii, has been
-- --- isolated from air samples taken dur-
s 10- ing African dust storms and has been
infecting sea fans on reefs in the
- USVI (Garrison et al., 2003).
DJFMAMJJAODFMAJJASNDJFMMJASONDJFMAM
1997 1998 g199 2000 2001
Sampling Period

Figure 4.3. Mean monthly and annual frequency of disease occurrence on Tektite
Reef, St. John, over 42 months. Source: Miller et al., 2003.


Tropical Storms
Tropical storms are a major force structuring coral reef communities in the Caribbean. Storms have the capac-
ity to degrade reefs in several ways. They increase terrestrial runoff, sedimentation, and pollution affecting
coral reefs, and cause extensive physical damage to the substratum. Several hurricanes have affected USVI
reefs since 1979, but Hurricanes David (1979) and Hugo (1989) were the most severe and destructive (Figure
4.4). The eye of David a category five hurricane traveled about 160 km southwest of St. Croix; the eye
of Hugo a category four hurricane passed directly over the island (Figure 4.4). Damage to reefs varied
with storm path, strength and velocity, wave height and direction, the dominant coral species, and reef depth
(Rogers et al., 1997; Bythell et al., 2000). The strongest evidence of storm damage to reefs was observed
at Lameshur Bay, St. John, and Buck Island, St. Croix. Hurrincane David resulted in large stands of elkhorn
coral on reef crests being replaced by mounds of dead elkhorn coral rubble at both Lameshur Bay and Buck
Island (Rogers et al., 1982; Beets et al., 1986). In Lameshur Bay, Hurricane Hugo caused significant declines
in total live coral cover, including star coral (Montastrea annularis), a dominant and slow growing coral species
(Edmunds, 1991; Rogers et al., 1991). At Buck Island, Hurricane Hugo resulted in significant declines in cover
of M. annularis and Porites porites at depths of 8-10 m, although M. annularis suffered greater mortality from
predation and tissue necrosis over a two-year period than from physical damage from the hurricane (Bythell et
al., 1993; Bythell et al., 2000). Hurricane Hugo also reduced areas on the south side of Buck Island to rubble
pavement and moved the reef crest off the island's south side 30 m landward (Hubbard, 1991).




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Fifteen years after Hurricane Hugo,
reefs in Lameshur Bay still have not
shown significant increases in live
coral cover (Rogers et al., 1997; C.
Rogers, pers. obs.; J. Miller, pers.
obs.). Exposure by Hugo of new
substrates for colonization, coupled
with a reduction in abundance of
urchins and herbivorous fishes that
consume macroalgae, may have fa-
cilitated extensive growth of macroal-
gae (Lessios et al., 1984; Levitan,
1988; Rogers et al., 1997). Macroal-
gae inhibit settlement and survival of
coral recruits and growth by existing
colonies, and mean benthic cover of
macroalgae sometimes reaches over
30% in the affected areas (Lessios et
al., 1984; Levitan, 1988; Rogers et
al., 1997). At Buck Island, recovery
Hurricanes(1979-2W4) Of elkhorn coral damaged by Hur-
cae sgory ricane David was hindered by WBD
-2 and by Hurricane Marilyn in 1995
-,"- WMIrc.2 m . . . (Rogers et al., 1982; Rogers et al.,
go."4 G w.." .. 2002). Some recruitment of elkhorn
Deep W*er
... coral has occurred since the 1995
..' l' . = .hurricane (Bythell et al., 2000; Rog-
S ":. w ers et al., 2002).
Figure 4.4. Hurrcanes that affected the coral reef ecosystems in the USVI between
1979 and 2001. Storm names are followed by year of occurrence and storm category
on the Saffir-Simpson hurricane scale. H1 to H5 = Hurricane categories one through
five. Arrows indicate the direction and path of the storms. Map: A. Shapiro. Source:
NOAA Coastal Services Center.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands

Coastal Development and Runoff
Sedimentation associated with runoff from coastal development poses a serious threat to water quality in the
USVI. In St. Thomas and St. John, the problem is worsened by the steep terrain of the islands (80% of the
slopes exceed 300 in incline), and runoff from unpaved roads after intense rain showers is considered the larg-
est contributor of eroded sediments to coastal waters (CH2M Hill Inc., 1979; Anderson and MacDonald, 1998;
IRF, 1999). Although published data on the temporal increase and spatial extent of coastal development in
the USVI are scarce, unplanned and poorly regulated development for a growing population, and a booming
tourism industry may have taken a toll on coral reef ecosystems through the years (see Tourism and Recre-
ation section). Nemeth and Sladek-Nowlis (2001) monitored the impacts of a hotel development on a nearby
fringing coral reef at Caret Bay, St. Thomas (Figure 4.5) monthly for two years. The hotel construction site was
on a steep hillside less than 50 m from the shoreline. The landward edge of the reef was 75-140 m from the
shoreline. Rates of sedimentation, changes in water quality, and changes in the abundance and diversity of
corals and other reef organisms were measured along five permanent transects from July 1997 to March 1999.
Sediment loads and suspended sol-
ids were highest at ravine outlets and
sheltered locations, increased during
large rainfall events, and decreased
after buildings and road pavements
were completed. Live coral cover
along the entire reef tract declined
about 14% and was lowest at sites
with the highest rates of sedimenta-
tion (Figure 4.6).

Severe rainfall events are problemat-
ic and can overwhelm existing sewer
and stormwater systems. During No-
vember 2003, a low pressure system
dropped 38 cm of rain in five days
throughout the territory and contrib-
uted to the formation of large sedi-
ment plumes along developed areas Figure 4.5. A large mound of dirt was excavated from a construction site located less
Sth coastline. ed ent p mes than 100 m from the shoreline at Solitude Bay, St. Croix. Photo: C. Jeffrey.
of the coastline. Sediment plumes
resulted in a decline in water qual- 0
ity, elevated turbidity on nearby reefs
and seagrass beds, and forced the -5
closure of swimming beaches. Ad-
ditionally, wastewater disposal and -10
sewage systems frequently malfunc-
tion and discharge raw sewage into .~ 15
nearshore areas. Despite the many
environmental problems associated & -20
with coastal development, major de- ,
velopment projects adjacent to en- 0 -25
vironmentally sensitive habitats are
being proposed and welcomed by -30
government officials as boosters of
the economy. Such projects may ex- -35
acerbate existing problems of coastal 7 8 9 10 11 12
pollution and runoff, and ultimately Sedimentation Rate (mg per em2 per day)
may harm the islands' economy that Figure 4.6. Percent change in live coral cover as a function of increasing rate of
is at least partly dependent on the sedimentation at Caret Bay, St. Thomas. Data were collected from five permanent
health of their natural resources. transects at Caret Bay Reef between July 1997 and March 1999 before, during, and
after the construction of the Caret Bay Villas. The decline in live coral was significant
(p <0.01). Source: Nemeth and Sladek-Nowlis, 2001.


U




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Coastal Pollution
Coastal pollution has led to several days of beach closings in the USVI. In 1999, the USVI was ranked third
in the number of beach closings per year among U.S. States and Territories with 307 beach-closing days (Na-
tional Resources Defense Council, http://www2.nrdc.org/water/oceans/ttw/sumvi.pdf, Accessed: 11/10/2004).
Beach closings decreased to eight days in 2002, but increased tenfold to 80 days in 2003. The continued de-
cline in coastal water quality has been linked to coastal development and runoff, as well as point and nonpoint
source discharges (USVI DPNR, 2004). In St. Croix, the Virgin Islands rum manufacturing plant discharges
a plume of wastewater that is visible from the discharge point to about 10 km westward along the shoreline.
Biological pollution of coastal water results largely from a failing, overloaded municipal sewage system that
frequently empties sewage directly into nearshore waters as well as the discharge of vessel wastes directly
into the sea by boat owners. Coastal waters also become polluted when groundwater that has been contami-
nated by failing septic tanks, sewage infiltration, and petroleum is carried to the marine environment during
flooding after intense rainfall (USVI DPNR, 2004). These pollution problems are worsened by a lack of public
awareness about the importance of USVI waters, which further contributes to the degradation of marine water
quality.

The USVI Department of Planning and Natural Resources (DPNR) conducts a Water Pollution Control Pro-
gram to monitor all known point source discharges of pollution such as outfalls, harbors, marinas, and main
recreational areas (USVI DPNR, 2004). The program also evaluates coastal water quality by monitoring
nonpoint source discharges through a signed Memoranda of Agreement and Cooperation with several partner
agencies, including the Virgin Islands Resource Conservation and Development Council, VI Conservation
District, University of the Virgin Islands (UVI), U.S. Geological Survey (USGS), Island Resources Foundation,
the NPS, and the St. Croix Environmental Association (SEA). Additionally, the NPS in St. John and UVI pub-
lish a local newsletter to inform and educate the public on nonpoint source pollution problems (USVI DPNR,
2004).


Tourism and Recreation
Historically dependent on agriculture and trade, the USVI has developed a robust tourism industry during the
last 34 years, which has shifted the islands' economy to one that is mainly tourism-based. The number of tour-
ist arrivals to St. Thomas and St. John has quadrupled between 1970 and 2000; tourist arrivals to St. Croix re-
mained relatively unchanged during the same period (Figure 4.7). In 2000, 108,612 USVI residents were joined
by 2.2 million visitors, but the annual number of tourists has remained fairly constant since then (U.S. Census
Bureau, 2003, http://www.census.gov/prod/cen2000/island/Vlprofile.pdf, Accessed 3/1/05; USVI Bureau of
Economic Research, http://www.us-
viber.org, Accessed 11/7/2004). 2.5
SSt. Croix
., St Thomas / St. John
Tourism accounts for more than 70% ThomasSt.
of the gross domestic product of the 2.0
territory, and as in other tourism-de-
pendent countries, the environmental -2
costs of tourism are evident. Visible >s
impacts of increased visitation in- "
clude physical damage to habitats,- 1,o0
poor treatment and control of solid =
waste and sewage, increased eutro-
phication, groundwater depletion and 05s
contamination, increased sediment
loads, and displacement of traditional
resource use (IRF, 1996; Bryant et 0.0
1970 1980 1990 2000
al., 1998; Burke and Maidens, 2004).
Snorkeling and diving are major rec- Date
rational activities that could cause Figure 4.7. Number of visitors to St. Croix and St. Thomas/St. John between 1970
physical damage to reefs. For exam and 2000. Source: USVI Bureau of Economic Research, http://www.usviber.org, ac-
physical damage toa cessed 11/7/04.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


pie, physical damage to corals has been observed at the BIRNM underwater snorkel trail, which attracts up to
200 visitors per day (Z. Hillis-Starr, pers. obs.). However, the more obvious impact of increased tourism has
been the exacerbation of solid waste disposal problems caused by the high density of tourists and residents
as well as an economy heavily dependent on high energy-consumption.

Tourists, residents, and poorly regulated development also contribute directly and indirectly to coastal pol-
lution. For example, participation by residents and tourists in diverse marine recreational activities such as
boating, fishing, diving, snorkeling, kayaking and beach camping negatively impact the marine environment in
overpopulated areas. Most reported oil spills in the USVI stem from the refueling of yachts, ferries, and cruise
ships (IRF, 1996). Poor lawn care practices on golf courses, in residential areas, and at tourist resorts are con-
sidered major sources of nitrate and phosphate contamination to nearshore areas through stormwater runoff
(IRF, 1996). Finally, the development of major tourism facilities in coastal areas further threatens the coastal
environment through increased sediment loads from the construction of buildings and roads, the operations of
facilities, and stormwater runoff (IRF, 1996).

More thought and effort is now being given to promoting sustainable tourism. Dive and anchor buoys have
been installed at popular dive sites to reduce the incidence of anchor damage. Environmental education and
outreach is on the agenda of the national and local non-governmental organizations (NGOs), as well as ter-
ritorial and Federal government agencies. For example, the NPS, UVI, SEA, and The Nature Conservancy
are providing eco-hikes and other educational tours and programs for the public.


Fishing
Fishery resources have declined in the USVI since the 1960s (e.g., Appeldoorn et al., 1992). Although fish-
ing is a visible and obvious impact to fisheries species in coral reef ecosystems, less tractable environmental
threats such as habitat degradation or loss and marine pollution have also undoubtedly contributed to the de-
cline in fisheries. Fishing has a long history in the USVI (Fiedler and Jarvis, 1932). Strongly integrated into the
Virgin Islands culture, fishing provides subsistence, supplemental income, recreation, or full-time employment
to the islanders. Residents and tourists consume a wide variety of marine species in relatively large quantities
(i.e., about 15 kg/person/year; Swingle et al., 1979; Olsen et al., 1984). Resource managers divide the USVI
fisheries into commercial and recreational fishing sectors. Presently, the bulk of information on USVI fisheries
is derived from the commercial sector.

The USVI commercial fishery is artisanal in nature and consists of about 380 registered fishers 240 on St.
Croix and 140 on St. Thomas and St. John (Brownell, 1971; Brownell and Rainey, 1971; Tobias et al., 2000;
CFMC, 2003; Gordon and Uwate, 2003; Kojis, 2004). Typically, fishers use small, open vessels (6 to 8 m in
length) powered by outboard motors to fish with a variety of gear types and methods, although traps or fish
pots are the most popular gear type (Sylvester and Dammann, 1972; Appeldoorn et al., 1992; Beets, 1997;
Kojis, 2004). Scuba diving is also a common commercial method used to harvest reef fishes (by spear) and
invertebrates (by hand or with snare; CFMC, 2003). In the past decade, gillnets and trammel nets used with
the aid of scuba equipment have become common fishing gear on St. Croix, and annual landing from nets
now exceed annual landings from traps on this island (Tobias and Toller, 2004). USVI commercial fishers must
submit monthly catch records as a stipulation for permit renewal.

Recreational fishing is also very important to the USVI economy. Boat-based recreational fishing may have
contributed as much as $5.9 million in fishing-related expenditures to the local economy in 2000 (UVI EEC,
2002), up from about $90,000 in 1986 (Jennings, 1992). Snappers were the most preferred species-group of
fishers. Collectively, however, much of the reported fishing effort was directed towards pelagic fish species
such as blue marlin, sailfish, dolphinfish, tuna, wahoo, and kingfish (UVI EEC, 2002). The recreational fishery
for pelagics has been routinely monitored for over a decade but is not the subject of this review (see Adams,
1995; Mateo, 2000). Less data exist on the impact of recreational fishing to reef-associated species primarily
because recreational fishers do not obtain fishing permits, and no records are kept on their population size or
activity (Appeldoorn et al., 1992).




