United States
Environmental Protection
Agency
EPA/600/R-11/206 | January 2012 | www.epa.gov/research
Quantifying Coral Reef
Ecosystem Services
I Off ice of Research and Development
I National Exposure Research Laboratory
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EPA/600/R-11 /206 | January 2012 www.epa.gov/research
Quantifying Coral Reef
Ecosystem Services
Peter P. Principe
Environmental Sciences Division
National Exposure Research Laboratory
Research Triangle Park, NC
Patricia Bradley
Atlantic Ecology Division
National Health and Environmental Effects Research Laboratory
Key West, FL
Susan H. Yee
William S. Fisher
Eric D. Johnson
Gulf Ecology Division
National Health and Environmental Effects Research Laboratory
Gulf Breeze, FL
Paula E. Allen
Environmental Sciences Division
National Exposure Research Laboratory
Las Vegas, NV
Daniel E. Campbell
Atlantic Ecology Division
National Health and Environmental Effects Research Laboratory
Narragansett, Rl
National Exposure Research Laboratory
Office of Research and Development
U.S. Environmental Protection Agency
Research Triangle Park, NC 27711
RecyciedlRecyclable
Printed with vegetable-based ink on paper thai
contains a minimum of 50% pnst-rtnnsumer
fiber and is processed chlorine free.
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Notice and Disclaimer
The U.S. Environmental Protection Agency through its Office of Research and Development funded the research described
herein. It has been subjected to the Agency's peer and administrative review and has been approved for publication as an EPA
document. Mention of trade names or commercial products does not constitute endorsement or recommendation for use.
This is a contribution to the EPA Office of Research and Development's Ecosystem Services Research Program, Coral Reefs
Project.
The appropriate citation for this report is:
Principe P, Bradley P, Yee S, Fisher W, Johnson E, Allen P and Campbell D. 2011. Quantifying Coral Reef Ecosystem
Services. U.S. Environmental Protection Agency, Office of Research and Development, Research Triangle Park, NC.
EPA/600/R-11/206, January 2012.
This document can be downloaded from EPA's website: www.epa.gov/research
Acknowledgements
This report was peer reviewed by Steve Jordan and Debbie Santavy of ORD's Gulf Ecology Division and Bob Leeworthy of
the National Oceanic and Atmospheric Administration.
Cover photo credit: Charles LoBue, EPA Region 2.
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Table of Contents
List of Figures vi
List of Tables vii
1 Coral Reefs: Introduction and Overview 1
1.1 Quantifying coral reef ecosystem services 1
1.2 Purpose and organization 5
1.3 Ecosystem services and economic benefits 5
1.4 Ecological integrity, ecological resilience, and biodiversity 8
1.4.1 Ecological integrity 8
1.4.2 Ecological resilience 9
1.4.3 Biodiversity 9
1.4.4 Insurance value 9
1.5 References 10
2 Coral Reefs: Tourism and Recreation 15
2.1 Describing the service 15
2.1.1 Definitions 15
2.1.2 Tourism and the economy 15
2.1.3 Tourism and recreation activities related to coral reefs 16
2.1.3.1 Recreational (scuba) diving 16
2.1.3.2 Sn orke ling 17
2.1.3.3 Underwater photography 17
2.1.3.4 Recreational (sport) fishing 18
2.1.3.5 Viewing nature and wildlife 18
2.1.3.6 Beach sunbathing and swimming 19
2.1.3.7 Collecting objects (beachcombing) 19
2.1.3.8 Surfing 19
2.1.4 Tourism and recreation: businesses 20
2.2 Necessary conditions for providing the service 20
2.2.1 Linkages among reef condition, reef structures, and
reef functions with respect to delivery of the service 21
2.3 Measuring the service 22
2.3.1 Stony corals (scleractinians) 23
2.3.2 Octocorals (gorgonians) 23
2.3.3 Sponges 23
2.3.4 Fishes 23
2.3.5 Mangroves and seagrasses 24
2.4 Valuing the service 24
2.4.1 Effect on production (EoP) 25
2.4.2 Financial analysis (FA) 25
2.4.3 Travel costs (TC) 26
2.4.4 Contingent valuation method (CVM) 26
2.4.5 National scale valuation of tourism 26
2.4.6 National scale valuation of recreational fishing 27
2.4.7 Noneconomic human dimensions measures 27
2.4.7.1 Importance-satisfaction ratings 28
2.4.7.2 Expectancy-discrepancy analysis 28
2.4.7.3 Norm curves 28
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2.4.7.4 Entergy 28
2.5 Reflections 28
2.6 Bibliography 31
Appendix 2-A: Studies quantifying tourism and recreation 39
3 Coral Reefs: Fish & Coral for Food, Aquaria,
& Ornaments 43
3.1 Describing the service 43
3.1.1 Definitions 43
3.1.2 Benefits of coral reef fisheries 43
3.1.2.1 Seafood 43
3.1.2.2 Live fish and coral for aquaria 44
3.1.2.3 Ornaments and jewelry 44
3.2 Providing the service 44
3.2.1 Linkages among reef condition, reef structures, and reef functions
with respect to delivery of fishes for aquaria and food 44
3.2.1.1 Scleractinian corals 44
3.2.1.2 Reef structural complexity 45
3.2.1.3 Seascape connectivity 45
3.2.2 Linkages among reef condition, reef structures, and reef
functions with respect to provision of stony corals, black
corals, and precious corals for aquaria stock and jewelry 46
3.3 Measuring the service 47
3.3.1 Fish 47
3.3.2 Stony corals 47
3.3.3 Reef structural complexity 47
3.3.4 Seascape mapping 47
3.3.5 Fisheries modeling 48
3.4 Valuing the service 48
3.5 Reflections 51
3.5.1 Threats to coral reef fisheries 51
3.5.2 Management options 51
3.5.2.1 Essential fish habitat (EFH) 51
3.5.2.2 Marine protected areas (MPAs) 51
3.5.3 Conclusions 52
3.6 Bibliography 53
4 Coral Reefs: Shoreline Protection 59
4.1 Defining the service 59
4.2 Providing the service 60
4.2.1 Presence of the reef 60
4.2.2 Reef attributes 60
4.2.3 Reef health 62
4.3 Measuring the service 62
4.3.1 Physical processes 62
4.3.2 Biological processes 62
4.3.3 Ecosystem service measures 63
4.3.4 Socioeconomic benefits 64
4.4 Valuing the service 64
4.5 Reflections 65
4.5.1 Improving current knowledge 65
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4.5.2 Connecting biophysical processes and ecosystem services
to socioeconomic benefits 66
4.6 Bibliography 69
Appendix 4-A: Studies quantifying shoreline protection 73
5 Coral Reefs: Natural Products 77
5.1 Natural products as sources and templates for Pharmaceuticals,
biochemicals, and biomaterials 77
5.1.1 Pharmaceutical uses of marine natural products 77
5.1.2 Marine natural products as structural templates for synthesis 83
5.1.3 Marine natural products as molecular probes 83
5.1.4 Nonpharmaceutical uses of marine natural products 84
5.2 Sustaining the presence of natural products in coral reef
ecosystems 85
5.2.1 Ecological role and sources of secondary metabolites 86
5.3 Measuring and valuing the service 88
5.3.1 Specific examples 89
5.3.2 Possible approach 89
5.4 Reflections 90
5.5 Bibliography 90
Appendix 5-A: Marine pharmaceutical products 95
6 Coral Reefs: Summary 133
6.1 Lessons learned 133
6.2 Next steps 134
Coral Reefs: Glossary 137
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List of Figures
Chapter 1
Figure 1-1 U.S. jurisdictions with coral reefs 3
a. The Atlantic Ocean, the Caribbean Sea, and the Gulf of Mexico
b. The Pacific Ocean
Figure 1-2 Conceptual model of a coral reef ecosystem illustrating local effects of driving
forces (urban development, industrial production, fishing, tourism),
pressures (industrial effluent, sedimentation and erosion, over fishing, groundings),
state (healthy coral reef on the left and degraded coral reef on the right), and
impacts (loss offish and coral species, conversion of reef habitat to rubble) 5
Figure 1 -3 Ecosystems provide services that, in turn, provide economic benefits 6
Figure 1-4 Ecological integrity as the overlap of physical, chemical, and biological integrity 8
Chapter 2
Figure 2-1 Annual diver certifications by the Professional Association of Dive
Instructors (PADI) 17
Chapter 4
Figure 4-1 Illustration of wave set-up and attenuation over a reef. 61
Figure 4-2 Relationship of wave energy with increasing reef depth and width and
with increasing reef friction 61
Figure 4-3 Reef influences on shorelines: salient and tombolo 64
Figure 4-4 Conceptual diagram showing how reef attributes, and various physical or
socioeconomic variables, can be used to determine various endpoints
quantifying shoreline protection 68
Chapter 5
Figure 5-1 Phyletic distribution of 160 reviewed marine species from which potential
antifouling natural products have been extracted 85
Figure 5-2 Marine phyla contributing secondary metabolites with pharmaceutical potential 88
Figure 5-3 Sources of 20 marine anticancer drugs in clinical trials or recently in clinical
trials: reported versus predicted based on subsequent metagenomic analysis 88
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List of Tables
Chapter 1
Table 1-1 Economic benefits provided by coral reefs 1
Table 1-2 Market and nonmarket value estimates for selected ecosystem services at local,
state/regional, and national/international levels 2
Table 1-3 Areal extent estimates for potential shallow water coral reefs inside 10 fathom
(18m) and 100 fathom (183 m) depths and local populations that benefit from
coral reefs 4
Table 1-4 Examples of the variety of ecosystem services valued by previous coral
reef studies 7
Table 1-5 Estimated annual economic impact from coral reef ecosystem services
at global, regional, national, and local levels 7
Chapter 2
Table 2-1 Economic impact of travel and tourism (2009 preliminary data) 15
Table 2-2 Features relevant to the perceived value of coral reefs 22
Table 2-3 Stony coral rapid bioassessment protocol coral condition indicators 24
Table 2-4 Methods used for valuing goods and services of coral reef ecosystems 25
Table 2-5 Final ecosystem services and supporting features for tourism and recreation 30
Chapter 3
Table 3-1 Benefits or amenities derived from fish production 43
Table 3-2 Important habitats for juveniles and adults of selected fish species 46
Table 3-3 Valuation methods 49
Table 3-4 Categories of values 50
Table 3-5 Final ecosystem services and supporting features for fish production 53