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Reef Fish Fishery
Reef fish assemblages and the composition of reef fish landings have changed markedly in the USVI over the
past 40 years (Appeldoorn et al., 1992; Rogers and Beets, 2001). Catanzaro et al. (2002) reviewed the lam-
entable collapse of the USVI fishery for Nassau groupers during the 1970s (Olsen and LaPlace, 1978) and red
hind during the 1980s (Beets and Friedlander, 1992). Landings of larger individuals of snappers (Lutjanidae)
and other groupers (e.g., the coney, Epinephelus fulvus) also declined in the 1980s (Appeldoorn et al., 1992).
Although fishery-dependent monitoring data for the 1990s are still unavailable, a growing number of fishery-in-
dependent studies, primarily utilizing visual survey methods, indicate that populations of targeted grouper and
snapper species have not recovered to date (Rogers and Beets, 2001; Beets and Friedlander, 2003; CCMA-
BT, unpublished data; Nemeth et al., 2004). On average, fewer than eight groupers per year were observed
during monitoring of reef fish assemblages at four reference sites within the VINP between 1989 and 2000
(Figure 4.8). Nassau grouper were observed in only 3% of 1,764 visual surveys conducted at the four ref-
erence sites during the entire study
period (Beets and Friedlander, 2003). 10- Hugo Marilyn
Of the 2,292 snappers and groupers Haulover
SNewfound
observed during 756 visual fish sur- wA
veys within the VINP and the BIRNM, 8- Teke
S1Large hurricanes
less than 2% were greater than 35 g Samping method a
cm in length (CCMA-BT, unpublished
data). On nearshore reefs throughout
the USVI, Nemeth et al. (2003c) also 6
found a similar trend where snappers 4
and groupers larger than 40 cm were E
mostly absent from the fish assem-
blage. Densities of snappers and 2-
groupers averaged five and three
fish/100 m2, respectively, with the
most common size class of both fam- 0
1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000
ilies being 11-20 cm (Nemeth et al.,
2003c). In contrast, the herbivorous Date
fishes (e.g., Acanthuridae and Scari- Figure 4.8. Abundance trends in groupers (Serranidae) among the four reference
dae) dominated the fish assemblage sites around St. John, U.S. Virgin Islands, from 1989-2000. Source: Beets and Fried-
lander, 2003.
with average densities between 10-
20 fish/100 m2.

Lobster Fishery
The Caribbean spiny lobster, Panulirus argus, is a species of tremendous commercial and recreational impor-
tance in the USVI. Spiny lobsters accounted for 6% of total reported landings in 1998-1999 (Tobias, 2000) and
its commercial value probably exceeds that of any other single reef-associated species in the USVI. Although
Bohnsack et al. (1991) concluded that USVI lobster populations appeared healthy, more recent studies found
a 10% decrease in mean size between 1997 and 2000, which suggests that overfishing is occurring (Tobias,
2000; Mateo and Tobias, 2002). The Virgin Islands Division of Fish and Wildlife (DFW) routinely monitors
commercial lobster landings (weight and carapace length) and has periodically monitored lobster recruitment
around St. Thomas, where recruitment appears to be highly variable but generally low (Gordon and Vasques,
in press). Limited field surveys around St. John also indicate that average lobster size and density have de-
creased since 1970 (Wolf, 1998).

Conch Fishery
The queen conch, Strombus gigas, forms an important fishery in the USVI. During the 1990s, conch accounted
for about 2% of total USVI landings, with most conch landed on St. Croix (5% of total St. Croix landings) and
less landed on St. Thomas and St. John (0.4% of total landings on St. Thomas and St. John; Valle-Esquivel,
2002). The value of reported USVI commercial conch landings in 1998-99 was about $340,000 (Tobias et
al., 2000). In the USVI, commercial fishers harvest queen conch by hand, primarily by scuba diving, although
some conch are also harvested by free diving (Rosario, 1995). Very little information exists presently on the
recreational harvest of conch, but concerns about declining stocks have been voiced for over 20 years (Wood




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


and Olsen, 1984; Valle-Esquivel, 2002). Although numerous territorial regulations were enacted in 1988 to
protect conch stocks including a five-year closure of the fishery on St. Thomas and St. John, conch stocks
have either not shown significant recovery or have continued to decline in the USVI (Friedlander, 1997; Gor-
don, 2002).

It is difficult to separate out the causal factors of fishery decline in the USVI. Overfishing, technological ad-
vances in fishery gear (larger boats, more powerful engines, and improved gear), and the deterioration of
habitats may have contributed to significant changes in the community structure of reef fish assemblages and
the observed decline in fishery yields. Several studies have documented the failure of existing regulations and
a lack of enforcement in protecting reef fishes or reversing the declines in the abundance of preferred species
such as the large groupers and snappers (Beets,1996; Wolff, 1996; Garrison et al., 1998; Rogers and Beets,
2001). Likewise, other studies have identified sedimentation and pollution as major factors in the decline of
nearshore reef ecosystems, which may have contributed to the decline of fisheries species (deGraaf and
Moore, 1987; Rogers and Beets, 2001).

On a more positive note, some USVI fisheries are beginning to show small signs of recovery. Since the clo-
sure of an important red hind spawning aggregation site south of St. Thomas in 1990, the average size of red
hind from the St. Thomas fishery increased significantly from 26 cm to over 34 cm total length in 2003 (Ne-
meth, in review). Moreover, tag and release studies conducted by the UVI on a red hind spawning aggregation
near a shelf-edge reef south of St. Thomas found that 78% of the fish were over 35 cm, 50% over 37.5 cm,
and fish greater than 40 cm (Nemeth, unpublished data). In March 2004, scientists at UVI discovered the first
evidence of a Nassau grouper spawning aggregation reestablishing itself south of St. Thomas. Underwater
surveys estimated that up to 100 Nassau groupers have aggregated and showed signs of reproductive activity
(Nemeth, in review). Unfortunately local fishers have targeted this unprotected reef for the past several years
and fishing mortality may seriously threaten this Nassau spawning aggregation. In this same area, yellowfin
grouper, tiger grouper, and cubera snapper have all been seen forming large spawning aggregations (Nemeth,
in review).


Trade in Coral and Live Reef Species
The trade in coral and live species is a minor problem in the USVI and has not received as much attention as
it has in other U.S. jurisdictions with coral reef ecosystems. The trade in live coral and fishes is prohibited by
the USVI Endangered and Indigenous Species Act of 1990 (Title 12, Chapter 2), the purpose of which is "to
protect, conserve, and manage indigenous fish, wildlife and plants, and endangered or threatened species for
the ultimate benefit of all Virgin Islanders, now and in the future." Very few permits have been issued for the
harvest or take of live coral and non-commercial or recreational fishes. Issued permits have been for research
and education purposes only. Thus, there are very few exports of live coral (W Coles, pers. obs.). Locally,
U.S. Customs and Department of Agriculture officials have confiscated small amounts of live coral that had
been recently collected by tourists departing from the territory. A much greater problem is the export of dead
coral (some of which may have been collected alive) and other marine organisms. Considerable amounts
of dead and dried coral, undersized conch, and shells have been confiscated at airports on the islands of St.
Thomas and St. Croix and at the U.S. Postal Service inspection facility in Puerto Rico.


Ships, Boats, and Groundings
Data on the impacts of resource use within the territory are limited, and research on this topic is needed. A
recent assessment of marine resource utilization identified boating as a major recreational activity in the USVI,
and damage from small boats anchored in corals and seagrasses as a primary concern (Link, 1997). A total
of 2,462 private and commercial boats were registered throughout the territory in 2000 (Uwate et al., 2001).
To reduce anchor damage to reefs, the VINP has installed over 300 mooring and protection buoys around
St. John and at the BIRNM. Another local program initiative called "Anchors Away" has recently installed 50
mooring buoys around the island of St. Croix. Additionally, funding from the NOAA Coral Reef Conserva-
tion Program has been approved for additional moorings within the East End Marine Park in St. Croix. Boat
groundings are also of concern. In 1988, a cruise ship destroyed 283 m2 of reef within the VINP, and coral
recovery after 10 years has been minimal (Rogers and Garrison, 2001).




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Marine Debris
Marine debris has become a problem in the USVI. Debris that washes out to sea via runoff (sewers, street
litter) pollutes the water and shorelines and can be life-threatening to marine organisms and humans. Fishing
line and nets, rope and other trash can wrap around animals and cause drowning, infection, or amputation, or
can settle on hard bottom areas and kill coral colonies. A major landfill that exists along the coast near Sandy
Point, St. Croix receives most of the solid waste from the island. Debris from this landfill consisting primar-
ily of fishing lines, bottles, plastic bags and other street litter is often washed out to sea where it becomes a
health hazard to marine life. The same may be true for landfills on St. Thomas and St. John. The SEA has
organized several beach cleanup campaigns to increase public awareness about marine debris and reduce
the amount of debris that litters the nearshore environment, but this threat is an ongoing concern.


Aquatic Invasive Species
Aquatic invasive species are not recognized as a major threat in this jurisdiction.


Security Training Activities
No security training activities currently occur in this jurisdiction.


Offshore Oil and Gas Exploration
No offshore oil and gas exploration currently occurs in the USVI.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands

CORAL REEF ECOSYSTEMS-DATA-GATHERING ACTIVITIES AND RESOURCE CONDITION

This section focuses on resource monitoring activities, data collection and analyses, and summaries of pub-
lished studies and data sets to provide an assessment of the current condition of resources in coral reef eco-
systems of the USVI. Information is presented to describe three functional or structural components of coral
reef ecosystems: marine water quality, benthic habitats, and coral reef-associated fauna (Table 4.2). A brief
summary of ongoing research and monitoring programs, methods, results and discussion, and an assessment
of overall condition are presented for each ecosystem component. Locations of monitoring and research ef-
forts are shown in Figure 4.9.

Table 4.2. Data sets selected for the descriDtion of the current condition and status of coral reef ecosystems in the USVI.





The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Figure 4.9. Locations of monitoring and research efforts in the USVI between 1988-2004. The boundary of the Buck Island Reef
National Monument was expanded in 2001 from 880 acres to 19,015 acres. Both the original and expanded boundaries are shown.
Water quality monitoring is conducted by the USVI DPNR. Coral monitoring is conducted by NOS, NPS, USGS, USVI DPNR, and
UVI-CMES. Map: A. Shapiro.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


WATER QUALITY

USVI DPNR/DEP Water Quality Monitoring

Methods
Water sampling in the USVI was initiated by the local health department in 1968. A network of fixed monitor-
ing stations was selected within the bays and nearshore waters of the islands to target areas of particular
concern, such as outfalls, harbors, marinas, and main recreational areas. The Division of Environmental Pro-
tection (DEP) within the USVI DPNR samples 135 sites quarterly each year (53 around St. Croix, 64 around
St. Thomas, and 18 around St. John). At each monitoring site, water samples are collected at the surface
to measure and record the chemical and physical parameters listed in Table 4.3. All data are uploaded into
STORET, a national online database maintained by the U.S. Environmental Protection Agency (EPA, http://
www.epa.gov/storet/dw_home.html, Accessed: 12/28/2004). Quarterly assessments are also complemented
by periodic assessments of water quality during episodic events (e.g., a sewage bypass), when USVI DPNR-
DEP collects daily samples until acceptable levels of water quality are reestablished.

Table 4.3. Water quality parameters measured by the USVI DPNR-DEP, NPS, and the U.S. Geological Survey.


In si&lu nSi multi parameter meter surface and near boitom
inl/L Ia L/ y S SI multi parameter meter surface and near bottom
Parts per thousand (ppti Ii u lL- Si multi parameter meter surface and near bottom
Scale of 1-141 n S1 u I SI multn parameter meter surface and near bottom
iTII In sILu S1 miulti parameter meter surface and near bottom
r i eters In eu erage depth of Sechi disk id sappearancie/appearancel
mingL ,srab near surface and send to certified lab
number of colonies/ 10 ml Grab near surface and send [o a certified lab
mg'L ,rab near surface and send to a certified lab



Results and Discussion
Data from the USVI DPNR-DEP water quality monitoring program indicate that water quality in the USVI is
generally good but declining because of an increase in point and nonpoint source pollution (S. Caseau, pers.
obs.). Almost all direct discharges in the USVI were traced to a failing and overloaded municipal sewage sys-
tem. Moreover, sewage treatment plants malfunction as the result of human error, old equipment, or unusual
conditions in the raw sewage.

Flooding is a major concern in the Virgin Islands. Watersheds have small areas, steep slopes, and increasing
amounts of impervious surfaces, which in turn can result in high-volume runoff after short periods of intense
rainfall. The territorial system consists of combined sewers, which are pipes designed to carry both raw sew-
age and stormwater. When the volume of rain becomes too great, the sewer system becomes overloaded,
and untreated sewage discharge flows into nearby marine waters (Figure 4.10A). In non-urban and suburban
areas, rainwater often flows directly over farms, golf courses, and lawns, washing pathogenic animal waste,
fertilizers, and pesticides into the water. Failure to use effective silt-control devices during construction ac-
tivities and improper discharge of waste by boat owners can result in pathogens that pollute beaches in less
densely populated areas (Figures 4.10B, C).

The Virgin Islands rum manufacturing process generates wastewater that is discharged on the south coast
of St. Croix. The effluent typically forms a plume visible from the discharge point to about 10 km westward
along the shoreline. As a result, a strong turbidity and color gradient decreases light penetration, which could
impede normal growth of submerged aquatic vegetation and corals. This effluent may be a reason for the
absence of significant coral reefs within direct influence of the discharge.




The State of Coral Reef Ecosystems of the U.S. Virc


Figure 4.10. Pollution of marine waters in the USVI. Left panel: Flooding of a sewer system in St. Croix after an intense rainfall.
Center panel: Poor land management practices associated with accelerating coastal development in St. John. Right panel: Improper
discharge of gasoline in marine waters from boating activities in St. Thomas. Source: V. Mayor, USVI DPNR-DEP

NPS and USGS Water Quality Monitoring

Methods
The NPS and USGS conduct assessments of water quality within the VINP in St. John. Monitoring of water
quality within and outside the park began in 1988. Thirty-one sites were originally identified for monitoring but
were reduced to 15 in 1996. Samples are taken every three months at each site for the parameters listed in
Table 4.3. Data through 2000 are available on-line at the EPA STORET website. Data from 2000 through the
present are being processed and analyzed for uploading to STORET (http://www.epa.gov/storet/dw_home.
html, Accessed: 12/28/2004). In June 2000, monthly sampling for Enterococcus spp. and fecal coliform began
at three park swimming beaches.

Results and Discussion
Data collected by the NPS and USGS from 1988-1998 indicate that marine water quality in the VINP is excel-
lent except at Cruz Bay. Horizontal visibility ranged from 10-20 m. Mean water temperature at 1m depth was
27.90C with a range of 20.2-31.90C. At a depth of 10 m, temperature varied from 24.5C to 30.80C. Average
salinity was 35 parts per thousand and average conductivity was 54 siemens. Marine systems in the region
experience little variation in these parameters. The extinction coefficient of photosynthetically active radiation
(PAR) was approximately 0.18, which is extremely good. Dissolved oxygen over dense seagrass beds was
7-8 mg/L. Dissolved oxygen over barren, muddy substrates (e.g., Cruz Bay) averaged 6.0 mg/L and ranged
between 5.0-7.1 mg/L.

Turbidity, a measure of particles in the water column, has been increasing. Additionally, turbidity was consis-
tently higher outside the park than in waters inside the park, which may have resulted from sediment erosion
caused by development of land outside the VINP. Total suspended solids (TSS) ranged from 1-15 mg/L ad-
jacent to heavily disturbed watersheds after large rainfall events. Likewise, nutrient analyses resulted in de-
tectable levels of micronutrients around mangroves, probably resulting from the natural production of organic
nutrients and in bays adjacent to the most developed watersheds such as Coral Bay, Cruz Bay, and Great
Cruz Bay. Clean, clear water is critical to maintaining healthy coral communities and seagrass beds.