Chapter 4
Table 4-1 Measures that have been used to quantify shoreline protection 59
Table 4-2 Reef attributes that contribute to wave attenuation 60
Table 4-3 Measures used to quantify shoreline protection, and the major reef attributes,
physical parameters, and socioeconomic parameters used to estimate that
protection's value 63
Table 4-4 Estimated values of shoreline protection 65
Table 4-5 Final ecosystem services and supporting features for shoreline protection 67
Chapter 5
Table 5-1 Testing stages for new pharmaceuticals 78
Table 5-2 Status of marine-derived natural products in clinical trials or clinical use 79
Table 5-3 Number of candidate drugs reaching different stages of clinical trials 81
Table 5-4 Timeline of drug development and amount of raw material and pure product
needed at each stage 82
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Table 5-5 Marine biochemicals used as molecular probes 83
Table 5-6 Current and potential nonpharmaceutical uses for marine natural products 84
Table 5-7 Percentage of secondary metabolites with pharmaceutical potential from
marine phyla 88
Table 5-8 Market value of pharmaceutical products 89
Table 5-9 Final ecosystem services and supporting features for natural products 90
Chapter 6
Table 6-1 Combined final ecosystem services and supporting features for coral reefs 134
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1
Coral Reefs: Introduction and Overview
1.1 Quantifying coral reef ecosystem services
Coral reefs have been declining during the last four decades as a result of both local and global anthropogenic stresses (MEA
2005). In response, governments, nongovernmental organizations (NGOs), and academia initiated numerous research efforts to
elucidate the nature, causes, magnitude, and potential remedies for the decline. This work has led to the widely-held belief that
the recovery of coral reefs is unlikely if public and private sector decisions that affect coral reefs continue to ignore the
economic value of the goods and services provided by these ecosystems (MEA 2005). If this perception is correct, successful
conservation will, in most instances, require that environmental benefits (or, ecosystem services) are routinely included in
economic and social decisions (Pearce & Turner 1990; Pearce & Moran 1994; Daily et al. 1997; Turner et al. 2003; Chee 2004;
Boyd & Banzhaf 2007; Turner 2010). However, this approach presupposes knowledge of what the ecosystem services are, what
their magnitudes and values are, what ecosystem characteristics provide them, how those characteristics are affected by human
activities, and how human activities may affect the future provision of ecosystem services. With this knowledge,
decisionmakers could have access to a more complete characterization of the consequences of different policy options.
Table 1-1. Economic benefits provided by coral reefs
A. Direct extractive uses
1. Commercial fishing
2. Subsistence fishing
3. Aquarium fish
4. Sport fishing
5. Coral jewelry
6. Pharmaceutical harvesting
7. Nonpharmaceutical harvesting
C. Indirect uses
1. Fish habitat
2. Nutrients
3. Reduced flooding
4. Less storm damage
5. Fewer deaths from storms & flooding
6. Reduced erosion from storms & flooding
7. Mangrove & seagrass protection
8. Sealife nursery protection
9. Global life support
B. Direct nonextractive uses
1. Scuba diving
2. Snorkeling
3. Boating
4. Pharmaceutical chemicals
5. Nonpharmaceutical natural products
D. Nonuse values
1. Existence value
2. Cultural value
3. Option value
4. Quasi-option value
5. Bequest value
6. Instrumental value
7. Intrinsic value
8. Scientific value
9. Scarcity value
Source: Beaumont et al. 2008; Burke et al. 2008; Cesar 2002; Cesar & Chong 2005; David et al. 2007;
Ghermandi et al. 2009; Moberg & Folke 1999; Naber et al. 2008; Nunes et al. 2009;
Remoundou etal. 2009; Spurgeon 1992.
The ecosystem services in Table 1-1 are organized in categories normally used for valuation purposes. The services can also be
organized according to their ecological role, an approach used in the Millennium Ecosystem Assessment (MEA 2005). The
MEA used four categories: provisioning, regulating, cultural, and supporting. Coral reefs, with high species diversity and
topographically complex habitat, are valued in all categories (McKinney 1998). Provisioning services include food (fish and
invertebrates), materials for construction (sand and coral blocks), pharmaceutical and cosmetic compounds (bio-mining),
jewelry, curios, and ornamental fish. Regulating services include land accretion and shoreline protection from waves and
currents (which allows growth of seagrass and mangrove communities), and carbon sequestration. Major cultural services
include recreation and tourism; in addition, coral reefs provide coastal communities an inherent sense of place. Sometimes less
obvious are the supporting services of coral reefs, such as sand for beach formation, biological primary and secondary
production, and biological diversity.
The economic value of some of these services (e.g., commercial fishing) are established in markets, while others have
nonmarket values for local, state/regional, and national/international segments of the population (Table 1-2).
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Table 1-2. Market and nonmarket value estimates for selected ecosystem
services at local, state/regional, and national/international levels
[small (+), moderate (+ +), or high (+ + +)]
Ecosystem Service*
Tourism
Recreation
Commercial Fisheries
Shoreline Protection
Non-Use Existence Value
Biodiversity
Sector Local State National
Market +++ + +
Nonmarket + +++ +++
Market + + +
Nonmarket ++ — —
Market + ++ ++
Nonmarket — — —
Market — — —
Nonmarket ++ ++ ++
Market — — —
Nonmarket ++ ++ ++
Market ++ + +
Nonmarket + +++ +++
* Tourism and recreation include snorkeling, diving, fishing, and viewing in reef areas; commercial fisheries
include spiny lobsters, shrimp, fmfish, and aquarium fish; shoreline protection is based on damage avoided
from storms; nonuse existence value is based on
what people are willing to pay to conserve coral reefs; biodiversity has both market value (e.g., potential
pharmaceutical products) and nonmarket value (existence value).
Source: EPA (2009)
An estimated 173,488 km2 of coral reefs exist in U.S. jurisdictions widely distributed across the Pacific Ocean, western Atlantic
Ocean, Caribbean Sea and Gulf of Mexico (Figure 1-1, Table 1-3) (Rohmann etal. 2005; Waddell 2005, p. 8). This is roughly
the areal extent of the State of Florida. Nearly 12 million people in these jurisdictions directly benefit from shoreline protection,
recreation, subsistence fishing, and sense of place provided by reefs. Some of them owe their livelihood to the services
provided by coral reefs. Others worldwide benefit from tourism, commercial fishery harvests, pharmaceutical products,
jewelry, and non-use services such as natural beauty and biodiversity. Many of these beneficiaries contribute to the success of
non-governmental organizations (e.g., The Nature Conservancy (TNC), Coral Reef Alliance, Reef Relief, Reef Environmental
Education Foundation (REEF), and the World Wildlife Fund (WWF)) in raising money for protection of coral reefs, and in
supporting actions by the U.S. Government to establish and fund the interagency Coral Reef Task Force, National Marine
Sanctuaries, National Parks, local action strategies, and legislation (e.g., Coral Reef Conservation Act) for protection of coral
reefs.
There is an urgent need to manage coral reefs and coastal zones differently. Over the past two decades, there have been
precipitous declines in coral reefs (Waddell & Clark 2008), despite their widely held status as highly valued and highly
important ecosystems. An estimated 19% of the world's reefs have been lost in the last 30 years, 15% are threatened with loss
by 2030, and another 20% with loss by 2050 (Wilkinson 2008). Many of the causes of this decline are well known. Stony corals
(Scleractinia spp.), which provide the calcareous reef infrastructure, are suffering worldwide from massive bleaching (resulting
from the loss of symbiotic algas) and higher than normal mortality. The most common trigger for bleaching events is elevated
sea surface temperatures. Also at the global level, increasing seawater acidity from elevated atmospheric CO2 is slowing the
calcification processes responsible for coral growth and repair. At local scales (Figure 1-2), corals are under chronic assault
from fishing and land use practices that release sediments, nutrients, toxic contaminants, and potentially pathogenic
microorganisms into the reef environment.
The loss or impairment of coral reefs directly results in the reduction or loss of ecosystem services provided by the reefs to
humans. When reef-building stony corals die, they become covered in algas and other organisms that eventually erode the
remnant skeletons. Unfortunately, coral reefs are among the few marine environments to exhibit disturbance-induced phase
shifts (akin to "tipping points") in which lush, complex coral communities dominated by reef-building stony corals are
transformed relatively quickly into "biologically impoverished wastelands overgrown with algas" (Bellwood et al. 2004; Work
et al. 2008). As coral skeletons erode and crumble, reefs lose the highly complex ecological and physical architectures that
provide shelter and resources for the uniquely abundant and diverse reef biota. The loss of the reefs' physical structure (such as
that currently being experienced in the Caribbean) is referred to as "flattening" (i.e., losing topographic complexity) (Alvarez-
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Filipi et al. 2009). This term would also be appropriate to describe the loss of ecological structure and biodiversity that make
these ecosystems so valuable to humans. Inevitably, these losses will adversely affect human well-being.