UVI-CMES and USVI DPNR-DEP Coral Bay Study




of Coral Reef Ecosystems of the U.S. Virgin Islands


Sediment coring and data on TSS were used to provide a rapid assessment of the state of Coral Bay. Sam-
pling site locations were chosen based on: 1) proximity to expected inputs of terrigenous sediment and 2)
achieving adequate spatial coverage throughout the bay. Sites were concentrated within Coral Harbor, the
area most expected to be impacted by the recent increase in development. Sites outside of the harbor were
chosen as control sites or sites at which to detect point sources of input from recent development. The as-
sessment of water quality conditions in Coral Bay, Coral Harbor, and at other sites around St. John was based
on a review of TSS data from the NPS water quality monitoring program at the VINP. Detailed descriptions of
the sampling protocol are provided in Devine et al., 2003.

Results and Discussion
Devine et al. (2003) found poor water quality in Coral Bay. Vibracores and surface sediment samples indi-
cated a seven-fold increase in the sedimentation rate and terrigenous input into Coral Bay as a direct result
of development within the watershed during the last 100 years, and more probably over the past 40 years.
Analyses of sediment samples also suggest that within the last 10-15 years, sedimentation rates in Coral Bay
were 1) 10-20 times greater than the rate of natural sediment deposition averaged over the last 5,000 years
and 2) the plantation era had a very small impact on sediment deposition. TSS was four times higher in Coral
Harbor than the average for all other sampled locations in St. John. Mapping of sediment deposition and data
on water chemistry indicate a growing problem within the harbor and the adjacent Coral Reef National Monu-
ment.

Coral Bay, an Area of Particular Concern (APC), is one of the largest watershed drainage areas within the
territory at 1,215 ha (Hubbard et al., 1987; Devine et al., 2003). The area also has the highest recorded rate
of population growth (79%) in the USVI between 1990 and 2000 (U.S. Census Bureau, 2001). The Coral
Bay Watershed has steep slopes that average 180 (several greater than 350), highly erodable soils, and very
diverse land use along the shoreline (Devine et al., 2003).

Coral Bay has a protected inner harbor with critically valuable fringing mangroves and salt ponds. The area
is also home to a cruising and live-aboard population of between 75-150 boats at any particular time. Many
boats (i.e., 15-20) are permanently anchored to the small mangrove fringe. No pump-out facilities exist to
handle vessel septic waste, no inspections are made of these vessels' holding tanks, and no regulations are
enforced to protect marine resources. Along the inner harbor, commercial businesses and an undeveloped
marina operate without containment for liquid spills or solid waste, paint, and dust. Residential roads, new
homes, and failing septic systems contribute unmeasured amounts of pollutants to the harbor.

The tremendous growth and diverse uses of the landscape and marine resources in this watershed have
visibly deteriorated marine water quality. Sedimentation and runoff are increasing in intensity and frequency,
with routine sediment plumes inundating the area during the rainy season. Chronic pollution from point and
nonpoint sources including the dumping of human and animal wastes, failing septic systems, and dumping of
boat tank materials has resulted in high nutrient levels in Coral Bay.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


BENTHIC HABITATS

NPS and USGS Coral Disease and Benthic Cover Abundance Monitoring

Methods
Long-term monitoring of coral diseases and abundance (percent cover) is conducted by the Inventory and
Monitoring Program around St. John and Buck Island in St. Croix. Diseases of corals are specifically moni-
tored using two different methods at two sites in VINP. The incidence and progression of the coral disease
white plague type II in 28 tagged coral colonies is being monitoring approximately quarterly using still photog-
raphy. Coral diseases are also monitored monthly with 1 m2 quadrats along eight 10-m transects. Details of
these monitoring projects, which began in 1997 and are still on-going, are given in Miller et al. (2003).

Benthic cover monitoring is conducted annually through the use of digital videography at four representative
sites around St. John US Virgin Islands (three within VINP, one outside of the park); and at two sites within
BIRNM in St. Croix. Monitoring of benthic cover began at two sites in 1999; four additional sites have been
added since 2000 (J. Miller, pers. obs.). The benthic sampling protocol involves the selection of random (tran-
sect origin) sites, which is accomplished by using a SONAR-based mapping system (AquaMap). Twenty
10-m transects are filmed using a digital video camera, and then the images are downloaded to a computer.
Random dots are placed upon images captured from transects. The substrate underneath each dot is identi-
fied to the lowest taxonomic unit possible (e.g., coral to species, algae to genus) and entered into a database.
Queries of the database produce values on the percent cover, diversity indices of species, and cover groups.
Qualitative data on coral disease are also collected. A detailed description of the protocol is available online
(http://science.nature.nps.gov/im/monitor/protocoldb.cfm, Accessed: 12/28/2004).

Results and Discussion
Live coral cover along disease monitoring transects decreased from 65.3% ( 7.41 standard deviation [SD]) to
43.4% ( 5.08 SD) between December 1997 and May 2001. The frequency of disease within transects ranged
from 3% to 58%, and the area of disease patches ranged from 0.25 to 9,000 cm2 within that same period. New
incidences of disease were observed every month with associated loss of living coral. Increases in disease
occurrence did not correlate with elevated water temperatures. The photos and observations revealed no re-
covery of diseased corals with all necrotic tissue being overgrown rapidly by turf algae, usually within less than
one month. Most coral colonies suffered partial mortality and some colonies greater than 1.5 m in diameter
were completely consumed in less than six months. Some limited recruitment (e.g., Porities spp., Agaricia
spp., Favia spp., and sponges) has been noted on the diseased areas.

In general, reefs monitored by the NPS and USGS were dominated by dead coral with turf algae (Figure 4.11).
Other benthic organisms such as gorgonians and sponges were not abundant and showed no significant tem-
poral patterns in percent cover. In contrast, the estimates of the percent cover of macroalgae, turf algae, and
abiotic substrates (sand, rubble, and pavement) varied substantially among sites and among sample periods.
A total of 19 coral species were recorded throughout the study period; the number of species varied among
sites and years. At Newfound Reef in St. John, the Montastraea annularis complex was the most abundant
and most frequently observed coral, accounting for approximately 70% of live coral cover and was present in
all 20 transects. The percent cover of live coral and algae was variable among sites, with Mennebeck Reef in
St. John having the highest estimates of live coral cover (Figure 4.11). Mennebeck Reef and Western Spur
and Groove Reef, St. Croix had more live coral than macroalgae, but live coral was twice as abundant on
Mennebeck as on Western Spur and Groove (Figure 4.11). At Yawzi Reef, the opposite pattern occurred, with
macroalgae being more abundant than live coral for all years (Figure 4.11). At South Fore Reef, St. Croix and
Newfound Reef, mean estimates of live coral cover were similar to those for macroalgae (Figure 4.11).

Significant changes in live coral cover occurred only at Newfound Reef in St. John, where the mean percent
live cover decreased by approximately 24% between 1999 and 2001 (p<0.01, Figure 4.11). Porites porites
was the only coral species to increase in both mean live coral cover and frequency at Yawzi Reef. At both
Yawzi and Mennebeck Reefs, Porites coral was consistently observed in more than 50% of belt-transects.
Haulover Reef in St. John had a high abundance of live coral (22.1%) based on one year's worth of data (Fig-
ure 4.11). Fifteen species were recorded at Haulover. The Montastraea annularis complex comprised 84% of
the live coral cover and occurred in all transects at Haulover Reef.





The State of Coral Reef Ecosystems of the U.S.


Virgin Islands


100
S. Fore Reef, St. Croix USVI
80

60
6Coral

40- Macroalgae
Dead coral wI turf algae
20

0
100
W. Spur and Groove, St. Croix USVI
80

60

40

20

0
100
Haulover Reef, St. John USVI
80

60

40

20

0
O 100
O Mennebeck Reef, St. John USVI
80

60

40

20

0
100
YNe wzi Reef, St. John USVI
80












p 20

0
100 1999 2000 2001 2002 2003
Newfoundland Reef, St. John USVI
80

60

40

20

0
1999 2000 2001 2002 2003

Year


Figure 4.11. Mean percent cover ( SE) of coral, macroalgae, and dead coral with turf algae at six sites in the USVI. Data were col-
lected according to video monitoring protocols developed by the USGS and NPS (http://science.nature.nps.gov/im/monitor/protocoldb.
cfm). Specific sites or years without data were not sampled or the data are not yet analyzed. Source: J. Miller, unpublished data.




The State of Coral Reef Ecosystems of the U.S. Virc


NPS, USGS, and UVI-CMES Elkhorn Coral (Acropora palmata) Monitoring

Methods
Researchers from the USGS, NPS, and UVI-CMES began an 18-month study of 66 tagged elkhorn coral colo-
nies at Haulover Bay, St. John in January 2003. The geographic coordinates of the perimeter of each monitor-
ing site and the locations of sampled elkhorn colonies are mapped onto geo-referenced aerial photographs.
Data are recorded on the depth, three-dimensional size of colonies, type of substrate, percent cover of live
and dead coral, presence/absence of specific diseases and lesions, and counts of damselfish territories and
predators (snails Coralliophila abbreviata and C. caribaea; and fireworms Hermodice spp.).

Results and Discussion
Observations of 66 tagged corals over an 18-month period showed that 17% died, 74% had disease, and
30% suffered physical breakage, most likely from careless snorkelers (Table 4.4). Although 92% showed new
growth throughout the study period, Table 4.4. The results of a study on the health and condition of Acropora palmata
15% of the new growth later lost 90- colonies (N=67). Data were collected through videography and in situ observation
100% of their tissue. White pox dis- by the USGS Caribbean field station since February 2003 at Haulover Bay, St. John,
USVI. Source: Rogers and Muller, unpublished data.
ease was the most significant cause
of coral mortality, however white pox
lesions can heal. Forty-eight percent
of the white pox lesions healed com- 40 ,.1 ,.1
pletely, mostly within three months. i.
The onset of a severe disease out-
break coincided with increasing sea IC
surface temperatures. Both the num-
ber of corals with white pox and the
total number of disease lesions start-
ed to rise in September and contin-
ued increasing into November, track-
ing the trend in SST. o
1 1 1




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


UVI-CMES Video Assessment of Benthic Substrates

Methods
UVI-CMES researchers used digital videography along belt-transects to characterize and monitor benthic
cover at permanent and rapid assessment sites in St. Croix and St. Thomas. The maximum width and height
and the percent of diseased coral cover were estimated from the videos for all coral colonies greater than 10
cm in diameter that were located directly under the transect lines. Data on diseases and bleaching were not
collected at rapid assessment sites. In St. Croix, divers filmed three to six permanent 10-m transects at 10
long-term and two rapid assessment sites between April 2001 and March 2003. In St. Thomas, digital video
transects were conducted at six coral reef sites between August 2002 and September 2003. The St. Thomas
reefs were placed into three categories based upon their location along the insular platform: nearshore reefs
(5-30 m deep, <2 km from the shoreline); mid-shelf reefs (5-30 m deep, 2-10 km offshore; and shelf-edge reefs
(>30 m deep, 10-50 km offshore). Detailed video sampling methods are discussed in Nemeth et al. (2002).
Results are reported separately here for St. Croix and St. Thomas.

Results and Discussion


St. Croix
The percent cover of living coral was
variable among sites and ranged
from 4.4% to 39.1%. Coral was the
most abundant component at only
one site. Turf algae covering dead
coral comprised 50% or more of the
substrata and was dominant at most
sites (Figure 4.12A, B). Dead coral
included both long dead and recently
killed coral covered with a layer of turf
algae. The percent cover of macroal-
gae ranged from 3.2% to 34.9% (Fig-
ure 4.12C). Percent cover of living
coral was similar among years but
a significant increase in turf algae
and a corresponding decrease in
macroalgae were observed at Buck
Island between years (Figure 4.12
B, C). These trends were reversed
at Sprat Hole, where an increase in
macroalgae corresponded with a de-
crease in turf algae (Figure 4.12B, C).
Significant increases in macroalgae
also occurred at Long Reef (Figure
4.12C). It is unlikely that the signifi-
cant changes observed in the abun-
dance of macroalgae between years
at Buck Island and Sprat Hole were
caused by urchins (D. antillarum) be-
cause macroalgae were very rare at
those sites. Sponges and gorgoni-
ans each comprised less than 10% of
the benthic cover at all sites (Figure
4.13D, E). Sand was the only non-


BI CB CS GS GP JB LB LR MS SRE SRW SH


100
80
60

a 40
20
0

100
80
60



20
0

100

80
60
* 40
20
0


A. Scleractinian Coral


BI CB CS GS GP JB LB LR MS SRE SRW SH


C. Macroalgae


BI CB CS GS GP JB LB LR MS SRE SRW SH
Sampling Site
Figure 4.12. Annual mean percent of (A) scleractinian corals, (B) dead coral with
turf algae, and (C) macroalgae for 12 sampling sites in St. Croix. Site codes are:
Bl=Buck Island, CB=Cane Bay, CS=Castle, GS=Gerson, GP=Great Pond, JB=Jacks
Bay, LB=Lang Bank, LR=Long Reef/Eagle Ray, MS=Mutton Snapper, SRE=Salt
River East Wall, SRW=Salt River West Wall, SH=Sprat Hole. Error bars represent
standard deviation. Asterisks denote significant differences: *=p<0.05; **=p<0.01;
***=p<0.0001. Source: Nemeth et al., 2002, 2003a.


living substrate type found at the sites, ranging from 0.0% to 9.5% (Figure 4.13F). No significant changes
occurred for sponges, gorgonians, or sand cover between years at any site.







The composition of the coral com- 18
munity was similar among sites, with 16a 2001 A. Sponges
14 ] 2002
10 species representing 95% of the 14
S12
coral community (Figure 4.14). Mon- lo
tastraea spp. were the most dominant o 8
corals except at Castle where Porites 6
porites was most abundant, at Great 4
Pond where Porites astreoides was o 0 0o oI
most abundant, and at Gerson and l8 c8 cs GS GP JB LB LR MS SRE SRW SH
Lang Bank where Diploria strigosa 18Gorgonians
was most abundant. Coral diver- 1
14
sity ranged from a Shannon-Weaver 12
diversity index (H') of 1.50 at Great 10-
Pond to 2.40 at Salt River West Wall
(Figure 4.15). 4
2- Die s0 a 0 0 0b
Coral condition varied greatly among 0 C J LB LR MS SRE SRW SH
sites. The incidence of coral disease
and bleaching ranged from 0-17% 1 C. Sand/Sediment
at several sites and 0-22% at Lang 14
Bank. Diseases and bleaching were S 12 -
observed among eight dominant cor- 10
al species, with Siderastrea siderea e
having the highest incidence of dis- 4
ease (50% of colonies) and bleaching 2 o o00 00 a o o 00
(80% of sampled colonies). Divers 81 C8 CS GS GP JB LB LR MS SRE SRW SH
observed white plague, dark spots Sampling Site
disease, yellow blotch/band disease,
disease, yellow blotch/band disease, Figure 4.13. Annual mean percent of (A) sponges, (B) gorgonians, and (C) sand/se
and white spots that were classified ment for 12 sampling sites in St. Croix. Site codes are: BI=Buck Island, CB=Cane B
as disease, but were unidentifiable to CS=Castle, GS=Gerson, GP=Great Pond, JB=Jacks Bay, LB=Lang Bank, LR=Lo
a specific disease. Reef/Eagle Ray, MS=Mutton Snapper, SRE=Salt River East Wall, SRW=Salt Ri\
West Wall, SH=Sprat Hole. Error bars represent standard deviation. Asterisks c
note significant differences: *=p<0.05; **=p<0.01; ***=p<0.0001. Source: Nemeth
al., 2002, 2003a.


DL CN MILC
MME 1% 1% 1%
MILA 2% Other
2%
DS
MACX
PP 42%
6%
SS AA Agaci agarites
7% CN Colpophy#ia natns
DL #plria labyinUtifomes
DS Diptona sfigo
AA /I MME Meanddna meandries
7% L MISeporna aicomis
MILC Milepora alcicomis complaneta
PA / MACX Montastrea annuteris complex
MC Montastma cavemose
9% PA PotLs asleroks
PP Poates pories
MC / SS Sklerestaa sdese
20 Other All other species combined

Figure 4.14. Percentage coral species composition at all sampled sites in St. Cro
'Other' denotes percent of all other coral species combined: Stephanocoenia michE
nii, Eusmilia fastigiata, D. clivosa, Madracid decactis, M. mirabilis, Mussa angulo,
Mycetophyllia danaana, M. ferox, M. aliciae, Dichocoenia stokesii, Manicina are
loata. and P divaricata. Source: Nemeth et al.. 2002. 2003a.







St. Thomas
The percent cover of living coral
ranged from a low of 8.3% at Benner
Bay to a high of 42% at Grammanik
Bank (Figure 4.16A). The percent
cover of dead coral covered with turf
algae ranged from 15% at Seahorse
Cottage Shoal to 45.6% at Benner
Bay (Figure 4.16B). The percent
cover of macroalgae ranged from
13.8% at Black Point to 42.7% at Sea-
horse Cottage Shoal (Figure 4.16C).
Sponges and gorgonians each com-
prised less than 10% of the benthic
cover at all sites (Figure 4.17D, E).
No gorgonians were observed on the
shelf-edge sites. The percent cover
of sand/sediment ranged from 3% at
the Grammanik Bank to 28% at Black
Point (Figure 4.17F). Almost all of the
substrate at Black Point was covered
by sediment, whereas the substrates
at other sites were predominantly
sandy areas mixed with vertical reef
structures.


The State of Coral Reef Ecosystems of the U.S. Virgin Islands


3.0
2001
2002
2.5


2.0


1.5


1.0


0.5


0.0
BI CB CS GS GP JB LB LR MS SRE SRW SH
Sampling Site

Figure 4.15. Annual diversity index (Shannon-Weaver H') for corals at 12 sites in
St. Croix. BI=Buck Island, CB=Cane Bay, CS=Castle, GS=Gerson, GP=Great Pond,
JB=Jacks Bay, LB=Lang Bank, LR=Long Reef/Eagle Ray, MS=Mutton Snapper,
SRE=Salt River East Wall, SRW=Salt River West Wall, SH=Sprat Hole. Source: Ne-
meth et al., 2002, 2003a.


60
50
40
S30
20
10
a

60
50
L 40
S3
20
10
0

60
50
a 40
| 30
20
10
0


8B BP SC FC G8 RN


BB BP SC FC GB RH
Nearshore Mid-shelf Shelf-edge


BB BP


SC FC GB8RH


Figure 4.16. Mean percent cover of benthic organisms in St. Thomas at: BB=Benner
Bay, BP=Black Point, SC=Seahorse Cottage Shoal, FC=Flat Cay, GB=Grammanik
Bank, RH=Red Hind Bank. BB and BP are nearshore sites; SC and FC are mid-shelf
sites; GB and RH are shelf-edge sites. n=6 transects for all sites. Error bars represent
standard deviation. Source: S. Herzlieb, unpublished data.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Nearshore sites tended to have low-
er percent cover of living coral and 40
higher percent cover of dead coral i Nearshoe A. Sponges
i Mid-sheff
covered with turf algae than mid- 30a She-edge
shelf and shelf-edge sites (Figure 9
4.18). Also, nearshore sites tended 8 20
to have lower percent composition of 10-
corals within the M. annularis com-
plex and higher percent composition o-
of the stress tolerant corals P astre- BB BP SC FC GB RH
oides and S. siderea than mid-shelf 40B. Gorgonians
and shelf-edge sites (Figure 4.18).
The coral reefs of St. Thomas were
generally dominated by coral species 20-
in the genus Montastraea (Figure a
4.18). The Shannon-Weaver Diver- 10
sity Index (H') for coral ranged from No Da
a high of 2.26 at Flat Cay to a low of BB BP SC FC GB RH
1.20 at Grammanik Bank. In general, 40
deeper shelf-edge sites (Seahorse C. Sand/Sediment
Cottage, Grammanik Bank, and Red 30 -
Hind Bank) had lower diversity than
the shallow sites (Figure 4.19). 20

Since most research and monitoring T
in the Virgin Islands in have gener- o ,
ally been concentrated on nearshore BB BP SC FC GB RH
fringing reefs, mid-shelf and shelf- Nearshore Mid-shelf Shelf-edge
edge sites were chosen to fill gaps in
the knowledge of other reef systems, Figure 4.17. Mean percent cover of A. sponges, B. gorgonians, and C. sand/sedi-
as well as to establish an experimen- ment in St. Thomas: BB=Benner Bay, BP=Black Point, SC=Seahorse Cottage Shoal,
tal design to test hypotheses involv- FC=Flat Cay, GB=Grammanik Bank, RH=Red Hind Bank. BB and BP are nearshore
sites, SC and FC are mid-shelf sites, and GB and RH are shelf-edge sites. n=6 tran-
ing differences between reefs located sects for all sites. Error bars represent standard deviation. Source: S. Herzlieb,
at different points along the insular unpublished data.
platform off the coast of St. Thomas.
The close proximity of nearshore fringing reefs to human populations and their relatively shallow depths, in-
creases the susceptibility of these reefs to both harmful human activities (overfishing, sedimentation, nutrient
enrichment, and physical damage) and the effects of natural disturbances (storm wave damage, high SSTs,
and high irradiance).

Due to their similar depths but greater distance from shore, mid-shelf reefs are less susceptible to the human-
induced stresses listed above, but are exposed to levels of natural impacts similar to nearshore reefs. Thus,
the mid-shelf reefs provide an ideal control for measuring the effects of human-induced stresses on nearshore
reefs. Deep reefs located along the edge of the insular platform are largely free from human induced stresses
(excluding fishing and anchoring) and natural impacts because of their greater distance from human popula-
tions and their greater depths. The shelf-edge deep reefs are quite extensive, but largely unstudied. Monitor-
ing of these systems will contribute greatly to an understanding of coral reef resources in the Virgin Islands.
Cross-shelf patterns in benthic composition in St. Thomas warrant special attention because they suggest that
overall reef quality is lower at nearshore sites compared with sites further offshore. However, only two reefs
of each reef type were surveyed, thus robust comparisons between reef types are difficult. Future monitoring
efforts involving a greater number of St. Thomas reefs will help to elucidate these and any further differences
among the near-shore, mid-shelf, and shelf-edge reef systems.











A. Nearshore B. Mid-shelf

PF SR Other MME PF Other
DS 1. 1.4% 4.3% AA 1.4% 1.4% 3.0%

2.3%5 2.2%
MILA MACX PP
2.9% 33.6% 2.3%
AA 2.3%
6.4% AC
2.3% MA AC Acropora cervicomis
4.1% 61.0% AA Agaricia agaricites
MC MC AL Agaricia lamarcki
10.1% pp 18.0% CN Colpophyllia natans
12.5% DS Diploria strigosa
SS MME Meandnina meandrites
10.7% PA
12.8% MILA Millepora alcicornis
MACX Montastrea annularis complex
MC Montastrea cavernosa
C. Shelf-edge D. All sites combined PA Porites asteroides
PF Porites furcata
AL CN Other AC PF AL MILA PP Porites porites
SS 1.3% 1.1? 1.8% MME 0.9% 8% .7% 0.7% Other SR Siderastrea radians
AA 2.2% .9 28% SS Siderastrea siderea
% ON Other All other species combined
MC 1.5% -
2.7% AA
PA 3.1%
3.3% PP
3.2%
SS
ss
4.5% MACX
PA6 MC 66.6%
PA MC
4.69%
MACX 9.9%
85.0%



jure 4.18. Percentage of coral species composition at nearshore sites, mid-shelf sites, shelf-edge sites and all sites combined for
Thomas. 'Other' denotes percent of all other coral species combined and includes: Agaricia grahamae, A. humilis, Dendrogyra
findrus, Diploria clivosa, D. labyrinthiformis, Eusmilia fastigiata, Manicina areolata, Mycetophyllia aliciae, M. danaana, M. lamarckia-
SP divaricata, Solenastrea bournoni, S. hyades, and Stephanocoenia michelinii. Source: S. Herzlieb, unpublished data.


2.5
S Nearshore
Mid-shelf
Shelf-edge
2.0



1.5


t 1.0-



0.5



0.0 ,
BB BP SC FC GB RH

Nearshore Mid-shelf Shelf-edge

jure 4.19. Shannon-Weaver Diversity Index (H') for corals at eight monitored sites
St. Thomas: BB=Benner Bay, BP=Black Point, SC=Seahorse Cottage Shoal, FC=
)t Cay, GB=Grammanik Bank, RH=Red Hind Bank. BB and BP are nearshore
se Z nrId C pr- mird_ehlf eif+C nrId ('P nrd D I-I r chalf_drln cif+e n-f +-rn_




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


UVI-CMES AGRRA Assessments of Benthic Substrates

Methods
Between May 1998 and August 2000, 16 sites within the USVI were surveyed (Nemeth et al., 2003a) using the
Atlantic and Gulf Rapid Reef Assessment protocol (AGRRA; Version 2.0). The AGRRA protocol focuses on
three aspects of benthic reef communities: coral condition, algae abundance, and sea urchin density along a
10-m transect. To assess coral condition, the dimensions of 50-100 coral colonies, occurring directly beneath
the transect line were measured. Coral colonies >25 cm in diameter were inspected for signs of disease,
predation, and overgrowth. The percent of old or recent tissue mortality was also estimated for each coral
colony from a planar view. Along these same transects, the point intercept method was used to estimate
percent coral cover, and the number of Diadema antillarium sea urchins occurring within 1 m of each side of
the transect line were counted. Finally at least 50 quadrats (0.25 m2) were placed along the transect lines to
estimate the percent cover and height of macroalgae, turf algae, and coralline algae, and to count the number
of coral recruits <2 cm in diameter.

The assessment sites included eight reefs on St. John, five reefs on St. Thomas and three reefs on St. Croix.
The data were summarized by depth (< 5.5 m and > 6 m) and geographic region (St. Thomas/St. John and St.
Croix). St. Croix was considered a unique geographic region because of its isolation from the northern Virgin
Island Archipelago, its unique geology (sedimentary/carbonate), and its location completely within the Carib-
bean Sea. St. Croix sites included Cane Bay, Salt River East Wall, and Long Reef. St. Thomas and St. John
were grouped as the northern Virgin Islands because of their close proximity, similar geographic origins and
topography (high volcanic islands), and exposure to both Atlantic waters from the north and Caribbean waters
from the south. Reefs around St. Thomas included Brewer's Bay, Buck Island, Caret Bay, Flat Cay and Sprat
Bay. Reefs around St. John included two sites in Great Lameshur Bay (Tektite, Yawzi Point) and two sites in
Fish Bay (outer east and west). Shallow reefs <5.5 m on St. John included two sites in Great Lameshur Bay
(Donkey Bight and VIERS) and two sites in Fish Bay (inner east and west). The AGRRA protocol is described
in detail in Ginsburg et al. (1996).


Results and Discussion
The percent cover of living coral ranged from 10% to 35% in the Virgin Islands. Average cover of living coral
on reefs deeper than 6 m was very similar between St. Thomas/St. John and St. Croix, but was significantly
lower on the shallow reefs of St. John (Table 4.5; Nemeth et al., 2003a). Large stony corals that were indi-
vidually surveyed were numerically dominated by the Montastraea annularis species complex in the shallow
and deeper reefs around St. Thomas/St. John whereas similar reefs in St. Croix were dominated by M. caver-
nosa. The second most common taxon was Siderastrea siderea. The differences in the AGRRA data for St.
Croix and the video assessment data presented above most likely resulted from differences in methods used
(AGRRA only assessed colonies greater than 25 cm whereas the video method included colonies of all sizes).
Moreover, the three sites surveyed forAGRRA were located on the north coast of St. Croix, whereas the larger
number of sites (n=12) assessed for the video method were distributed around the entire island.

Table 4.5. Summary data for corals from AGRRA assessment of USVI reefs around St. John, St. Thomas, and St. Croix and the shal-
low reefs <5.5 m around St. John. Source: Nemeth et al., 2003b.




of Coral Reef Ecosystems of the U.S. Virgin Islands


Between 1998 and 2000 the condition of coral colonies varied among island groups. Coral bleaching was
recorded at all sites with the highest average values occurring on St. Croix, the lowest occurring around St.
Thomas, and moderate levels around St. John (Table 4.6). Alternatively, incidence of disease was lowest on
St. Croix and the shallow reefs of St. John but higher on the deeper reefs of the northern Virgin Islands. Divers
were able to recognize four general disease types: black band, yellow blotch, white plague, and dark spots.
The coral species most susceptible to disease included M. faveolata, M. franksi, M. cavernosa, M. annularis,
Colpophyllia natans, and Siderastrea siderea. The high percentage of coral colonies with fish bites contrib-
uted to the elevated level of recent tissue mortality on shallow reefs of St. John. These shallow nearshore
reefs were also affected by sedimentation especially those outside the boundaries of the VINP (i.e., Fish Bay)
that had high levels of old tissue mortality.

Table 4.6. Summary data for algae (macro, turf, crustose coralline) and coral recruitment from AGRRA assessment of reefs around
St. John, St. Thomas, and St. Croix and the shallow reefs <5.5 m around St. John. Source: Nemeth et al., 2003b.




4 S- ,' 41 P 4. 1 71 24 11 1:.

10,.. 2'.2 25 509 '14 177 1TT
11 '. 170 1 1 7.41 10 10 1

Stony coral recruitment varied considerably from site to site, but on average, it was similar among reefs great-
er than 6 m depth (Table 4.6). Coral recruitment on the shallow reefs of St. John was about 50% of that on
deeper reefs. With the exception of S. siderea, coral recruits were dominated by species that brood their lar-
vae. The five most abundant taxa S. siderea (23%), Agaricia spp. (17%), Porites astreoides (15%), P porites
(13%) and S. radians (6%) comprised 70% to 80% of the recruits on all islands (Nemeth et al., 2003b). The
relative abundance of macroalgae was significantly lower on St. Croix compared with the northern Virgin Is-
land reefs and the shallow reefs of St. John, which had over two times the number of Diadema spp. urchins
than deeper reefs (Table 4.6).