Figure 1-la. U.S. jurisdictions with coral reefs. The Atlantic Ocean, the Caribbean Sea,
and the Gulf of Mexico, (figure from Waddell & Clarke 2008, p. 10)
Jff
COMMONWEALTH
OF THE NORTHERN
MARIANAS ISLANDS
^s
FEDERATED STATES
OF MICRONESIA
MARSHALL
ISLANDS
_
~~~
V
500 1.000 km
Figure 1-lb. U.S. jurisdictions with coral reefs. The Pacific Ocean. (Waddell & Clarke 2008, p. 12)
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Table 1-3. Areal extent estimates for potential shallow water coral reefs inside 10 fathom
(18 m) and 100 fathom (183 m) depths and local populations that benefit from
coral reefs
U.S. Jurisdiction
<1(
Florida13
Puerto Ricod
U.S. Virgin Islandsf
Navassa Island
Flower Gardens NMS8
Main Hawaiian Islands'1
Northwestern Hawaiian Islands (NWHI)1
American Samoak
Pacific Remote Islands1
Northern Mariana Islands (CNMI)111
Guam
Marshall Islands
Federated States of Micronesia
Palau
Total
Reef Area (km2)
) fathoms3
30,801
2,302
344
3
0
1,231
1,595
55
252
124
108
13,456n
14,517"
2,529n
67,317
< 100 fathoms"
113,092
5,501
2,126
14
164
6,666
13,771
464
436
476
276
13,456n
14,517"
2 529n
173,488
Population
5,213,884C
3,971,020e
109,825e
0
0
1,283,388'
0
65,628e
100e
86,6 16e
178,430e
64,522e
107,434e
20,796e
11,774,315
a Areas estimated from NOAA nautical charts except for the Marshall Islands, the Federated States of Micronesia, and
the Republic of Palau, which were estimated from Landsat satellite imagery of shallow waters (<15 m).
b Florida corals extend along the Atlantic Ocean coast of Florida to Jupiter Inlet, Florida and along the Gulf of Mexico
coast of Florida to Tarpon Springs, Florida.
c 2007 estimate for five coastal counties: St. Lucie, Martin, Palm Beach, Broward, Miami-Dade and Monroe counties.
Total population of Florida is 18,251,243 (Census Bureau 2010).
d Puerto Rico includes the islands of Puerto Rico, Desecheo, Culebra, Vieques, and Mona.
e Projected 2009 population (CIA 2010)
f The U.S. Virgin Islands includes the islands of St Thomas, St John, and St Croix.
g The NOAA nautical chart depicts only the 100 fathom depth curve for this location.
h The Main Hawaiian Islands includes the islands of Hawaii, Maui, Molokai, Lanai, Kahoolawe, Oahu, Kauai, and
Niihau.
i 2007 estimate (Census Bureau 2010)
j The NWHI 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.
k American Samoa includes the islands of Tutuila, Ofu, Olosega, Tau, Swains, and Rose Atoll.
1 The Pacific Remote Islands include Howland, Baker, and Jarvis Islands, Palmyra, Johnston, and Wake Atolls, and
Kingman Reef.
m 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.
n 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.
Source: Rohmann etal. (2005); Waddell (2005, p. 8).
The Ecosystem Research Program (ESRP) is a concerted research effort within the U.S. Environmental Protection Agency
(EPA) Office of Research and Development (ORD) to better incorporate consideration of ecosystem services in decision
processes. The overall goal of the ESRP is to provide decision-makers information on the nature and magnitude of
environmental benefits provided by ecosystems and the means to understand how policy or management choices will affect the
provision of those benefits (i.e., the nature and magnitude of the economic benefits [see Table 1-1 for a list of benefits and
Table 1-2 for their relative magnitudes]).
Completion of research in the Coral Reef Project (CRP) is expected to advance the understanding of coral reef ecosystem
services, how they are affected by human activities, and how management and policy decisions influence their delivery. If this
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information is incorporated into public and private decision-making and the policy analyses that precede it, the resulting
decisions will undoubtedly have a firmer scientific basis and a more inclusive consideration of the benefits provided by coral
reef ecosystems. This report provides a review of previous studies of ecosystem services and economic benefits provided by
coral reefs and how those ecosystem services are linked to characteristics (attributes) of the reef.
Fishing:
loss of grazers
by-catch
Groundings
anchor damage
dive tourism
Loss and decline
of coral, fish
habitat, fish food
Loss of fish
Conversion
from reef to
simple hard
bottom
Figure 1-2. Conceptual model of a coral reef ecosystem illustrating local effects of driving forces (urban
development, industrial production, fishing, tourism), pressures (industrial effluent, sedimen-
tation and erosion, over fishing, groundings), state (healthy coral reef on the left and degraded
coral reef on the right), and impacts (loss of fish and coral species, conversion of reef habitat to
rubble). Adverse impacts reduce the natural benefits and value of coral reefs unless steps are
taken to reduce pressures, (figure and caption from Bradley et al. 2010)
1.2 Purpose and organization
The purpose of this report is to provide an overview of the published literature with respect to three aspects of ecosystem
services provided by coral reefs: (1) which services have been identified; (2) what, if any, methods were used to quantify the
services; and, (3) what connections were identified between the services and the attributes of the reef. The goal is to lay the
foundation for selecting research questions that, if answered, would fill gaps in our current understanding of the links between
coral reef attributes and the delivery of services by the coral reef ecosystem. Ultimately, the goal is to identify coral reef
indicators that can be used to estimate the quantity of ecosystem services being delivered by coral reef ecosystems and to
predict the extent to which the current quantities are likely to change in the future.
1.3 Ecosystem services and economic benefits
There exists a certain imprecision prevalent in the current use of the term "ecosystem services". Many authors tend to conflate
"ecosystem services" and "economic benefits", thereby creating a confusing logical muddle. In this report, ecosystem services
are viewed as an ecological construct that creates wholly, or partially, the economic construct of economic benefits, which is
the core of policymaking. Ecosystem services can always be quantified in physical terms (e.g., kg of fish, board feet of lumber),
but they are always linked to economic benefits, which are usually valued in monetary terms (e.g., market value of fish catch or
lumber). In the case of shoreline protection provided by coral reefs, the presence of the reef attenuates the energy contained in
waves reaching the shoreline, and that attenuation reduces the wave-induced erosion of the shoreline and, during storms, the
extent of near-shore flooding, damage to buildings and properly, and danger to humans and livestock. The shoreline protection
ecosystem service could be characterized as the amount of wave energy attenuation attributable to the presence of the reef, or in
the case of a foregone ecosystem service, the increase in wave energy attributable to a diminishment of the reef's size. The
economic benefits from shoreline protection would be the value of the properly, lives, and well-being preserved by the
attenuation of the wave energy, or the value lost due to the increase in wave energy. However, the value of the economic
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benefit is a function of how much erosion is prevented, how much property is protected, and how many human and livestock
lives are saved. Transforming energy attenuation into a physical quantification of economic benefits should be a joint exercise
for scientists and economists, with the former leading the conversion modeling and the latter leading the identification of the
economic benefits to be quantified, but the subsequent valuation of the benefits would be the bailiwick of economists. To
determine which ecosystem services are the most important for policymaking requires working backward from the economic
benefits to identify the services that created those benefits (Figure 1-3).
Analytical path of this document
Ecosystem
State &
Attributes
Ecosystem
Services
Transfor-
mation
Economic
Benefits
Valuation
Joint ecological-economic analysis
Figure 1-3. Ecosystems provide services that, in turn, provide economic benefits. Some services have a direct,
one-to-one correspondence to benefits, but more commonly services have to be transformed,
disaggregated, or combined to yield benefits. Services and many benefits can be quantified in physical
terms; the value of benefits is usually estimated in monetary terms; services are not valued per se, but
their importance can be inferred from the value of the benefits they create.
The journey traversed in creating this report, then, began by identifying the most valuable economic benefits arising from coral
reefs, followed by an assessment of which ecosystem services contribute to those benefits. Subsequently, the literature was
reviewed to find examples where these ecosystem services had been wholly or partially quantified and where the links between
these services and coral reef attributes were investigated.
Identification and quantification of coral reef ecosystem services began in the 1980s (Hodgson & Dixon 1988; Mattson &
DeFoor 1985; McAllister 1988; Munro & Williams 1985), followed in the early 1990s by the first unified estimates of the value
of the economic benefits generated by those services. Numerous studies have estimated quantities and values for specific
services and the derived benefits received from coral reefs, but fewer have done so for the whole range of coral reef services
and benefits (see Pendleton [2008] and Conservation International [2008] for reviews; also see Table 1-1 references). In most
cases, reefs are valued to determine or demonstrate the economic impact of a planned or implemented decision. The decision
under consideration plays a role in what is quantified and how the services are valued. For example, one early coral reef study
focused on the economic value of biodiversity, because the study area was an existing marine reserve established to protect
biodiversity (Bakus 1983); another study approached valuation more broadly because it was intended to reinforce the creation
of a Marine Protected Area in exchange for international debt reductions (David et al. 2007). The services valued can vary from
study to study (Table 1 -4), depending on the decision, the availability of information, and the expertise or insight of the authors.
Despite the wide variety of services that have been measured and valued, there are certain services that are almost always
considered, because they make very significant contributions to the total economic value (TEV includes all direct and indirect
use values plus all nonuse values). The values can be calculated at a global, regional, or local scale (Table 1-5).
The values reported in Table 1-5 suggest that the four most valuable economic benefits are: fisheries, natural products,
shoreline protection, and tourism and recreation. This report examines each of these in a separate chapter.
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Table 1-4. Examples of the variety of ecosystem services valued by previous coral reef studies
Study
Direct Use
Indirect Use
Nonuse
Spurgeon 1992
Bergetal. 1998
Pet-Soede et al. 1999
Burke et al. 2002
Cesar 2002
Fisheries, aquarium and curio trade,
pharmaceutical, construction,
tourism, research, education, social
value
Biological support, physical
protection, global life support,
social services
Tourism, mining, fishing Coastal protection
Blast fishing, nondestructive fishing, „ . ...
Coastal protection
Truincm *•
tourism
Tourism, recreation, fishing, blast
fishing, poison fishing, mining
Coastal protection
Live reef fish, man culture, aquarium . . . . .. . .
... 4-14- Biological support (habitat),
trade, pharmaceutical, tourism, . . . . .
. . . . physical protection, carbon
recreation, research, education,
,, . store
aesthetic
Existence value, option
value, intrinsic value
Food security, biodiversity
Aesthetic, biodiversity
Future uses, new
information, bequest
value, existence value
Burke & Maidens 2004 Tourism, recreation, fishing
MEA 2005
Spurgeon 2006
Burke et al. 2008
Hicks et al. 2009
Food, medicines, cultural and
amenity, aesthetic, recreational
Fisheries, tourism, recreation,
mining,
Tourism, recreation, fishing
Fishery, cultural, research, aesthetic,
recreational
Coastal protection —
Biodiversity, biological
regulation, nutrient cycling,
climate regulation, disease —
control, waste processing, flood
protection, erosion control
Biodiversity, coastal protection, Intergenerational equity,
carbon storage existence value
Coastal protection —
Biological control,
habitat/refuge, waste regulation,
coastal protection
Bequest, option, and
existence values
Table 1-5. Estimated annual economic impact from coral reef ecosystem services at global, regional, national,
and local levels
Scale/Location
Global
Caribbean
Indonesia
Philippines
St. Lucia
Tobago
Turks & Caicos Islands
Jamaica Portland Bight
Jamaica Montego Bay
Jamaica Montego Bay
Tourism
$9.6 billion
$2.1 billion
$103m
$108m
$160m-$194m
$101m-$130m
$18.2m
$llm
$3 15m
$400m
Fisheries
$5.7 billion
$0.3 billion
$1.221 billion
$620m
$0.5m-$0.8m
$0.8m-$1.3m
$3.7m
$19m
$1.3m
$4m
Coastal
Protection
$9.0 billion
$1.5 billion
$314m
$326m
$28m-$50m
$18m-$33m
$16.9m
$0.4m
$65m
$65m
_.. .. .. Carbon
Biodiversity _.
Storage
$5.5 billion —
$9m —
$10m —
$4.7m —
$18m $4m
$19.6m —
Ref.
1
2
7
7
6
6
8
3
4
5
References:
\: Cesar et al. (2003)
5: Gustavson (1998)
Source: Partially adapted
2: Burke & Maidens (2004) 3: Cesar et al. (2000) 4: Ruitenbeek & Cartier (1999)
6: Burke et al. (2008) 7: Burke et al. (2002) 8: Carleton & Lawrence (2005)
from Conservation International (2008).
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1.4 Ecological integrity, ecological resilience, and biodiversity
Ecological integrity, resilience, and biodiversity are all concepts that portray slightly different aspects of ecosystem condition
but convey the same sense that the attributes, processes, and functions of an ecosystem are all integral to the ecosystem itself
and necessary for sustaining it as well as its continued provision of services. These concepts represent the "glue" (Pearce &
Moran 1994, p. 22; Turner et al. 2000) of the properly functioning ecosystem, supporting the growth of reef-building corals for
shoreline protection, the presence of unique and diverse species to attract tourists, the creation of potentially useful natural
products, and the maintenance of habitat and nurseries for harvestable fish stocks. Assessing the extent to which an ecosystem
possesses any of these concepts provides insight into the many elements and processes that we do not know, understand, and
cannot otherwise measure. The development of concise, rigorous definitions and unambiguous metrics for these three attributes
continues but remains unfinished.
In choosing which topics to cover in this report, considerable thought was given to the role of ecological integrity, ecological
resilience, and biodiversity. Are these economic benefits, ecosystem services, or something else? How, or should, they be
included in our analysis? In the end, we decided that they were natural features rather than ecosystem services or economic
benefits (see the Definitions section at the bottom of Table 6-1, page 153), and consequently should not be included in this
report.
1.4.1 Ecological integrity
The Millennium Ecosystem Assessment (MEA 2005) describes supporting services as "those that are necessary for the
production of all other ecosystem services", including soil formation, photosynthesis, primary production, and nutrient cycling,
among others. Some have called these "biotic services" (Moberg & Folke 1999) or "fundamental services" (Holmlund &
Hammer 1999). Although called services, these could otherwise be characterized as functions of the ecosystem that ensure
ecosystem persistence and resilience, which are required for the sustainable delivery of services. The same characteristics of the
reef ecosystem have also been called "regulation functions" (de Groot 1992) and "primary" or "glue" value of the ecosystem
(Pearce & Moran 1994, p. 22; Turner et al. 2000). In many respects, these are all descriptions of ecological integrity, a concept
that has evolved from the stated objective of the Clean Water Act (CWA): "The objective of this Act is to restore and maintain
the chemical, physical, and biological integrity of the Nation's waters." (CWA 1972).
Although often debated, a generally accepted definition of ecosystem integrity is:
the capability of supporting and maintaining a balanced, integrated, adaptive community of organisms having a species
composition, diversity, and functional organization comparable to that of natural habitat of the region (Frey 1975; Karr
& Dudley 1981; Karr et al. 1986; Angermeier & Karr 1994).
Ecological integrity thus encompasses a variety of scales (genetic, taxonomic, ecological), elements (genes, species,
populations, landscapes), processes (colonizations, mutations, extinctions), and a dynamic biology in terms of evolution and
biogeography. Intended or not, the concept of ecological integrity appears to encompass the natural functions of an ecosystem
that provide services. Ecological integrity is generally decomposed into the components described by the CWA—represented as
the overlap of physical, chemical, and biological integrity (Figure 1-4). Yoder (1995) argues that the three components are not
equal and that physical and chemical integrity are components of biological integrity. Regardless, many authors (e.g., Turner
et al. 2000) suggest that ecological integrity is a component that should be valued as an ecosystem service.
Healthy reefs maintain their structure and function and provide the supporting services that allow for the fulfillment of
reasonable human needs through more direct provisioning of ecosystem services, such as shoreline protection, fisheries
production, and recreational opportunities (McField & Richards Kramer 2007). A number of indicators have been proposed and
monitored for assessing reef health (van Beukering & Cesar 2004; Healthy Reefs Initiative 2010).
Figure 1-4. Ecological integrity as the overlap of physical, chemical, and biological integrity.
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1.4.2 Ecological resilience
Ecological integrity is often equated with ecological health and ecological resilience. Although "health" is an anthropocentric
concept, it implies that a stressed ecosystem is similar to a diseased individual. While use of the term is sometimes neither
accurate nor precise (Ehrenfeld 1992), carefully structured definitions have been proposed using Energy Systems models
(Campbell 2000). Ecological resilience—"the capacity of a system to absorb disturbance and reorganize while undergoing
change so as to still retain essentially the same function, structure, identity, and feedbacks" (Walker et al. 2004; Folke 2006)—
is a more potent concept. Moberg and Folke (1999) conclude that managing resilience is the key to maintaining delivery of
ecosystem services. Clearly related to ecological integrity, resilience emphasizes the ecosystem's capacity to buffer stress and
reduce rates of system change. Rollings (1973) states that ecological resilience "determines the persistence of relationships
within a system and is a measure of the ability of these systems to absorb changes of state variables, driving variables, and
parameters, and still persist". Ecosystem integrity and resilience are thus closely linked, but provide a few distinctions.
Ecosystem integrity emphasizes a natural condition with implied stability, whereas resilience emphasizes stability with an
implied natural condition.
In the case of resilience, more than one stable state can exist. When considering a single stable state, resilience is measured by
the amount of time (after disturbance) required to return to the stable state (referred to as engineering resilience). When
considering multiple stable states, resilience is measured by the amount of disturbance needed to shift the system to an
alternative stable state (referred to as ecological resilience). This approach allows that an ecosystem can slide into a highly
stable, but less desirable state—in fact, less desirable states are often extremely resilient and difficult to revert to desired
conditions (Scheffer et al. 2000; Gunderson & Rolling 2002; Walker et al. 2004). Reversion to a desired state is the goal of
ecosystem restoration. Although ecosystem resilience is not usually considered an ecosystem service, it is similar to low
ecosystem integrity in that a less desirable state is expected to provide fewer ecosystem services.
1.4.3 Biodiversity
Biodiversity is frequently cited as an ecosystem service (e.g., Cesar 2002; CI 2008). Taken literally, biodiversity is simply taxa
richness, which is not an ecosystem service but is an ecosystem attribute that can contribute to a variety of reef services. High
biodiversity may contribute to ecological integrity, but there are many examples of natural systems with high integrity that do
not exhibit high biodiversity. Biodiversity more likely plays a contributory role in ecological resilience—high species richness
usually provides functional redundancy in the ecosystem, which increases resilience. If a keystone species is lost, another
functionally redundant species may take its place, and the ecosystem persists in its original state.
1.4.4 Insurance value
While the preponderant view seems to be that ecological integrity, resilience, and biodiversity are not ecosystem services, a
recent proposal by The Ecology and Economics of Biodiversity (TEEB) initiative combines the three concepts under the rubric
of "insurance value", which is defined as "the value of ensuring that there is no regime shift in the ecosystem with irreversible
negative consequences for human wellbeing" (Pascual et al. 2010). A similar notion put forward by Balmford et al. (2008) is
"infrastructure value", which is the capacity of an ecosystem to maintain its provision of ecosystem services despite variability
and disturbance.