NOAA CCMA-BT Benthic Habitat Mapping
NOAA's Center for Coastal Monitoring and Assessment-Biogeography Team (CCMA-BT) completed a near-
shore benthic habitat mapping project for the USVI in 2002. Aerial photographs were collected by a NOAA ci-
tation jet in 1999 and used to delineate habitat polygons in a geographic information system (GIS). The habitat
polygons were defined and described according to a hierarchical habitat classification system consisting of 26
discrete habitat types. The project mapped approximately 490 km2 of nearshore habitat in the islands includ-
ing coral reefs, mangroves, seagrass beds, and other tropical marine bottom types. A series of 55 maps are
now available via a CD-ROM, and on-line (http://biogeo.nos.noaa.gov/products/benthic. Accessed 1/19/05).
Major habitat types are depicted in Figure 4.20.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


*~-
E Land
Benthic Habitats
SCoral Reef and Colonized Hardbttom


Unconsolidated Sediments
O0tw Deineations
Deep Water


Figure 4.20. Nearshore benthic habitat maps were developed in 2001 by CCMA-BT based on visual interpretation of aerial photog-
raphy and hyperspectral imagery. For more info, see: http://biogeo.nos.noaa.gov. Map: A. Shapiro.






ASSOCIATED BIOLOGICAL COMMUNITIES

ata from four monitoring and assessment programs were used to characterize community structure, bio-
ass, trophic structure, and the size frequency distribution of fish assemblages in the USVI. Mean estimates
standard reef fish assemblage variables were determined from each data set. Species richness is the total
timber of species observed per sample. Abundance is the mean number of individuals per sample. Biomass
the estimated live wet weight of individuals per sample. Live wet weight (W) of each fish was estimated
)m the mean visually estimated fork-length (FL) with the equation: W = a(FL)b, where a and b are known
irameters of the length-weight relationship for each species (Randall et al., 1967; Froese and Pauly, 2000;
tp://fishbase.org., Accessed 12/28/2004). For species not in these databases, estimates from available
erature on the species or congeners were used. The biomass of all fishes recorded in all censuses was
>tained by multiplying the mean live wet weight for each size class for each species by the total number of
dividuals observed in that size class.

3S Long-term Monitoring of Reef Fish Assemblages

methods
inual trends in total species richness, fish abundance, and biomass were analyzed and are presented
parately for the NPS long-term reef fish monitoring dataset. NPS has been monitoring reef fish populations
monthly at four reference sites in the VINP on St. John for 12 years (1988-2000). This data set represents
ie of the longest time series data sets on reef fishes for the territory. An investigation to study the monthly




The State of Coral Reef Ecosystems of the U.S. Virgin Islands

Results and Discussion
In most tropical fisheries, many changes go relatively unnoticed and undocumented. Data acquisition and
monitoring programs are frequently initiated following large resource changes. While this is true for the USVI,
the area has fortunately received much scientific investigation at other times as well. A comparison of histori-
cal data (1958-1961) and more recent monitoring data (1989-2000) provides a view of changes in reef fish
abundance over 60 years.


NPS reef fish monitoring data docu-
mented numerous significant de-
clines in the abundance of several
reef fish over a 12-year period (Beets
and Friedlander, 2003; Figure 4.21).
However, numerous species that
were historically common in landings,
such as the Nassau grouper, demon-
strated no significant trend over the
monitoring period (Figure 4.21). This
may be because their abundance is
presently too low to show significant
trends, assuming that a decline in
abundance has occurred. Further-
more, large declines in abundance for
species, such as the Nassau grouper,
may have occurred before monitoring
projects were initiated.


0.30

0.25

0.20

0.15

0.10

0.05

0.00
0.4

0.3

0.2

0.1

0.0
0.20

0.15

0.10

0.05

0.00


A. Gray angelfish (Pomacanthus arctuatus)


B. Queen angelfish (Holacanthus)


C. Red hind (Epinephelus guttatus)


D. Nassau grouper (Epinephelus striatus)









1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001


Date


Figure 4.21. Significant (A-C) and non-significant (D) declines in abundance of four
commercially-targeted species observed in visual monitoring data from four reefs
around St. John, US Virgin Islands from 1991-2000. Source: Beets and Friedlander,
2003.







Historical data collected by previ-
ous investigators provide compara-
tive information, although compara-
tive abundance data frequently are
not available. For example, Randall
(1967) collected many species of
fish for his landmark studies of Ca-
ribbean reef fishes around St. John
from 1958-1961. Although few were
quantitative, Randall's studies pro-
vided relative abundance and size
structure of species. Large grou-
pers frequently captured by Randall
in 1958-61 were in very low relative
abundance in the 1989-2000 moni-
toring data (Figure 4.22). The two
smaller-sized groupers, red hind and
coney, were much more common in
the recent monitoring data. These
long-term comparisons suggest that
large changes have occurred in Vir-
gin Islands fisheries, similar to pat-
terns observed throughout the Carib-
bean. Over-exploitation by fisheries
certainly has been a strong contribu-
tor to the observed declines.

The most apparent temporal signal
in reef fish assemblage character-
istics around St. John over the 12-
year monitoring period resulted from
the influence of large storm events
(Beets and Friedlander, 2003). The
Virgin Islands have been greatly in-
fluenced by numerous large storms
since 1988. Data were separated
into two periods (1989-1994 and
1996-2000) representing the post-
storm recovery periods following the
two major storms affecting St. John
(Hurricane Hugo, Sept. 1989; Hurri-
cane Marilyn, Sept. 1995). As data
for 1995 were collected just prior to
Hurricane Marilyn, those data were
excluded from analysis. Assemblage
characteristics (species richness,
abundance, and biomass) showed
statistically significant increases dur-
ing the five-year period following
Hurricane Hugo (1989, Figure 4.23).
While species, number of individuals,
and biomass all increased following
Hurricane Marilyn (1995), none of
these trends were significant for the


The State of Coral Reef Ecosystems of the U.S. Virgin Islands


0.7
S1951 -1968
0.6I 1989- 2000
0.6

0.5
E
0.4
0




0.1
0.0

0.0
Nassau Red hind Yellowfin Tiger Rock hind Coney
Species of Grouper
Figure 4.22. Comparison of the relative abundance of groupers collected by Randall
from 1958-1961 and groupers sampled during 1989-2000 around St. John. Source:
Beets and Friedlander, 2003.


o 35-

S30-

S25-

W 20-

0
0S
6


(A) Species Richness


* *0


**
2 = 0.306
p=o.0115


o
0o o
0
o 8 o o

= 0.026
p = 0.5010


1989 1990 1991 1992 1993 1994 1995 1998 1997 1998 1999 2000 2001

7
(B) Abundance

6 -
e.
S* 0

8
r=0.267 r2=0.029
p=0.0197 p = 0.4710
1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001


. 1 -

S10


oE
a 8.

I -


(C) Biomass


* 0
e

S r = 0.322
S p = 0.0091


o o
o 0

o
r2 = 0.019
p = 0.5585


1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001
Sampling Year


Figure 4.23. Trends in assemblage characteristics during the five years following two
hurricanes which affected St. John (Hugo, Sept. 1989; Marilyn, Sept. 1995). Average
values for each of the four reference sites are represented by circles for each year.
Regression lines and coefficients were obtained from linear regression analysis. Data
for 1995 were excluded from these analyses. Source: Beets and Friedlander, 2003.


a




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


five-year period following the storm (Figure 4.23). Large storms that passed near the USVI in 1998 and 1999
may have had a significant negative impact on reef fish assemblage recovery, as lower values in assemblage
characteristics were noted for 2000. Without long-term consistent data, the ability to evaluate such events is
limited.

Current Status of Reef Fish Assemblages in the USVI

Methods
The current status of reef fish assemblages in the USVI was determined from the CCMA-BT, DFW, and UVI-
CMES reef fish monitoring programs. These programs present the most recent data on the status of reef
fishes in the USVI. In 1998, UVI-CMES joined the Caribbean-wide effort to assess reef fish assemblages at
16 sites throughout the USVI. Since Table 4.8. Species of commercially important snappers (Lutjanidae) and groupers
2001, CCMA-BT has surveyed reef (Serranidae) for which estimates of mean biomass density (g/m2) were calculated
fishes semi-annually for three years for the Virgin Islands National Park, St. John and the BIRNM, St. Croix. Source: Ap-
at 309 and 128 hard bottom sites in peldoorn et al., 1992.
St. Croix and St. John, respectively. I 1
Most recently, the DFW conducted 80 Luljanus anal/, mutton snapper
visual surveys and collected 120 trap Lutlanus ap,-is sc hoolmrster
samples at eight permanent hard bot-
weight of groupegran snapper
tom areas in St. Croix during spring Lutian iS
and fall of 2002. LulFanus ]ocu cog snapper
Lutjanus a.-,o mahogany snapper
Mean biomass density of 12 com- Lutantus sinangis lane snapper
mercially important species of grou- Oc.n ius Ce.u isui us yellor..tail snapper
pers and snappers (Table 4.8), the Epnephelu c, genrtatus graysby
trophic biomass ratio of three broad EpitepeIus fi tusL cone
feeding guilds (Table 4.9), and the
size frequency distribution of select- Epnephel t red hind
ed species were calculated for each Epnepeus moan red grouper
site. Biomass density is the live wet IA Tclei opeica aiq.aI tiger grouper
weight of groupers and snappers ob-
served per area (m2) sampled. Trophic biomass ratio is the proportion of live wet weight of fishes in one of
three feeding guilds. Fishes were assigned to trophic guilds according to Randall (1996). However, Randall's
trophic classification was reduced to three trophic groups to simplify the interpretation of the results. Randall's
"mobile invertebrate feeders/piscivores" were integrated into the group "piscivores"; herbivores were not re-
classified; all other trophic groups ("detritivores", "sessile invertebrate feeders", "zooplanktivores", and "om-
nivores") were combined into one category called 'generalized carnivores' (Table 4.9). Size class frequency
is the proportion of individuals of a species belonging to one of eight size classes. Size classes were based
on visual estimates of fork length (FL). Size class frequency was estimated for three commercially important
species red hind grouper (Epinephelus guttatus), coney (E. fulvus), and red band parrotfish (Sparisoma
aurofrenatum) and the bluehead wrasse (Thalassoma bifasciatum), a commonly occurring species with no
commercial importance. These assemblage and species variables were chosen because they can provide
a relative index of the condition of coral reef fish assemblages. Current estimates of these assemblage vari-
ables will be used as a baseline for comparison with estimates from future monitoring data to determine how
reef fish assemblages are changing over time.

Table 4.9. Trophic guilds used to determine trophic biomass ratio of fishes in the USVI. Source: Randall, 1967.


Herbl.ores Marine plants C lamiselflshes parrotfishes surgeonfishes
Flisc.,,ores mobile in .ert .orestpisci ores theire r fish crabs Red hind other groupers snappers
Mobile in erii ores sessiie in..erti ores Cruistaceans corals Spanish hogfish raisess gobies filefish buitterflYfish
zooplankti ores generalized carni ores zooplankton etc blennies cardinal fishes angelfishes squirrel fishes
goatfishes scadblennies cardinal fishes




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


NOAA CCMA-BT
Since August 2000, NOAA's CCMA-BT has led a collaborative effort to monitor coral reef ecosystems through-
out the U.S. Caribbean, including the USVI. This regionally-integrated monitoring effort explicitly links ob-
served fish distributions to shallow (<30 meters) benthic habitats recently mapped by CCMA-BT and its many
partners (Kendall et al., 2001). Objectives of this work include: 1) developing spatially-articulated estimates of
the distribution, abundance, community structure, and size of reef fishes, conch, and lobster; 2) relating this
information to in situ data collected on associated habitat parameters; 3) using this information to establish
the knowledge base necessary to implement and support "place-based" management strategies for coral reef
ecosystems of the Caribbean; and 4) quantifying the efficacy of management actions.

This regional monitoring program has been conducted in partnership with the UVI, NPS, USGS, and DPNR,
and provides standardized monitoring data for portions of the entire U.S. Caribbean. Since the inception of
this effort, over 600 surveys of reef fish populations and associated benthic habitats have been conducted
in southwestern USVI (see Figure 4.9). The foundation of this work is the nearshore benthic habitats maps
created by CCMA-BT in 2001. Using ArcView GIS software, the benthic habitat maps are stratified to select
monitoring stations along a cross-shelf depth gradient. Because the program was designed to monitor the
entire coral reef ecosystem, CCMA-BT and its partners survey seagrass meadows, mangroves, sand flats,
as well as various coral reef formations. Survey sites are selected at random within each habitat stratum to
ensure complete coverage of the study region. At each site, fish, conch, lobster, and benthic habitat informa-
tion is collected using standard visual survey techniques (Christensen et al., 2003). Since 2003, CCMA-BT
has also been collecting water quality and oceanographic characteristic data at each survey location. These
water quality data are not yet available, but will be provided in a future report.

By correlating monitoring data to the habitat maps, CCMA-BT and its partners are able to map and model
(predict) species and community level parameters throughout the seascape. Furthermore, by integrating this
work with other studies being conducted concurrently by its partners on fish migration patterns, home range
size, fish dispersal, and recruitment, CCMA-BT is in a unique position to answer questions about marine zon-
ing strategies (e.g., placement of marine protected areas [MPAs]), and evaluate management efficacy through
long-term monitoring.


USVI-DPNR-DFWS
Surveys of reef fishes were conducted by divers from the DFW. Surveys occurred during fall of 2003 at eight
permanent long-term monitoring sites surveyed annually by researchers from UVI-CMES. The permanent
sites were selected because they were considered representative of the reefs around St. Croix (Nemeth et al.,
2002). Sites were hard-bottom, less than 15 m in depth, and considerably varied in the composition of benthic
flora and fauna (Nemeth et al., 2002).

A60 m2 rectangular transect was used to assess reef fish assemblage structure (Nemeth et al., 2003c). Visual
fish counts were conducted along 10 replicate transects at each site. During fish transects, transect width
and fish lengths (measured in 5-cm increments up to 35 cm) were measured with a 1 m t-bar marked in 5 cm
increments. Using transects as replicates, the average density (no./100m2) and size (cm) of each species and
family were calculated for each site.

The DFW also conducts independent fisheries monitoring of reef fishes with fish traps and hand-lines through
the Southeast Area Monitoring and Assessment Program for the Caribbean (SEAMAP-C; Tobias et al., 2002).
SEAMAP-C is a cooperative program among NOAA Fisheries, the Puerto Rico Department of Natural Re-
sources, and DFW. SEAMAP-C was implemented to collect data needed to assess the status of marine
resources of the U.S. Caribbean and to monitor any changes in status (Tobias et al., 2002). Briefly, 12 baited
fish traps and three hand-lines (each with three hooks) were used to sample reef fishes in a 52 km2 area
northeast of St. Croix on 10 sampling missions between January 2001 and April 2002 (Figure 4.9). Traps were
placed randomly at two depth strata (0-18 m and 19-36 m). Total trap soak time was 59 hours and total hand-
line fishing time was also 59 hours. A detailed description of the SEAMAP-C sampling methods is provided
in Tobias et al. (2002).