The concept that ecosystem biodiversity and resilience provide insurance against a variety of uncertainties has been generally
accepted for some time in the environmental economics literature. While this literature's view of insurance value has been
evolving over the past 20 to 30 years, the notion that insurance value is a distinct economic nonuse benefit that can be valued is
not generally accepted, because of the difficulties described below. Generally speaking, the insurance value of ecosystems is
considered part of option value and quasi-option value (Pearce & Moran 1994, pp. 19-21; Pearce 2001; Weikard 2001), but our
limited understanding of ecological processes precludes our being able to be sufficiently certain of ecological outcomes to
move from uninsurable uncertainty to insurable risk (Pearce & Pearce 2001; Turner et al. 2003; Barbier 2006; Turner 2010;
Abson & Termansen 2011).
In the 1990s, Pearce and Moran (1994), Perrings (1995a; 1995b; 1998), Swanson et al. (1994), Turner (1992), and others began
describing how biodiversity and resilience might have an insurance value analogous to crop insurance, and included it as an
element of option value. They took care to distinguish between the insurable risk (the probability that a known adverse outcome
might occur) and the uninsurable uncertainly (wherein neither the outcome nor the probability are known) (Perrings 1995b).
That biodiversity and ecological resilience provided insurance in a more general sense was recognized, but the tangled
complexity of the relationship between that insurance and those "natural features" was viewed as an almost insurmountable
hurdle to its proper characterization (Pearce 2004). In the past five years, Baumgartner and colleagues have developed a more
generalized and rigorous definition of insurance value (Baumgartner 2007; Baumgartner & Quaas 2006; Quaas & Baumgartner
2008; Baumgartner & Strunz 2010), but the means to measure it as a distinct value not embedded in whole or in part within
other use and nonuse benefits (specifically, option value and quasi-option value) remains to be defined. The importance of
viewing insurance value (or the value of any ecosystem service) in terms of the value of a marginal change rather than a lump
sum value is regularly emphasized (Pearce 1998; Toman 1998; Heal 2000; Pearce & Pearce 2001), and to properly assess the
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value of marginal changes, ecological models must be able to predict the effect of small state changes (an ability that remains
beyond our reach).
Even though our incomplete understanding of ecological functions and processes and our resulting inability to predict
ecological outcomes prevent us from moving from uninsurable uncertainly to insurable risk, it may be that the inherent
instability of ecosystems near thresholds (or, tipping points) may preclude a quantification of the insurance value. Though these
limitations suggest that insurance value may be best characterized in qualitative terms (Fromm 2000), insurance value remains
a powerful and tangible concept that should be included in any ecological assessment.
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Work TM, Aeby GS and Maragos JE. 2008. Phase Shift from a Coral to a Corallimorph-Dominated Reef Associated with a Shipwreck
on Palmyra Atoll. PLoS ONE, 3(8):e2989. doi:10.1371/journal.pone.0002989.
Yoder CO. 1995. Policy issues and management applications for biological criteria. In: Davis WS and Simon TP (eds), Biological
Assessment and Criteria: Tools for water resource planning and decision making. Boca Raton (FL): Lewis Publishers, pp. 327-
343.
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Coral Reefs: Tourism and Recreation
Tourism is one of the most important economic sectors
worldwide and few environments are more important
for tourism and recreation than coastal zones.
(Moreno & Amelung 2009)
2.1 Describing the service
2.1.1 Definitions
It is important to understand the terms "tourism" and "recreation", particularly in the context of ecological economics.
Webster defines recreation as "refreshment of strength or spirits after work; also: a means of refreshment or diversion"
(Merriam-Webster 2010). Recreational activities are enjoyed by both tourists and residents of a given geographic location.
However, the common practice of economists is to differentiate between tourism and recreation based upon the source of
demand.
Tourists: people who "travel to and stay in places outside their usual environment for more than twenty-four (24)
hours and not more than one consecutive year for leisure, business and other purposes not related to the
exercise of an activity remunerated from within the place visited" (UNWTO 1995).
Residents: people who live at a particular place for a prolonged period (Leeworthy 2002; Princeton WorldNet
Glossary 2010a).
2.1.2 Tourism and the economy
Tourism is a significant sector of the global economy. According to the UN World Tourism Organization, tourism has become
one of the largest and fastest growing economic sectors in the world—as much as 30% of the world's exports of commercial
services and 6% of overall exports of goods and services are due to tourism. Globally, tourism ranks fourth after fuels,
chemicals, and automotive products in exports. For many developing countries, it is a main income source and the number one
export category (UNWTO 2009).
According to Goeldner and Ritchie (2009), global travel and tourism in 2011 is expected to account for:
• $7.0 trillion of economic activity
• 260 million jobs
The travel and tourism industry accounts for 24% of all U.S. service exports and 8% of total U.S. exports. Tourism is our
nation's third largest retail industry and the nation's largest service export. There are currently more than 7m Americans
employed directly in the travel industry, 9 million indirectly, for a total of over 16m jobs. In 29 states, tourism is the first,
second, or third largest employer (Goeldner & Ritchie 2009; Table 2-1).
Congress passed "The Travel Promotion Act of 2009", which highlights the importance of travel and tourism to our national
economy. The act creates a public-private partnership—the Corporation for Travel Promotion—to help bring more international
visitors to the United States (Sanchez 2010).
Table 2-1. Economic impact of travel and tourism (2009 preliminary data)
Travel expenditures*
Travel-generated payroll
Travel-generated tax revenue
Travel trade surplus
$704.4 billion
$186.3 billion
$11 3.0 billion
$22.0 billion
Travel-generated employment (direct)
Travel-generated employment (indirect)
7. 4m jobs
9m jobs
* includes spending by domestic and international travelers in the U.S. on travel related expenses
(i.e., transportation, lodging, meals, entertainment & recreation, and incidental items)
Source: Goeldner & Ritchie (2009); U.S. Travel Association (2010)
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Tourism demand is strongly dependent on the economic conditions in major generating markets. Discretionary income is
particularly dependent upon economic conditions. Tourism tends to account for a relatively large part of discretionary income,
particularly in emerging economies. And most significantly, "the growth of international tourism arrivals significantly outpaces
growth of economic output as measured in Gross Domestic Product (GDP)" (UNWTO 2010).
Marine tourism is the industry sector relevant to coral reef ecosystems. Orams (1999) provides a working definition for marine
tourism: "Marine tourism includes those recreational activities that involve travel away from one's place of residence and that
have as their host or focus the marine environment (where the marine environment is defined as those waters that are saline and
tide-affected)." This definition excludes business or work-related activities such as commercial fishing, shipping, oil
exploration, and scientific research.
The definition includes activities that have as their focus the marine environment such as shore-based fishing, land-based
whale-watching, or reef walking (Orams 1999). It also includes marine tourism development (accommodation, restaurants, food
industry, and second homes) and the infrastructure supporting marine development (e.g., retail businesses, marinas, and activity
suppliers) (Hall 2001).
2.1.3 Tourism and recreation activities related to coral reefs
Coral-reef based tourism and recreation is a subset of marine tourism. Leeworthy and Wiley (2001) provide estimates for a
comprehensive list of coastal and marine recreational uses. Recreational activities on or associated with coral reefs include:
• Scuba diving • Snorkeling
• Underwater photography • Recreational (sport) fishing
• Viewing nature and wildlife • Beach sunbathing
• Surfing • Collecting objects (e.g., dead shells, fragments of corals, driftwood)
2.1.3.1 Recreational (scuba) diving
Scuba diving is a type of diving that uses Self-Contained Underwater Breathing Apparatus (SCUBA) equipment for the
purpose of leisure and enjoyment.
The first documented underwater diving took place in 500 BC, when Scyllis demonstrated the practical use of breath-hold
diving by performing military exploits for the King of Persia. In the 1600s, dive bells were invented, with air being pumped in
from the surface. In the 19th century, Bert and Haldane researched the effects of water pressure on the body, and began to
define safe limits for compressed air diving. Also in the 19th century, improvements in technology (e.g., compressed air pumps,
carbon dioxide scrubbers, regulators, etc.) made it possible for people to stay under water for long periods, although still using
surface air (Martin 1997).
But it was not until the 20th century, with the invention of SCUBA, that recreational diving was really possible. Further
developments in technology have reduced the cost of training and equipment. Swim fins, masks, dive computers and other
scuba gear are available and affordable (Freeland 2010). Today, scuba diving is well regulated, with internationally governed
training programs and a code of conduct (Basiron 1997).
Scuba diving has become a popular leisure activity and globally, there are many locations that derive significant income from
scuba diving tourists. Recreational diving is an industry that is dependent upon the condition of the underwater resources.
Certain types of features make an interesting dive, and most of these are found on coral reefs including:
• Wildlife at the site (e.g., coral, sponges, fish, rays, mollusks, cetaceans, sharks and crustaceans).
• Complex topography of the site (e.g., coral reefs, drop offs).
• Historical or cultural items at the site (e.g., ship wrecks, aircraft) provide both historical value and form artificial habitats
for marine fauna.
• Good underwater visibility. Poor visibility is caused by particles in the water, such as mud, sand and sewage.
• Temperature. Warm water diving is comfortable and convenient.
The growth in recreational scuba diving can be measured by analyzing the increase in Professional Association of Dive
Instructors (PADI) diver certifications (Figure 2-1).
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Annual Diver Certifications (thousands)
-LMU£oin-jcD
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The goal of underwater photography is to bring back an image that will inform, entertain or educate. Attributes that contribute
to good underwater photographs include:
• Clear water. If the water is not clear, underwater pictures will not be either.
• Colorful fish, corals and other biota.
2.1.3.4 Recreational (sport) fishing.
Recreational fishing, fishing for pleasure or competition, is included as a component of tourism and recreation. Commercial
fishing (i.e., fishing for profit) and subsistence fishing (i.e., fishing for survival) are covered in the following chapter on fishing.
Many common salt water game fish spend some portion of their lives in coral reef ecosystems including tarpon, shark,
bonefish, permit, sailfish, marlin, bonito, pompano and mackerel. Historically, sport fishers, even if they did not eat their catch,
almost always killed them to bring them to shore to be weighed or for preservation as trophies. To protect recreational fisheries,
sport fishermen now often catch and release or tag and release, which involves fitting the fish with identity tags, recording vital
statistics, and sending a record to a government agency.
Recreational fishing is governed by a variety of conventions, rules, licensing restrictions and laws that typically restrict
sportfishing to hook, line, rod and reel rather than with nets or other aids.
Recreational fishing is big business, generating more than $125 billion in economic output and more than one million American
jobs. At nearly 40 million, more Americans fish than play golf (24.4 million) and tennis (10.4 million) combined. If fishing
were ranked as a corporation, it would be 47 on the 2007 Fortune 500 list of America's largest companies based on total sales
(Allen & Southwick 2008).
The recreational fishing industry consists of enterprises such as the manufacture and retailing of fishing tackle, the design and
building of recreational fishing boats, and the provision of fishing boats for charter and guided fishing trips. The American
Sportfishing Association (ASA) is the sportfishing industry's trade association, committed to looking out for the interests of the
entire sportfishing community (ASA 2010).
While recreational fishing is possible in almost any body of water, sport fishermen prefer locations where there are minimal
waves (calm water). Coral reefs are particularly attractive to recreational fishers because they:
• occur in calm waters;
• are found where the weather is generally hot and sunny; and,
• are populated with a wide diversity of fish species, including large, hard-fighting fish (tarpon, sailfish, sharks, mahi
mahi).
2.1.3.5 Viewing nature and wildlife.
Nature and wildlife watching is the practice of observing nature and wildlife (e.g., birds, dolphins, fish, manatees, turtles,
whales) in their natural habitat. Viewing is performed with the naked eye or through a visual enhancement device like
binoculars. The wildlife found in coral reefs and the adjacent seagrasses and mangroves are diverse and fascinating creatures.
They range from the charming and intelligent dolphin to the gentle, slow-moving manatee.
Guidelines for responsible wildlife viewing have been developed by federal and state agencies and NGOs. These agencies
support responsible wildlife viewing as a positive way to promote conservation and respect for the animals and the marine
environment.
Wildlife viewing is a significant commercial activity. For example:
• Whale-watching is estimated to be worth up to $2.1 billion per annum worldwide to whale watching operations,
employing around 13,000 workers (Cisneros-Montemayor et al. 2010).
• Bird-watchers contributed $36 billion per annum in the U.S. alone, and a fifth (20%) of all Americans identify
themselves as birdwatchers (U.S. Fish & Wildlife Service 2006).
Wildlife viewing can be done on an individual basis or through organized tours with knowledgeable marine naturalists. Guided
bird tours have become a major business with at least 127 companies offering tours worldwide (Wikipedia 2010a). Guided
kayak nature tours are a popular way to view birds, manatees, fish, and other wildlife in mangroves adjacent to reefs.
Recreational wildlife viewing is an industry that is dependent upon the condition of the supporting habitat. Coral reefs and the
adjoining mangroves and seagrasses are particularly attractive to wildlife viewers because they:
• occur in calm waters;
• are found where the weather is generally hot and sunny; and,
• are populated with a wide diversity of species, including birds, dolphins, fish, manatees, turtles, and whales.
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2.1.3.6 Beach sunbathing and swimming.
Sunbathing on wide sandy beaches is a popular recreational activity. Sandy beaches are an important part of the attraction for
major tropical and subtropical destinations, and many tourists spend at least some time at the beach. White, soft sandy beaches
composed of coral and shell particles are a favorite. The contrast between white sand beaches and the emerald, light blue, and
sparkling azure colors of tropical and subtropical sea waters is especially beautiful.
Coral reefs and mangroves play an important role in building and maintaining white sand beaches. Coral reefs serve as a sand
source. The sand is made of limestone of recent biological origin (e.g., corals, foraminifera, calcareous algas, mollusks, and
crustaceans) (UNEP/GPA 2003). Production of sand results from two processes: wave erosion of the coral reefs and fish
feeding on the coral and excreting coral sand (e.g., parrot fish, butterfly fish, and trigger fish).
However, not all beaches are made up of coral sand. Many Caribbean beaches are composed of sand derived from weathered
rock, and others consist of a mixture of coral and terrigenous sands.
Reefs also reduce wave energy creating calm waters desirable for swimming. Many sunbathers spend some portion of their
time swimming or wading in the water.
Attributes that contribute to good sunbathing beaches include:
• White coralline sands;
• Generally hot, sunny weather; and,
• Calm, warm waters
2.1.3.7 Collecting objects (beachcombing).
Beachcombing is the recreational activity of searching the beach and the intertidal zone for items that have washed in with the
tide (e.g., corals, seashells, sponges, sea fans, fossils, pottery shards, artifacts, sea beans, sea glass, and driftwood).
Beachcombing provides the opportunity to achieve better emotional, physical and spiritual health at little or no financial cost
(Ritterbush 2008, 2010).
Many beachcombers use knowledge of storms, geography, ocean currents, and seasonal events to determine the arrival and
exposure of rare finds (LaMotte 2004; Robinson & Robinson 1995; McRee 2009). Beachcombers tend to focus on the area
from just above the high tide line, usually marked by a row of seaweed and debris (wrack), down to the water's edge (McRee
2009).
Beachcombing equipment can be as simple as a bag or bucket to put the shells in and a shell identification guide. Many
beachcombers also use a metal detector, which aids in locating jewelry, coins and artifacts buried in the sand. Beachcombers
tend to be environmentally conscious and serve as stewards of the seashore (Ritterbush 2008).
Currently recognized beachcombing experts include: oceanographer Dr. Curtis Ebbemeyer (Flotsamterics and the Floating
World); eco-educator Dr. Deacon Ritterbush (A Beachcomber's Odyssey); sea glass experts Richard LaMotte (Pure Sea Glass)
and C.S. Lambert (Sea Glass Chronicles); geologist Margaret Carruthers (Beach Stones); shell specialists Chuck and Debbie
Robinson (The Art of Shelling); and, zoologists Dr. Blair Witherington and Dawn Witherington, (Florida's Living Beaches: A
Guide for the Curious Beachcomber) (Wikipedia 2010b).
Beaches adjacent to coral reefs are particularly attractive to beachcombers because they:
• provide unique, beautiful shells, corals, and sponges;
• often have wide sandy beaches;
• generally have hot and sunny weather; and,
• experience occasional tropical storms that bring in treasures.
2.1.3.8 Surfing.
Surfing is the sport of riding a surfboard toward the shore on the crest of a wave (Princeton WorldNet Glossary 201 Ob).
Ancient Polynesians (e.g., Hawaiians, Samoans, Tongans, Tahitians, and Maori) integrated surfing into their culture and
considered surfing an art. Hawaiians referred to this art as he 'e nalu, which translates into English as "wave sliding". Samoans
call surfing^a 'ase 'e or se 'egalu (Kramer 1994). The most skilled surfers were often members of the upper class, which
included chiefs and warriors, who had access to the best waves (Young 1983).
In 1779, Lieutenant James King, the newly promoted captain of the HMS Discovery (following Captain James Cook's demise),
devoted two full pages of the ship's log to a description of surfboard riding, as practiced by the locals at Kealakekua Bay on the
Kona coast of the Big Island. His entry is the earliest written account of surfing (Marcus 2010). The sport was also recorded in
print by Augustin Kramer, a 19th century German ethnologist, author, collector, and expert on Polynesian and Samoan culture.
European missionaries forbade or discouraged many Polynesian traditions including surfing. By the 20th century surfing had
almost disappeared. Only a few Hawaiians continued to practice surfing and the art of crafting surfboards. In 1905, Duke
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Kahanamoku and his friends created a surf club in the Waikiki area of Hawaii. Duke and his friends are credited for bringing
surfing back to Hawaii and exposing the world to surfing. In 1907, Jack London went to Hawaii and was introduced to surfing
by Alexander Hume Ford, an eccentric journalist and wanderer. London subsequently wrote A Royal Sport: Surfing in Waikiki,
which included descriptions of Waikiki and Alexander Hume Ford. His story was published in the October 1907 edition of The
Lady's Home Companion and again in 1911 as part of The Cruise of the Snark. In 1907, another member of the Waikiki surf
club, George Freeth, introduced surfing to California. Duke Kahanamoku introduced surfing to Australia in 1915.
Until the 1960s only a small number of people were involved in surfing, mainly in Hawaii, Australia and California. The
release of the movie Gidget moved surfing from an underground culture to a national fad. Since then, films about surfing have
continued to play a part in the evolution of surfing.
Globally, there are over 10 million surfers. In the Indo-Pacific, the commercial surf tourism industry is strongly linked to the
clothing, fashion, and entertainment industries, and marketed through specialist surfing magazines and surfing media (Buckley
2002).
Coral reefs and other shallow water formations (e.g., rocks, sandbars, etc.) allow waves to break, thereby forming a surfable
wave. Collectively, these breaks are known as surf breaks (Silmalis 2007). A reef break is a wave that breaks over a coral reef.
When a coral reef is exposed to open ocean, there is potential for a fast and hollow wave. As the wave swell approaches from
deep water, it hits the shallower reef, escalating in height before pitching and curling over the reef. The waves at Pipeline in
O'ahu, Cloudbreak in Fiji, and Jaws in Maui are among the most famous and photographed reef-break waves. Experienced
surfers are rewarded with a fast tubing ride on a ramp-like wave. Most surfing competitions take place on reef breaks (The
Surfing Site 2010).