The State of Coral Reef Ecosystems of the U.S. 1


UVI-CMES
Between May 1998 and August 2000 16 sites within the USVI were surveyed (Nemeth et al., 2003b) with the
AGRRA protocol (Version 2.0; Ginsburg et al., 1996). Visual fish counts along at least 10 60-m2 transects were
conducted at each site. On St. Croix, additional surveys were conducted at Cane Bay (n=3), Long Reef (n=5)
and Salt River (n=6). Transect width and fish lengths (measured in 5-cm increments up to 35 cm) were esti-
mated using a 1 m wide t-bar constructed of pvc. Using transects as replicates, the average density (no./100
m2) and size (cm) of each species and family were calculated for each site and island group (see below).
Parrotfish and grunts less than 5 cm were counted and identified to species when possible at all sites except
St. Croix. Sites were identical to those listed in the UVI-CMES AGGRA Assessments of Benthic Substrates
section of this chapter.


Results and Discussion
Despite the variation in sampling techniques, spatial extent, and temporal coverage, analysis reveals patterns
in the abundance and assemblage structure of reef fishes that were consistent among the data sets. These
patterns are described below.

The biomass of commercially important groupers and snappers was very low for all monitoring programs.
Mean biomass density of groupers and snappers was 5.67 0.55 g/m2 (CCMA-BT) and 8.76 1.17 g/m2 (UVI-
CMES). Furthermore, the NPS long-term reef fish data show clearly that the average number and frequency
of occurrence of groupers decreased at reference sites during 12 years of sampling (Figure 4.22). Low esti-
mates of biomass also reflect low abundance of groupers and snappers in USVI waters, and indicate a lack
of recovery of local grouper populations to fishable levels. Intense fishing pressure, degradation of coral reef
habitats, and tropical storm events have contributed to the demise of several large-sized grouper and snapper
species such as the Nassau grouper, Epinephelus striatus and the dog snapper, L. jocu in the USVI (Olsen
and LaPlace, 1978; Beets and Friedlander, 1992; Rogers and Beets, 2001). Now, the abundance of smaller-
sized groupers (e.g., red hind, E. gutatus; coney, E. fulvus; graysby, E. cruentatus) that have replaced the
decimated fisheries are so low that they too are rarely caught in recreational or commercial fisheries (W To-
bias, pers. comm.). Continued monitoring of grouper/snapper biomass density would provide an easy way to
assess the future trends and health of USVI reef fish assemblages.

Overfished reef fish assemblages typically are characterized by a higher proportion of herbivores and fewer
piscivores compared with unfished assemblages. Many large-bodied predatory species (e.g., groupers and
snappers) usually are the primary targets of fishers, which results in the depletion of the largest and most valu-
able fishes from reef fisheries. As the
abundance of these larger species 100
decrease to unfishable levels, fishers m Herbivores
are forced to switch to smaller and Generalized camivores
SPiscivoMres
more undesirable fishes, a phenom- so80 is
enon known as "serial overfishing"
(see Ault et al., 1998). Assuming a
reduction in fishing pressure may re- o -
suit in an increase in the abundance
of predators, monitoring temporal M
changes in the trophic structure could a 40
provide another way to determine
the status of USVI reef fish assem- 20


UdLd U UIItUL; -U UY LI IIt II IUI IILUII lIY dl U I I It-- I JIUY L dli II UY II CrID I, LoV I L)FVV,
-4 I\/I rhRA Q nllrc'c VanrniI lt aIi onnil nImith at i )nnqn rn





The State of Coral Reef Ecosystems of the U.S. Virgin Islands

fish biomass. Other trophic guilds accounted for 38% (CCMA-BT), 67% (DFW), and 64% (UVI-CMES) of the
observed biomass. Comparisons of current baseline data with future estimates of trophic biomass ratios could
indicate whether fishing pressure on UVSI reef fish assemblages is increasing or decreasing.

The size frequency distributions of groupers suggest that grouper populations in the USVI consist predomi-
nantly of small-sized individuals. The average adult size of a red hind grouper ranges from 25-38 cm, with a
maximum known length of 61 cm (Humann and Deloach, 2002). The coney is smaller with an average adult
size ranging from 15-25 cm and a maximum length of 40 cm (Humann and Deloach, 2002). Eighty-three per-
cent of the 909 red hind and coney groupers observed by CCMA-BT were smaller than 25 cm in size (Figure
4.25A). DFW caught 513 red hind and 46 coney groupers during trap and hand-line fishing in St. Croix. Of
these, 89% were smaller than 25 cm in size (Figure 4.25B). UVI-CMES divers observed 30 red hind and 72
coney groupers, 94% of which were smaller than 30 cm in size (Figure 4.25C).

Most redband parrotfish observed in the USVI were smaller than the average size of an adult. The average
adult-sizde redband parrotfish ranges from 15-25 cm (Humann and Deloach, 2002). A total of 3,043 redband
parrotfish were observed during 373 CCMA-BT surveys (Figure 4.25D). Thirty-four percent (1,035 individuals)
were 0-5 cm in length. The number of individuals decreased consistently as size-class increased, and only
21% were larger than 15 cm in length. DFW divers observed 590 red band parrotfish grouped into four size-
classes, and 93% were less than 20 cm (Figure 4.25E). UVI-CMES reported 721 redband parrotfish grouped
into five size-classes, with 82% being less than 20 cm in size (Figure 4.25F). The size frequency distribution


CCMA-BT reef fish assessments
n-43710~ teansect


] I


SRedhind grouper
I Coney


L,


0-5 5-10 10-1515-2020-2525-3030-35 >35


I= Redband parroish


0-5 5-10 10-1515-2-220-2525-3030-35>3

1 Blueheadwss
M__________an ur


USVI DFW independent fisheries monitoring
n (Red hind grouper & Coney) 120 trap samples
n (Redband parrotfih & Blueheed wresse) = 80 60m transcts
300-
25o- a


o 5 5 10151-2020-2525-303535-4
<5 5-10 10-'1515-2020-2525-03030-35540


0-5 5-10 10-20 20-30 30-40 >40


UVI CMES reef fish monitoring
n 184O0m' transcts


0-5 6-10 11-20 21-30 31-40 >40


0 I 1
0-5 6-10 11-20 21-30 31-40 >40


0- 5-10 10-2 20430 3040 > 40
Size (cm)


0-5 6-10 11-20 21-30 31-40 >40
Size (cm)


Figure 4.25. Size frequency histograms for four reef fishes based on data collected by three monitoring and assessment programs:
CCMA-BT, USVI DFW, and UVI CMES. Sources: Kendall et al., 2003; Nemeth et al., 2003a,c.


12000 -
10000 -
BW-
8000 -
6000-
zo-
4000-
2000 -


05 5-10 10-155-2020-25253030-35 >35
Size (cm)




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


indicates that redband parrotfish populations in the USVI generally consisted of immature individuals.

Bluehead wrasse populations in the USVI also consisted primarily of individuals smaller than the average
adult size (10-13 cm; Humann and Deloach, 2002). CCMA-BT divers observed 15,337 bluehead wrasse on
398 of 437 surveys (Figure 4.25G). Most (98.5%) were less than 10 cm in length. DFW divers observed 4,959
bluehead wrasse in three size-classes and 99% were less than 10 cm in length (Figure 4.25H). UVI-CMES
reported 375 bluehead wrasse grouped into three size-classes, and 98% were smaller than 10 cm in size (Fig-
ure 4.251). The size frequency distribution of the bluehead wrasse indicates that the USVI populations consist
primarily of juveniles and immature adults.

In summary, fish species composition on reefs and in fisheries catch has shifted to more herbivorous species
since 1988. Additionally, there has been a decline in the number of grouper and snapper species as well as
the average size of fishes observed on reefs during field surveys. Commercially important species such as
large grouper and snapper species, that once abounded on USVI reefs during the 1950s and 1960s are cur-
rently of low abundance in fisheries landings. Continued monitoring of the status of reef populations and reef
fisheries as well as commercially important macroinvertebrates (e.g., conch and lobster) is important.


CURRENT CONSERVATION MANAGEMENT ACTIVITIES

The U.S. Department of the Interior (DOI), U.S. Department of Commerce (DOC), and Virgin Islands Territo-
rial Government all have jurisdiction in overlapping sections of submerged lands within the USVI (Table 4.10).
These agencies have conducted several research and monitoring activities to aid in the management of USVI
coral reef ecosystems. Both Federal and territorial agencies in the USVI use a variety of management tools
to address issues such as fishing, recreational use, and land-based sources of pollution to protect the marine
resources of the territory.

Table 4.10. Authorities with jurisdiction over waters and submerged lands with coral reefs in the USVI.


Leas~Ing ret ponr sibilit for Federal submerged lands f.,ethin 2tii nmni of










Mapping
BUCk Island Reef IJational Monument Salt RI .er Bay r latonal H .ltorical
Park and Ecological Preser 'e ,., rgin Islands rJajtonal Park ,.,rgin Island.
Coral Reef I national F..1onument
3.-20i nmi
cll1 other aters and submerged lands from the shoreline to 3 nmi



Mapping
In 2000, an extensive seafloor mapping project around the USVI was completed by NOAA, local partners, and
the U.S. Coral Reef Task Force. For this project, much of the insular shelf of the USVI from the shoreline to
a depth of approximately 20 m was mapped using visual interpretation of aerial photographs, a 26-category
classification scheme, and a minimum mapping unit (MMU) of 1 acre (NOAA, 2001). Completed maps cover
approximately 490 km2 of benthic features including mangroves, seagrass, and coral reefs. These maps have
been used for a wide variety of research and management applications, including stratification of sampling
effort in reef fish monitoring projects, distribution and abundance surveys for elkhorn coral and lobster popula-
tions, and an inventory of cryptic fish. Mapping projects since 2000 have primarily covered smaller areas of
particular interest in the USVI, focused on specific bottom types, and had smaller MMUs. Aerial photos have
been used recently to map Buck Island and Salt River, St. Croix at a finer scale (100 m2 MMU) than was done
previously by CCMA-BT and NPS. These activities are focused on providing a more refined inventory of habi-
tat types in the national parks located at those sites.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Automated computer analysis of historical and current aerial photos was recently completed to detect changes
in seagrass beds northeast of St. Croix (Kendall et al., 2004). This information is currently being used to
establish records of this critical habitat in a location where anchor damage has historically been a problem.
Lidar has also been collected by USGS northeast of St. Croix and around St. John for fine-scale bathymetry
and habitat mapping. Several groups at NOAA are using satellite data to map benthic cover and bathymetry.
LandSat has been used to map bathymetry around the USVI (EarthSat), with cover mapping currently un-
derway. IKONOS is being used by CCMA-BT to map bathymetry and bottom types around Buck Island, St.
Croix.

Several sonar-based projects have also been completed or are underway in the region. These projects cover
areas too deep or too turbid to map with either aerial or satellite-based sensors. Side scan sonar has been
used by the Caribbean Fishery Management Council and DPNR to map the marine conservation district south
of St. Thomas along with some nearby areas to aide fisheries management. An upcoming project by CCMA-
BT and partners will use multi-beam sonar to map bottom features below 20 m around the BIRNM, St. Croix
and along the mid-shelf reef south of St. John and St. Thomas. The cumulative result of these and future
projects will be continuous map coverage of benthic cover and bathymetry from the shoreline to deep water
areas beyond the insular shelf.

Marine Protected Areas
MPAs are used as management tools to protect, maintain, or restore natural and cultural resources in coastal
and marine waters. They have been used effectively both nationally and internationally to conserve biodiver-
sity, manage natural resources, protect endangered species, reduce user conflicts, provide educational and
research opportunities, and enhance commercial and recreational activities (Salm et al., 2000).

Many different types of MPAs have been established throughout the USVI to provide different levels of protec-
tion for resources based on their size, management goals, and intended purpose (Table 4.11). Over the years,
the number and size of these protected areas have grown steadily, thereby providing protection to a greater
proportion of coral reef ecosystems. Recently, additional marine areas have been set aside for protection
through Federal and local legislation.

The BIRNM is a large coral reef national park located off the island of St. Croix. The monument, originally
established in 1962 by Presidential Proclamation, included a tropical dry forest island (0.7 km2) and 2.9 km2
of submerged land. Created to protect the island's elkhorn coral (Acropora palmata) barrier reef, the original
park boundaries did not fully encompass all essential coral reef habitats or the unique "haystack" formations
along the north side of the reef.

When the USVI was highlighted by the U.S. Coral Reef Task Force in 1999, the Secretary of the Interior ac-
tively sought to improve protection for coral reef areas under DOI jurisdiction. In 2001, this effort resulted in
two Presidential Proclamations, one expanding BIRNM by adding over 75 km2 acres of submerged lands,
and another creating the Virgin Islands Coral Reef National Monument (VICRNM) on St. John. The VICRNM
contains 48.9 km2 of marine waters adjacent to the VINP, including bank shelf and spur-and-groove reef
formations, mangrove shorelines, hardbottom habitat, and seagrass beds. VICRNM is almost entirely a no-
take area (fishing for baitfish and blue runner in a specified zone is allowed) and anchoring is prohibited. The
BIRNM expansion not only added many of the missing and essential coral reef habitats (seagrass, sand,
shallow and deep shelf-edge reefs, and deep pelagic areas), but it also made the entire park a no-take area.
Anchoring at BIRNM requires a permit.

BIRNM and VICRNM are two of the four units in the National Park System that contain fully-protected marine
reserves. The parks were given two years to develop new general management plans and a vessel manage-
ment plan, but the expansion of BIRNM and new regulations prohibiting all extractive uses were legally chal-
lenged by the USVI territorial government. To determine if the President had the right to expand BIRNM and
create VICRNM, the Virgin Islands' delegate to Congress requested that the U.S. General Accounting Office
review the Presidential Proclamations. The review, which took almost 18 months, found that the Proclama-
tions were valid. The regulations for both monuments went into effect on May 5, 2003.







Table 4.11. USVI Marine Managed Areas (MMA) and their management agencies.