According to Baker (2007), the Caribbean has fewer surf breaks than the Pacific, but offers great surfing during winter and
spring. The waves provide short powerful rides, sometimes sweeping over the coral reefs, creating demanding tubes. The best
surfing conditions occur when Atlantic storms push through the Caribbean in late May through early September (hurricane
season) and December through March (when Atlantic storms push through the Caribbean). When coral reefs are destroyed,
waves may diminish, so preserving coral reefs is critical to preserve the sport of surfing (NOAA 1997).
2.1.4 Tourism and recreation: businesses
A diverse range of businesses forms the coral reef tourism industry. Those directly associated with coral reef tourism include
small businesses such as charter fishing boat operators, sea kayak tours, and scuba diving instructors, etc. They also include
moderate-sized private companies like coral reef dive-boat operators and large corporations such as those that manufacture and
retail fishing tackle and recreational fishing boats. An even greater number of businesses are indirectly associated with coral
reef tourism (e.g., boat maintenance shops, coastal resorts, island ferry services, and artists). Government agencies monitor and
manage coral reef tourism (e.g., park authorities, fisheries control agencies, tourism marketing and promotion bodies, law
enforcement agencies, and marine safety organizations). Nonprofit groups also form an important component of the industry
(e.g., clubs for scuba diving and fishing) (Orams 1999).
2.2 Necessary conditions for providing the service
Coral reefs are the most biologically diverse marine ecosystems on earth, rivaled only by tropical rainforests (Sebens 1994;
Odum 1997). Coral reefs cover less than 0.1 % of the ocean's surface (an area about half the size of France) but support about
25% of all marine species (Wilkinson 2004; Mulhall 2007).
Coral reefs occur in seas with very specific environmental and climatic conditions.
• Mainly in tropical and subtropical seas—between 30°N and 30°S latitudes.
• Warm ocean temperatures (68-82°F, or 20-28°C). Warm water flows along the eastern shores of major land masses.
• Generally at depths of less than 150 ft (46 m), where sunlight penetrates. Because reef- building corals have a symbiotic
relationship with zooxanthellas, a type of microscopic algas, sunlight is necessary for these corals to thrive and grow.
• High salinity, low CO2 concentration, and low acidity, facilitating precipitation of calcium from the water necessary to
form a coral polyp's skeleton.
• Strong wave action. Waves carry food, nutrients, and oxygen to the reef, distribute coral larvas, and prevent sediment
from settling on the coral reef.
• Most corals grow on a hard substrate.
Coral reefs depend on the interaction of many species, including hard and soft corals, fish, sponges, crustaceans (including
shrimp, lobsters, and crabs), echinoderms (including starfish, sea urchins, and sea cucumbers), sea turtles, and cetaceans. Hard
and soft corals provide the structural habitat that supports this high abundance and diversity. Bryozoans encrust coral skeletons
and reefs debris, cementing the reef structure. Fish, crabs, and lobsters find shelter in the reef structure and play a vital role in
the reef's food web.
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2.2.1 Linkages among reef condition, reef structures, and reef functions with respect to delivery of the service
The complex three-dimensional coral reef structure provides habitat for the high numbers and diversity of marine organisms
that support tourism and recreation (Bradley et al. 2008; Fisher et al. 2007; Courtney et al. 2007; Cesar 2002; Wilkinson 2002;
Done etal. 1996; Hoegh-Guldberg 1999; Muscatine 1980, 1990; Reaka-Kudla 1996; Sebens 1994; Sale 1991; Grassland et al.
1991; Sutton 1983; Loy a 1972). Stony corals are the basic building blocks of the coral reef (Human & DeLoach 2002). Massive
species (e.g., Montastrea spp., Diploria spp.) and branching corals (e.g., Acropora spp.) provide significant three-dimensional
surface area that functions as essential habitat for fish and other reef-dwelling animals (Mumby & Steneck 2008; McField &
Richards Kramer 2007; Moberg & Folke 1999). Tourism and recreation are, therefore, directly or indirectly dependent upon the
reef-building corals (Moberg & Folke 1999).
Most reef-building corals have photosynthetic algas (zooxanthellas) that live in the coral tissue. The coral provides the
zooxanthellas with a protected environment and compounds they need for photosynthesis. The zooxanthellas provide glucose,
glycerol, and amino acids (the products of photosynthesis) that the coral uses to make proteins, fats, and carbohydrates, and
produce calcium carbonate (Barnes 1987; Barnes & Hughes, 1999; Lalli & Parsons 1995; Levinton 1995; Sumich 1996).
Zooxanthellas also give stony corals their beautiful, bright coloration.
When stony corals become physically stressed, they expel their zooxanthellas and the coral colony bleaches (Barnes & Hughes
1999; Lalli & Parsons 1995). This, in turn, results in a reduction of energy (in the form of various photosynthates) being
provided to the host (Hoegh-Guldberg & Smith 1989; Brown 1997), and a subsequent loss of tissue biomass (Porter et al. 1989;
Szmant & Gassman 1990; Fitt et al. 1993), coral skeletal deposition (Goreau & MacFarlane 1990), and fecundity (Szmant &
Gassman 1990). Extended periods of coral bleaching can result in the coral's death.
Poor coral health may also adversely impact fish production. While overall abundance of reef fishes is correlated mainly with
structural complexity of reefs (Jones & Syms 1998; Done 1999), several short-term studies have documented decreases in fish
productivity, species richness, fish biomass, and potential yield (reef biocapacity) (Warren-Rhodes et al. 2003; Graham et al.
2006; Sano 2004) resulting from lost coral structural complexity. Over the long term, however, coral reef structure will reduce
in complexity as corals die and bio-erode.
Coral reefs are part of a tropical marine "seascape that functionally links them with the adjacent tropical ecosystems (i.e.,
mangrove forests and seagrass meadows)" (Mumby & Steneck 2008). This seascape mosaic: (McField & Richards Kramer
2007; Mumby et al. 2004)
• provides critical foraging areas, nurseries and refugia;
• provides physical and chemical buffering;
• facilitates energy and material flows; and,
• creates corridors for transient species.
For example, mangroves strongly influence the community structure of fish on neighboring coral reefs (Mumby et al. 2004),
and also trap sediments, nutrients and pollutants, improving the water quality on nearby reefs (Grimsditch & Salm 2006).
Seagrasses contribute nutrients to the coral reefs and produce colored dissolved organic matter (CDOMs), which can protect
coral against bleaching by screening harmful solar radiation (Salm & West 2003).
Reef fish respond to this spatial mosaic, many showing pronounced associations with specific habitat types (Sale & Kritzer
2008). Coral reefs provide essential habitat for adult fish. The three-dimensional coral reef structure protects shorelines and
creates calm waters necessary for seagrass meadows and mangrove forests to thrive. The rainbow parrotfish, grunts, barracudas,
and several snapper species depend on these mangrove forests and seagrass beds for nursery habitat (McField & Richards
Kramer 2007).
The tropical marine mosaic also supports "charismatic megafauna", large animal species with widespread popular appeal (e.g.,
manatees and dugongs, sea turtles, rays, sharks, and dolphins). Some of these species (e.g., manatees and sea turtles) use a
variety of habitats during different life stages (McField & Richards Kramer 2007).
Many of the attributes discussed above make coral reefs particularly attractive destinations for tourists. The warm, sunny
weather, clear, calm waters, and wonderful species diversity and richness all contribute to this appeal. The desirability of these
attributes for tourism and recreation has been well documented in the literature. For example, Pendleton (1994) points out that
scuba divers look for high-quality coral reef habitats (as indicated by live coral coverage), coral and fish diversity, and water
clarity. Leujack and Ormand (2007) reported that 51% of survey respondents were interested in both fish and corals, whereas
36.5% were only interested in fish and 5.2% only in corals, while another 5.2% stated that besides being interested in corals
they also looked for other things on the reef. Uyarra et al. (2005) found that divers correctly perceived differences between sites
in the condition of biological attributes such as:
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• fish species richness;
• total number offish schools;
• live coral cover;
• coral species richness; and,
• reef structural complexity.
Leeworthy et al. (2004) grouped survey respondents' preferences into three categories:
• natural resource attributes (e.g., clear water, amount of living coral, fish/biota diversity, megafauna, and beach quality);
• natural resource facilities (e.g., parks, shoreline access, marina facilities, mooring buoys, boat ramps); and,
• other facilities (parking, roads, rest rooms) and services.
Studies in Israel (Weilgus 2004) and Tobago (Beharry-Borg & Scarpa 2010) have documented the importance of water clarity,
coral cover, and fish abundance for divers and snorkelers.
There are other factors that influence the selection of a tourism venue, including the availability of facilities and amenities (e.g.,
boats, dive shops, fishing guides, hotels and restaurants, etc.), the perceived "healthy" condition of the waters (e.g., lack of
pollution, absence of debris), and even perceptions of crowding (Park et al. 2002; Beharry-Borg & Scarpa 2010). Inglis et al.
(1999) and Leujack and Ormand (2007) documented the effect of perceived crowding on snorkelers' enjoyment. Table 2-2,
below, summarizes some of the most important attributes that have been identified in the literature.
Table 2-2. Features relevant to the perceived value of coral reefs
Natural Features
Biotic Features
Species richness (coral, fish,
sponges, etc.)
Variety of species characteristics
(coral, fish, sponges, etc.)
• colorful
• large
• rare
Charismatic megafauna species
diversity (birds, marine mammals,
turtles)
Coral health
3 -dimensional reef structure
Coral/macroalgas ratio
Abiotic Features
Warm ocean temperatures
Water clarity
Calm waters
White coralline sands
Proximity to deep ocean & waves
Connectivity with the adj acent
tropical ecosystems
Social Features
Perceptions of crowding
• # of divers/snorkelers
• # of proximal boats
Lack of pollution
Absence of debris
The coral reef tourism industry depends upon high-quality, pristine or undisturbed assets (Basiron 1997). Coral reef
degradation directly impacts delivery of the tourism and recreation services (Cooper et al. 2009). For example, recreational
fishing is extremely dependent upon the health of coral reefs since many marine game fish species (e.g., tarpon, groupers,
snook, barracuda, and dolphin) utilize reef habitats for at least part of their life cycles (Bryant et al. 1998).