Cas Cad Ale..e Small Pond at Fianl< St Con,. East End lai,,ne Pati< USvl
La..goon., Pes.e, Ea \\Idllife and A anne established 200'. In early, stages o eminent
Sancituai, of implementation
St Jarmes Pese eP:- Sail Pieti Ba\ National HiaSIncal I JPS
Pal riand Ecologcal Pteseie
Compass ,Poi, nt established 1992 e-panded
Alaine Peserte ando 1975 and 20,0:. by P'residential
S\\ildllife Sanctiuat Proclamations
\ ,tain IslaQ 's Co ,-al P-ee-f ELCI islatt Peer Nati-onal I IF'PS
\National monument Aliiumenti established 19152
established 2001 by expanded 1'-975and 2001 by
Presidential Proclamalions Presidential Proclamations

S\i.in Islatnds National Salt Ptet ea\ B iatltonal Histoincal I IPS
Patl, establihsned 1'9".:?. Patr and Ecolo. cal Pe.ee "
expanded marine portions established 19';92 expanded 2002
added in 191i.2 ointl. th territorial go emin ent
Ped n17'dti ClIOSI e Ped 1ai d C71 e closed iUSvi
Closed year round !Decenmber 1 February 28
Vu&Jitlon snappCe Cnlosuie clOsed Joint Federal an
I,,'larch 1 June ;.0 Territorial
0_-,o eGrnment
<41tona Lacoon and d-,3eat Potnd Territorial
shrimp management area .o..ernment
restricted gear ulse


The most recent addition to the existing network of protected areas is the St. Croix East End Marine Park, t
first territorial park designated by the USVI Legislature in January 2003. With this designation, the USVI Le




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Other Management Tools
The DFW has deployed several fish 4 + St Thomas port
aggregating devices (FADs) in terri- -o- St. Thomas SPAG
trial waters and the adjacent Exclu-
sive Economic Zone in order to take
fishing pressure off the reefs and
promote a shift to pelagic fishes (To- E
bias, 2001). Six FADS are currently
deployed around St. Croix and three 32
around St. Thomas. Efforts are un- ,
derway to increase these numbers 30
during 2004.
28-
Mooring buoys have been installed
throughout the territory by Federal 26
and territorial agencies as a manage-
ment tool to decrease recreational 24
impacts on coral reefs and related 1974 1977 1980 1983 19i 1989 1992 1995 1998 2001 2004
ecosystems. Mooring buoys are well Date
used by dive operators, recreational
fishers and boaters. Funding has Figure 4.26. Length of red hind from fishery port surveys conducted over 30 years and
been secured by the territorial gov- from red hind spawning aggregation (SPAG). Modified from Nemeth (in review).
been secured by the territorial gov-
ernment to increase the number of
mooring buoys throughout the ter- 30o
ritory, especially within the St. Croix
East End Marine Park.
25"
Outside of managed areas, fishing is E
regulated under Federal and territori- 20
al rules and regulations. Size restric- 8
tions exist for whelks, conch, and lob-
ster. The harvest of goliath grouper a. is -
(E. Itajara) and Nassau grouper (E.
striatus), as well as the commercial *
harvest of billfish is prohibited. Other 10
restrictions are in place. The territo- a
rial fishing rules and regulations are 5 T
currently under review and will be re-
vised in the near future.
1977 2000 2001
Date
Figure 4.27. Biomass of spawning red hind at the Marine Conservation district south
of St. Thomas USVI. Source: 1997 data: Beets and Friedlander (1999); 2000-2001:
Nemeth (in review).



OVERALL CONCLUSIONS AND RECOMMENDATIONS
This report has identified several threats facing coral reef ecosystems in the USVI. Current assessments
indicate that water quality is generally good, but it is declining because of an increase in point and nonpoint
source discharges into the marine environment. Coral diseases remain abundant and epidemic, and the per-
cent cover of coral remains low, while macroalgae abundance on reefs remains high. Dense stands of elkhorn
coral that occurred on reefs during the 1960s and 1970s have not recovered to date. Likewise, populations of
large-sized grouper and snapper species, which were abundant on reefs and were common in fisheries land-




The State of Coral Reef Ecosystems of the U.S. 1


wings 4u years ago, nave not reDounaea. iNotwitnsianaing, ine size ana numbers or groupers ana snappers
spawning within some enforced MPAs may be increasing, which is very encouraging. Due to the existence of
several MPAs, current coral reef ecosystem conditions could improve with: 1) a reduction in the number and


Gaps, Problems, and Recommendations
Although the importance of MPAs has been recognized and much effort has been put into their establishment
by government agencies and NGOs, a lack of enforcement of MPAs is a major problem. Minimal enforcement
stems from a lack of management capacity caused by understaffed teams and limited project funding. The
establishment of MPAs is meaningless unless rules and regulations governing those areas can be properly
enforced. This is also true for territorial and Federal fishing regulations, which are not enforced because en-
forcement offices are understaffed. This issue must be addressed before additional or stricter regulations can
be proposed and enacted.

Another problem caused by a lack of capacity is the absence of a flow of information between research and
management programs within and among management agencies. For instance, monitoring programs have
collected several years' worth of data, but analyses of these data have been delayed. Limited human resourc-
es caused by a lack of funding are a primary reason why the results and recommendations from data analyses
sometimes are not available to local managers in a timely manner. Thus, management decisions concerning
resource issues usually are not proactive. Additionally, research and management programs run by territorial
agencies are supported mainly through funding from Federal agencies (EPA, NPS, NOAA and USGS) rather
than through funding from the territorial government. Consequently, the direction and emphasis of research
programs in the USVI are often directed by the programmatic mandates of non-territorial funding agencies
rather than by specific resource issues that affect the territory.

The fact that several jurisdictions are involved in resource management has led to conflicts in the past. Ap-
proaches are very different among management agencies and jurisdictions, and conflicts have arisen where
jurisdictions overlap. Whereas the territorial government tried to involve stakeholders and communities, the
establishment of monuments and national parks has been a top-down approach. This approach has led to
conflicts between managers of monuments and territorial management agencies, such as when objections
were raised, and are still being raised to the 2001 presidential proclamation that expanded the BIRNM. The
territory is also in need of management plans for all designated APC. As of now, these areas exist only on pa-
per and are useful only for supporting permit decisions for coastal development. Along with efforts to address
the issues here, the following actions would be very valuable in helping to manage and protect resources in
the USVI. These actions include the following:

1. Establishing acceptable limits of change or carrying capacity for protected areas;

2. Training judges on adjudication of environmental issues and concerns;

3. Hiring trained environmental prosecutors (as environmental crimes are currently of low priority in the terri-
tory);
4. Shifting toward eco-tourism and increased support and promotion of sustainable and ecologically sound
coastal development by the territorial government; and

5. Establishing pollution control criteria.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


REFERENCES

Adams, A. 1995. Final Report, Recreational Fishery Assessment Project. Division of Fish and Wildlife, Department of
Planning and Natural Resources, U.S. Virgin Islands. 50 pp.

Adey, W.H. 1975. The algal ridges and coral reefs of St.Croix their structure and Holocene development. Atoll Research
Bulletin 187: 1-67.

Adey, W.H., W Gladfelter and R.D. Ogden. 1977. Field Guidebook to the reefs and reef communities of St. Croix, Virgin
Islands. Atlantic Reef Committee, University of Miami, Miami Beach, Florida. 52 pp.

Anderson, D.M. and L.H. MacDonald. 1998. Modelling road surface sediment production using a vector geographic infor-
mation system. Earth Surface Processes and Landforms 23: 95-107.

Appeldoorn, R., J. Beets, J. Bohnsack, S. Bolden, D. Matos, S. Meyers, A. Rosario, Y. Sadovy and W Tobias. 1992.
Shallow Water Reef Fish Stock Assessment for the U.S. Caribbean. National Oceanic and Atmospheric Administration
Technical Memorandum NMFS-SEFSC-304. 70 pp.

Ault, J.S., J.A. Bohnsack and G.A. Meester. 1998. A retrospective (1979-1996) multi-species assessment of coral reef
fish stocks in the Florida Keys. Fishery Bulletin 96: 395-414.

Beets, J. 1993. Long-term monitoring of fisheries in Virgin Islands National Park: Chapter I. Baseline data, 1988-1992,
with emphasis on the impact of Hurricane Hugo. Virgin Islands National Park, Technical Report, VINP 1/93.

Beets, J. 1996. The effects of fishing and fish traps on fish assemblages within Virgin Islands National Park and Buck
Island National Monument. U.S. Virgin Islands National Park Technical Report 5/96. 44pp.

Beets, J. 1997. Can coral reef fish assemblages be sustained as fishing intensity increases? Proceedings of the 8th In-
ternational Coral Reef Symposium 2: 2009-2014.

Beets, J., L. Lewand and E. Zullo. 1986. Marine community descriptions and maps of bays within the Virgin Islands Na-
tional Park/Biosphere Reserve. Virgin Islands Resource Management Cooperative. Biosphere Reserve Research Report
2. National Park Service. 117 pp.

Beets, J. and A. Friedlander. 1990. Long-term monitoring of fisheries in the Virgin Islands National Park: Impact of Hur-
ricane Hugo. Annual Project Report to Virgin Islands National Park. 20 pp.

Beets, J. and A. Friedlander. 1992. Assessment and management strategies for red hind, Epinephelus guttatus, in the
U.S. Virgin Islands. Proceedings of the 42nd Gulf and Caribbean Fisheries Institute 42: 226-242.

Beets, J. and A. Friedlander. 1999. Evaluation of a conservation strategy: a spawning aggregation closure for red hind,
Epinephelus guttatus, in the U.S. Virgin Islands. Environmental Biology of Fishes 55: 91-98.

Beets, J. and A. Friedlander. 2003. Temporal analysis of monitoring data on reef fish assemblages inside the Virgin
Islands National Park and around St. John, U.S. Virgin Islands, 1988-2000. Final Report to the U.S. Geological Survey
Caribbean Field Station, St. John, Virgin Islands.

Bohnsack, J.A. and S.P. Bannerot. 1986. A stationary visual census technique for quantitatively assessing community
structure of coral reef fishes. NOAA Technical Report NMFS 41. 15 pp.

Bohnsack, J.A., S. Meyers, R. Appeldoorn, J. Beets, D. Matos and Y. Sadovy. 1991. Stock Assessment of Spiny Lobster,
Panulirus argus, in the U.S. Caribbean. Report to the Caribbean Fishery Management Council. Miami Laboratory Con-
tribution No. MIA-90/91-49.

Brownell, W.N. 1971. Fisheries of the Virgin Islands. Commercial Fisheries Review 33 (11-12): 23-30.

Brownell, W.N. and WE. Rainey. 1971. Research and development of deep water commercial and sport fisheries around
the Virgin Islands plateau. Virgin Islands Ecological Research Station, Contribution No. 3. 88 pp.

Bruckner, A.W 2001. Coral health and mortality: Recognizing signs of coral diseases and predators. pp. 240-271 In: P.
Humann and N. Deloach (eds.) Reef Coral Identification: Florida Caribbean Bahamas. New World Publications, Inc.,
Jacksonville, FL.

Bryant, D., L. Burke, J. McManus and M. Spalding. 1998. Reefs at risk: a map-based indicator of threats to the world's
coral reefs. World Resources Institute. Washington, D.C. 59 pp.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Burke, L. and J. Maidens. 2004. Reefs at Risk in the Caribbean. World Resources Institute, Washington, DC. 80 pp. Avail-
able from the internet url: http://marine.wri.org/publications.cfm.

Bythell, J.C., E.H. Gladfelter and M. Bythell. 1992. Ecological studies of Buck Island Reef National Monument, St Croix,
US Virgin Islands. A quantitative assessment of selected components of the coral reef ecosystem and establishment of
long-term monitoring sites. Part II. US National Park Service, Cruz Bay, St John, U.S. Virgin Islands. 72 pp.

Bythell, J.C., E.H. Gladfelter and M. Bythell. 1993. Chronic and catastrophic natural mortality of three common Caribbean
reef corals. Coral Reefs 12: 143-152.

Bythell, J.C., Z. Hillis-Starr and C.S. Rogers. 2000. Local variability but landscape stability in coral reef communities fol-
lowing repeated hurricane impacts. Marine Ecology Progress Series 204: 93-100.

Catanzaro, D., C. Rogers, Z. Hillis-Satrr, R. Nemeth and M. Taylor. 2002. Status of coral reefs of the U.S. Virgin Islands.
pp. 131-150 In: Turgeon D.D., R.G. Asc, B.D. Cause, R.E. Dodg, W Jaap, K. Banks, J. Delaney, B.D. Keller, R. Speiler,
C.A. Mato, J.R. Garcia, E. Diaz, D. Catanzaro, C.S. Rogers, Z. Hillis-Starr, R. Nemeth, M. Taylor, G.P. Schmahl, M.W
Miller, D.A. Gulko, J.E. Maragos, A.M. Friedlander, C.L. Hunter, R.S. Brainard, R. Craig, R.H. Richmond, G. Davis, J.
Starmer, M. Trianni, R. Houk, C.E. Birkeland, A. Edward, Y. Golbuu, J. Gutierrez, N. Idechong, G. Paulay, A. Tafileichig
and N.V. Velde. The state of coral reef ecosystems of the United States and Pacific freely associated states: 2002. NOAA/
NOS, Silver Springs, MD. 265 pp.

CFMC (Caribbean Fishery Management Council). 2003. Draft Environmental Impact Statement for the Generic Essential
Fish Habitat Amendment to: Spiny Lobster Fishery Management Plan, Queen Conch Fishery Management Plan, Reef
Fish Fishery Management Plan, Coral Fishery Management Plan for the U.S. Caribbean. Caribbean Fishery Managment
Council Volume 1.

CH2M Hill Inc. 1979. A sediment Reduction Program. Report to the Department and Cultural Affairs. U.S. Virgin Islands
(USVI). Government of the USVI. 88 pp.

Christensen, J.D., C.F.G. Jeffrey, C. Caldow, M.E. Monaco, M.S. Kendall, and R.S. Appeldoorn. 2003. Cross-shelf habitat
utilization patterns of reef fishes in southwestern Puerto Rico. Gulf and Caribbean Research 14: 9-27.

Coles W 2004. Virgin Islands. Department of Planning and Natural Resources. St. Croiz, US Virgin Islands. Personal
communication.

deGraaf, J. and D. Moore. 1987. Proceedings of the conference on fisheries in crisis. Government of the United States
Virgin Islands, Department of Planning and Natural Resources, Fish and Wildlife Division. 147 pp.

Devine, B., G. Brooks, and R. Nemeth. 2003. Coral Bay Sediment Deposition and Reef Assessment Study. State of the
Bay Final Project Report, submitted to the V.I. Department of Planning and Natural Resources, Division of Environmental
Protection. Non-Point Source Pollution Grant Program MOA# NPS -01801. University of Virgin Islands, St. Thomas, U.S.
Virgin Islands.

Edmunds, P.J. 1991. Extent and effect of black band disease on a Caribbean reef. Coral Reefs 10: 161-165.

Edmunds, P.J. and J. Witman. 1991. Effect of Hurricane Hugo on the primary framework of reefs along the south shore
of St John, US Virgin Islands. Marine Ecology Progress Series 78: 201-204.

Fiedler, R.H. and N.D. Jarvis. 1932. Fisheries of the Virgin Islands of the United States. U.S. Dept. of Commerce, Bureau
of Fisheries. Investigational Report 14. 32 pp.

Friedlander, A.M. 1997. Status of queen conch populations around the northern US Virgin Islands with management rec-
ommendations for the Virgin Islands National Park. Biological Resources Division, United States Geological Survey.

Froese, R. and D. Pauly (eds.). 2000. Fishbase 2000: concepts, design, and data sources. ICLARM, Los Banos, Laguna,
Phillipines.

Garrison, V.H., C.S. Rogers and J. Beets. 1998. Of reef fish, overfishing, and in situ observations of fish traps in St. John,
U.S. Virgin Islands. Revista de biologia tropical 46: 41-59.

Garrison, V.H., E.A. Shinn, W.T. Foreman, D.W Griffin, C.W. Holmes, C.A. Kellogg, A. Christina, M.S. Majewski, L.L.
Richardson, K.B. Ritchie and G.W Smith. 2003. African and Asian Dust: From Desert Soils to Coral Reefs. BioScience
53: 469-480.







nsburg, R.N., R.P.M. Bak, WE. Kiene, E. Gischler and V. Kosmynin. 1996. Rapid assessment of reef condition using
ral vitality. Reef Encounter 19: 12-24.

adfelter W.B., E.H. Gladfelter, R.K. Monahan, J.C. Ogden and R.D. Dill. 1977. Environmental studies of Buck Island
eef National Monument, St. Croix, US Virgin Islands. National Park Service. 140 pp.

)rdon, S. 2002. USVI queen conch assessment. Final Report to the Southeast Area Monitoring and Assessment Pro-
am-Caribbean. Division of Fish and Wildlife, Department of Planning and Natural Resources, U.S. Virgin Islands. 65


)rdon, S. and K.R. Uwate. 2003. 2002 opinion survey of U.S. Virgin Islands commercial fishers and the marine recre-
onal industry. Division of Fish and Wildlife, Department of Planning and Natural Resources, US Virgin Islands. 31 pp.

)rdon, S. and J.A. Vasques. In press. Spatial and temporal variation in postlarval settlement of the spiny lobster, Panu-
is argus, between 1992 and 2003 with the Cas Cay/Mangrove Lagoon and Great St. James Marine Reserves, St.
lomas. Gulf Caribbean Fisheries Institute.

Ilis-Starr, Z. 2004. U.S. National Park Service, St. Croix, US Virgin Islands. Personal communication.

ibbard, D. K., J. D. Stump and B. Carter. 1987. Sedimentation and reef development in Hawksnest, Fish, and Reef
iys, St. John, U.S. Virgin Islands. Virgin Islands National Park/Biosphere Reserve, Virgin Islands Resource Manage-
ent Cooperative, National Park Service. Biosphere Reserve Research Report 21. 99 pp.

ibbard, D.K., K.M. Parsons, J.C. Bythell and N.D. Walker. 1991. The effects of Hurricane Hugo on the reefs and as-
ciated environments of St Croix, US Virgin Islands a preliminary assessment. Journal of Coastal Research Special
sue 8: 33-48.

ibbard, D.K., E.H. Gladfelter and J.C. Bythell. 1993. Comparison of biological and geological perspectives of coral-reef
immunity structure at Buck Island, U.S. Virgin Islands. In: R.N. Ginsburg (ed.) Proceedings of the global aspects of coral
efs: health hazards, and history. Rosentiel School of Marine and Atmospheric Sciences, University of Miami.

Imann, P. and N. Deloach. 2002. Reef Fish Identification: Florida Caribbean Bahamas, 3rd edition. New World Publica-
ns, Inc., Jacksonville, FL. 481 pp.

F (Island Resource Foundation). 1996. Tourism and coastal resources degradation in the wider Caribbean. Island Re-
urces Foundation, Washington, DC. 53 pp. Available from the internet URL: http://www.irf.org/irtourdg.html.

F (Island Resource Foundation). 1999. Abbreviated 1998 water quality assessment of the United States Virgin Islands.
ibmitted to the U.S. Environmental Protection Agency. Division of Environmental Protection, Department of Planning
d Natural Resources (DPNR), Government of the Virgin Islands (USVI) of the United States. 101 pp.

rrell, J.D., M. Mayfield and E.N. Rappaport. 2001. The deadliest, costliest, and most intense United States hurricanes
im 1900 to 2000 (and other frequently requested hurricane facts). NOAAAOML Hurricane Research Division, Miami.
3p.

innings, C.A. 1992. Survey of non-charter boat recreational fishing in the U.S. Virgin Islands. Bulletin of Marine Science
i (2): 342-351.

,ndall, M.S., C.R. Kruer, K.R. Buja, J.D. Christensen, M. Finkbeiner, R.A. Warner and M.E. Monaco. 2001. Methods
ed to map the benthic habitats of Puerto Rico and the U.S. Virgin Islands. NOAA, NOS, NCCOS. Silver Spring MD.
i pp.







)bertson and J.D. Cubit. 1984. Spread of Diadema mass mortality through tl


Levitan, D.R. 1988. Algal-urchin biomass responses following the mass mortality of the sea urchin Diadema antillarum
Philippi at St. John, U.S. Virgin Islands. Journal of Experimental Marine Biology and Ecology 119: 167-178.

Link, E.G. 1997. Information transfer between managers and boaters and compliance with mooring and anchoring regu-
lations in the Virgin Islands National Park. Masters thesis for the University of Rhode Island. 138 pp.

Mateo, I. 2000. Final Report, Recreational Fishery Assessment Project. Division of Fish and Wildlife, Department of Plan-
ning and Natural Resources, U.S. Virgin Islands. 52 pp.

Mateo, I. and W.J. Tobias. 2002. Preliminary estimations of growth, mortality, and yield per recruit for the spiny lobster
Panulirus argus. pp. 58-75. In: St. Croix, USVI. Proceedings of the Gulf and Caribbean Fisheries Institute 53.

Miller, J., C. Rogers and R. Waara. 2003. Monitoring the coral disease, plague type II, on coral reefs in St. John, U.S.,
Virgin Islands. Revista de biologia tropical 51: 47-55.

Monaco, M.E., J.D. Christensen and S.O. Rohmann. 2001. Mapping and monitoring of U.S. coral reef ecosystems-the
coupling of ecology, remote sensing and GIS technology. Earth System Monitor 12 (1): 1-16

Monaco, M.E., A.M. Friedlander, C. Caldow, J.D. Christensen, J. Beets, J. Miller and C.S. Rogers. In preparation. A Pre-
liminary Assessment of Virgin Islands Coral Reef National Monument: Comparison of Fish Populations and Habitat Inside
and Outside of a Marine Protected Area.

National Resources Defense Council. 2004. Testing the waters: a guide to water quality at vacation beaches. Available
from the internet URL: http://www.nrdc.org/water/oceans/ttw/sumvi.pdf.

Nemeth, R.S. In review. Recovery of a US Virgin Islands red hind spawning aggregation following protection. Marine
Ecology Progress Series.

Nemeth, R.S. and J. Sladek-Nowlis. 2001. Monitoring the effects of land development on the near-shore reef environ-
ment of St. Thomas, USVI. Bulletin of Marine Science 69: 759-775.

Nemeth, R.S., S. Herzlieb and M. Taylor. 2002. Coral reef monitoring in St. Croix and St. Thomas, United States Virgin
Islands. Year 1 final report. Division of Fish and Wildlife, Department of Planning and Natural Resources. US Virgin Is-
lands. 33 pp.

Nemeth, R.S., L. Whaylen and C. Pattengill-Semmens. 2003a. A rapid assessment of coral reefs in the Virgin Islands
(Part 1: stony corals and algae). Atoll Resesearch Bulletin 496: 544-566.

Nemeth, R.S., L.D. Whaylen and C.V. Pattengill-Semmens. 2003b. A rapid assessment of coral reefs in the Virgin Islands
(Part 2: fishes). Atoll Resesearch Bulletin 496: 567-590.

Nemeth, R.S., S. Herzlieb, M. Taylor, S. Herald and W Toller. 2003c. Coral reef monitoring in St. Croix and St. Thomas,
United States Virgin Islands. Year 2 final report, submitted to Department of Planning and Natural Resources.

Nemeth, R.S., S. Herzlieb, E.S. Kadison, M. Taylor, P. Orthenberger and S. Harold. 2004. Coral reef monitoring in St.
Croix and St. Thomas, United States Virgin Islands. Year 3 final report. Division of Fish and Wildlife, Department of Plan-
ning and Natural Resources, US Virgin Islands. 74 pp.

NOAA (National Oceanic and Atmospheric Administration). 2001. National Ocean Service, National Centers for Coastal
Ocean Science, Biogeography Program. Benthic Habitats of Puerto Rico and the U.S. Virgin Islands. CD-ROM. Silver
Spring, MD.

Olsen, D.A. and J.A. LaPlace. 1978. A study of a Virgin Islands grouper fishery based on a breeding aggregation. Pro-
ceedings of the 31st Gulf and Caribbean Fisheries Institute 31: 130-144.

Olsen, D.A., D.W Nellis and R.S. Wood. 1984. Ciguatera in the eastern Caribbean. Marine Fisheries Review 46: 13-18.

Randall, J.E. 1967. Food habits of reef fishes of the West Indies. Studies in Tropical Oceanography 5: 665-847.

Randall, J.E. 1996. Caribbean Reef Fishes, 3rd edition. T.F.H. Publications, Inc., Neptune City, NJ. 368 pp.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


Rogers, C.S., T Suchanek and F. Pecora. 1982. Effects of Hurricanes David and Frederic (1979) on shallow Acropora
palmata reef communities: St Croix, USVI. Bulletin of Marine Science 32: 532-548.

Rogers, C.S., L. McLain and C. Tobias. 1991. Effects of Hurricane Hugo (1989) on a coral reef in St John, USVI. Marine
Ecology Progress Series 78: 189 199.

Rogers, C.S., V. Garrison and R. Grober-Dunsmore. 1997. A fishy story about hurricanes and herbivory: seven years of
research on a reef in St John, US Virgin Islands. pp. 555-560. In: H. A. Lessios and I. G. Macintyre (eds.) Proceedings of
the 8th International Coral Reef Symposium. Smithsonian Tropical Research Institute, Balboa, Panama.

Rogers, C.S. and J. Beets. 2001. Degradation of marine ecosystems and decline of fishery resources in marine protected
areas in the US Virgin Islands. Environmental Conservation 28: 312-322.

Rogers, C.S. and V.H. Garrison. 2001. Ten years after the crime: lasting effects of damage from a cruise ship anchor on
a coral reef in St. John, US. Virgin Islands. Bulletin of Marine Science 69: 793-803.

Rogers, C.S and J. Miller. 2001. Coral bleaching, hurricane damage, and benthic cover on coral reefs in St. John, U.S.
Virgin Islands: a comparison of surveys with the chain transect method and videography. Bulletin of Marine Science 69:
459 470.

Rogers, C.S., W. Gladfelter, D. Hubbard, E. Gladfelter, J. Bythell, R. Dunsmore, C. Loomis, B. Devine, Z. Hillis-Starr and
B. Phillips. 2002. Acropora in the U.S. Virgin Islands: A Wake or an Awakening? A Status Report Prepared for the National
Oceanographic and Atmospheric Administration. pp. 99- 122. In: A.W. Bruckner, A. (ed.) NOAA Technical Memorandum
NMFS-OPR-24. National Oceanic and Atmospheric Administration, Silver Spring, MD.

Rosario, A. 1995. Queen conch stratification survey. SEAMAP-Caribbean Program. CFMC/NMFS. 33 pp.

Salm, R.V., J. Clark and E. Sirila. 2000. Marine and Coastal Protected Areas: A Guide for Planners and Managers. Wash-
ington, DC: IUCN-The World Conservation Union. 371 pp.

Swingle, WE., A.E. Dammann and J.A. Yntema. 1979. Survey of the commercial fishery of the Virgin Islands of the
United States. pp. 22: 110-121. In: Proceedings of the 22nd Gulf and Caribbean Fisheries Institute.

Sylvester, J.R. and A.E. Dammann. 1972. Pot fishing in the Virgin Islands. Commercial Fisheries Review 34: 33-35.

Tobias, W USVI Dept of Planning and Natural Resources. St. Croix, USVI. Personal communication.

Tobias, W. 2000. U.S. Virgin Islands/National Marine Fisheries Service inter-jurisdictional fisheries program final progress
report, 1 October 1997 30 September, 2000. 25 pp.

Tobias, W 2001. Fish attracting device (FAD) construction, placement, monitoring and maintenance. Final Report to
U.S. Fish and Wildlife Service Sport Fish Restoration Program. Division of Fish and Wildlife, Department of Planning and
Natural Resources, U.S. Virgin Islands. 35 pp.

Tobias, W, R. Gomez, I. Mateo and B. Kojis. 2000. Three Year Summary Report, 1 April 1997-31 March 1999, Coopera-
tive Statistics Program NA77FT0093. USVI Department of Planning and Natural Resources, Division of Fish and Wildlife,
Bureau of Fisheries, 40 pp.

Tobias, W, W Toller, H. Rivera and W Ventura. 2002. SEAMAP-C USVI St. Croix fisheries independent trap and line sur-
vey. Summary report: Caribbean/NMFS Cooperative SEAMAP Program NA07FS0100-01. Division of Fish and Wildlife,
Department of Planning and Natural Resources, U.S. Virgin Islands. 27 pp.

Tobias, W and W Toller. 2004. Netfishing overview-St. Croix, U.S. Virgin Islands. Management implications for restric-
tions on the use of gill and trammel nets. Report to the Commissioner of Department of Planning and Natural Resources,
Division of Fish and Wildlife, US Virgin Islands. 21 pp.

U.S. Census Bureau (USCB). 2001. Census 2000 population counts forthe U.S. Virgin Islands. USCB, Washington, DC.
Available from the internet URL: http://www.census.gov/Press-Release/www/2001/cb01cnl72.html.

U.S. Census Bureau. 2003. Population and Housing Profile: 2000: 2000 census of population and housing, U.S. Virgin
Islands. Washington, DC. 219 pp. Available from the internet URL: http://www.census.gov/prod/cen2000/island/Vlprofile.
pdf.

USVI (United States Virgin Islands) Bureau of Economic Research. 2004. U.S. Virgin Islands annual tourism indicators.
Charlotte Amalie, St. Thomas, USVI.




The State of Coral Reef Ecosystems of the U.S. Virgin Islands


USVI DPNR (United States Virgin Islands. Department of Planning and Natural Resources). 2004. The 2004 Integrated
Water Quality Monitoring and Assessment Report for the United States Virgin Islands. DPNR, Division of Environmental
Protection, U.S. Virgin Islands. 193 pp.

UVI-ECC (United States Virgin Islands. Eastern Caribbean Center). 2002. Telephone survey of boat-based marine recre-
ational fishing in the U.S. Virgin Islands, 2000. University of the Virgin Islands, ECC. Department of Planning and Natural
Resources, Division of Fish and Wildlife, U.S. Virgin Islands. 56 pp.

Uwate, K.R., W. Tobias, P. Nieves, H. Rivera, W Ventura and L. Critchley. 2001. Survey of U.S. Virgin Island commercial
fishery opinions and usage of New National Monument areas (Buck Island and South of St. John). Department of Plan-
ning and Natural Reosurces, Division of Fish and Wildlife, Bureau of Fisheries, U.S. Virgin Islands. 13 pp.

Valle-Esquivel, M. 2002. U.S. Caribbean queen conch (Strombus gigas) data update with emphasis on the commercial
landings statistics. University of Miami, Sustainable Fisheries Division Contribution SFD-01/02-169.

Weil, E. and G. Smith. 2003. Status, variability, and extended host and geographic range of Caribbean coral diseases in
Bermuda. In: Proceedings, 7th International Conference on Coelenterate Biology. University of Kansas, Lawrence.

Wolff, N. 1996. The Fish Assemblages within Four Habitats found in the Nearshore Waters of St. John, USVI: With some
Insights into the Nature of Trap Fishing. M.S. Thesis for the University of Rhode Island. 207 pp.

Wolff, N. 1998. Spiny lobster evaluation within Virgin Islands National Park (Summer of 1996). Report to U.S. Geological
Survey (USGS), Biological Resources Division, Washington DC. 16 pp.

Wood, D. and D. Olsen. 1984. Application of biological knowledge to the management of the Virgin Islands conch fishery.
pp: 112-121. In: Proceedings of the 35th Gulf and Caribbean Fisheries Institute.




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