Recreational diving is especially sensitive to reef condition, and thus particularly vulnerable to degradation (Cooper et al.
2009). As an example, dive tourism in Zanzibar decreased by 20%, snorkeling in Sri Lanka declined substantially, and there
was an estimated $1.5m annual loss in tourism dollars in the town of El Nido, Philippines, after the mass bleaching and coral
mortality of 1998 (Wilkinson et al 1999; Cesar 2000; Bruno 2008). The relative cover of benthic habitats in an area may be
indicative of snorkeling or swimming opportunities, with certain types of benthic habitats, such as patch reefs or Montastrea-
dominated reefs, having greater recreational value than those dominated by seagrass or macroalgas (Mumby et al. 2008).
2.3 Measuring the service
The recreational ecosystem services provided by coral reefs (the opportunity to dive, snorkel, and fish) have not been directly
measured. However, there are surrogate measurements related to the natural features that support these ecosystem services that
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can (and have) been measured. These could potentially serve as, or be combined into, multi-metric indicators of final ecosystem
services.
A wide range of biological indicators have been developed and are documented in Jameson et al. (1998). Methods for
monitoring stony corals, octocorals, fish, benthos, and sponges have been developed and tested by EPA.
2.3.1 Stony corals (scleractinians)
Stony corals (phylum Cnidaria, class Anthozoa, subclass Hexacorallia) build and maintain the physical infrastructure that
supports all other organisms in the community (Fisher et al. 2007; Fisher et al. 2008). Stony corals constitute the basic
framework and substrate for many other organisms that penetrate the skeletal mass (sponges, polychastes, sipunculides,
bivalves, and gastropods). The complex skeletal structure of stony corals also provides habitat for the high numbers and
diversity of marine organisms that support fisheries and tourism (Grassland et al. 1991; Done et al. 1996; Hoegh-Guldberg
1999; Muscatine 1980, 1990; Reaka-Kudla 1996; Sebens 1994; Wilkinson 2002). Coral structures also protect coastal
shorelines from wave and current erosion (Costanza et al. 1997; Pernetta 1992).
Because the health, growth, and recruitment of stony corals are crucial to reef sustainability and future benefits, these corals are
often considered the primary indicator organisms for reef communities (Loy a 1972; Brown 1988; Done 1997). Stony coral
colony size is an extremely important attribute, because colony size determines the contribution of each colony and species to
community habitat, biomass, photosynthetic activity, metabolism, and calcium carbonate deposition. Colony size is a major
determinant of growth, reproduction, population dynamics and community interactions (Fisher 2007).
The stony coral rapid bioassessment protocol (RBP) (Fisher 2007) relies on three observations (colony identification, colony
size, and proportion of live tissue), that can be combined in different ways to generate multiple indicators that characterize the
value and sustainability of coral reefs. These indicators can be used to assess the capability of reefs to continue providing
ecosystem services such as reef-based tourism and recreation (Table 2-3). Unique to the RBP is the ability to generate three-
dimensional indicators that help to quantify the complex three-dimensional structure of the reef that is so important to
providing the ecosystem services.
2.3.2 Octocorals (gorgonians)
Marine octocorals (phylum Cnidaria, class Anthozoa, subclass Octocorallia [aka Alcyonaria]) (including gorgonians, blue
coral, soft corals, and sea pens) are sessile invertebrates that provide substantial spatially varied biogenic habitat for adult and
juvenile fish and other invertebrates (Pugliese 1998; Lybolt 2003). Octocoral surface area and topographical heterogeneity are
therefore extremely important attributes.
EPA has developed a method that can be performed in conjunction with stony coral monitoring to estimate the total three-
dimensional surface area of octocorals on coral reefs. Divers classify marine gorgonians by colony morphology and measure
their maximum height and diameter. Morphology-specific equations are used to calculate surface area. These indicators can be
used to assess habitat availability for fish and macrobenthos.
The protocol may be supplemented to identify gorgonians by taxonomy and report adverse health conditions (e.g., bleaching,
disease, predation, etc.) These additional observations can be used to estimate additional indicators (gorgonian abundance,
density, and richness) (Santavy et al. in review). These additional indicators could be used to assess features that would be
attractive to divers and snorkelers.
2.3.3 Sponges
Marine sponges (phylum Porifera) provide habitat for fish and other invertebrates, cement and reinforce reef structure,
contribute to nitrogen and carbon cycling through microbial symbionts, and efficiently filter sediment, algas, and small
organisms from the water column. Three-dimensional sponge area is the critical attribute that supports these services.
EPA has developed a method that can be used to estimate the three-dimensional surface area of marine sponges. Divers classify
sponges by colony morphology, then measure height and maximum diameter. Dimensions are converted to surface area using a
formula derived for each morphological type. If the necessary expertise is available, additional data collection can include
taxonomic identification and adverse physical condition (e.g., bleaching, disease, and predation). Such data will permit
estimation of sponge abundance, density, surface area, and, if included in the protocol, taxa richness and physical condition
(Santavy et al. in review).
2.3.4 Fishes
Reef fish are major components of coral reef ecosystems. The coral reef ecosystem is a very complex environment with many
niches. Reef fish fill these niches, helping to sustain the balance of the reef. Reef fish also provide a readily available food
source and are an important aspect of tourism and recreation (e.g., sportfishing and diving/snorkeling).
Species diversity and richness are critical attributes that support these services. NOAA has developed and tested fish survey
techniques that can be performed in conjunction with stony coral monitoring. Divers assess the species, numbers, and sizes of
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all reef fishes within an underwater transect. This protocol is a noninvasive, timed, rapid assessment that takes approximately
30 minutes to complete (Menza et al. 2006). The fish are classified according to size categories. Visual data are used to
estimate abundance, species richness, and biomass for the fish populations and different feeding guilds sampled. Although not
yet included in the protocol, the visual data collection could be expanded to include indicators of fish color, which is an
attribute of interest to divers and snorkelers.
2.3.5 Mangroves and seagrasses
Remote sensing provides an efficient way to track the change in the areal extent of mangrove forests and seagrass meadows
over time. Landsat or other satellite images can be used to measure the extent and spatial patterns of coral reefs, seagrass
meadows, and mangrove forests (McField & Richards Kramer 2007). The development of indicators that relate landscape
composition and pattern attributes (including hydrology and transitional coastal systems) to coral reef condition is a new
research area for EPA.
Table 2-3. Stony coral rapid bioassessment protocol coral condition indicators*
Abundance and Composition
Abundance
Density
Relative species abundance
Species (taxa) richness
Species frequency
Species diversity
Protected species
Community composition
number of colonies
number of colonies per m2 sea floor
abundance of a particular species per total abundance
number of species occurring in a reef or region
proportion of sites where a species occurs
index of taxa richness and relative abundance
richness and abundance of protected coral species
relative richness or abundance of a species or groups of species with
some discretionary biological or physical attribute (e.g., tolerance)
Physical Status
Colony surface area (CSA)
Total surface area (TSA)
3D total coral cover (3DTC)
Average colony surface area (AvCSA)
Population structure
Community structure
3D skeletal surface area of an entire colony (m )
!L CSA for all colonies at a transect, station or reef
TSA per m2 sea floor
TSA / # colonies
colony size distribution for a species compared to colony number or
other attribute
colony size distribution for all species compared to colony number or
other attribute
Biological Condition
Percent live tissue (%LT)
Average percent live tissue (Av%LT)
Colony live surface area
Live surface area (LSA)
3D live coral cover (3DLC)
Percent Live Surface Area (%LSA)
proportion of live coral tissue on each colony
2 %LT / # colonies
live tissue on a colony (m2) = (CSA x [%LT / 100])
IL colony live surface areas at a transect, station or reef (m )
LSA per m2 sea floor
comparative ratio of live and total surface area = ([LSA / TSA] x 100)
* Indicators are derived from three core observations (colony identification, colony size, and proportion of live tissue)
on stony coral colonies and can represent cumulative or average values for transects, stations, and reefs or for a
particular species or group of species.
2.4 Valuing the service
Across the globe, nearly half a billion people are located within 100 km of a coral reef, and therefore, receive some benefit
from the ecosystem services provided by coral reefs (Park et al. 2002). Local communities depend on coral reefs as an
important source of employment, income, and tourism revenues (Ahmed et al. 2007). Consequently, tourism (and recreation) is
one of the most commonly valued ecosystem services. Tourism (and recreation) is a direct, largely nonconsumptive use of coral
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reef ecosystems. Sportfishing is largely catch and release, so we consider it to be nonconsumptive even though a portion of
sportfishing remains consumptive.
There is a considerable body of literature on valuation of tourism and recreation in coral reef environments. Many different
valuation methods have been developed, tested, and applied globally; however, the results are rarely comparable (Burke et al.
2008). Values for ecosystem services (UNEP-WCMC 2006) will vary according to:
• the location (e.g., reefs that are major tourist destinations will have a higher value in terms of diving and other reef-
related activities than those where tourism has not been developed);
• the length of time being considered and whether a prediction for the future is involved (e.g., all reefs are potentially of
value for diving tourism, but some may have no value at present);
• visitor responses to marginal changes in reef quality (e.g., some people are more sensitive to changes and will place a
higher value on maintaining reef quality than those less sensitive to such changes); and,
• the method used and the assumptions made.
Table 2-4 below describes commonly used approaches to economic valuation and relates them to particular services.
Approaches relevant to tourism and recreation are Travel Cost (TC), Effect on Production (EoP), Financial Analysis (FA), and
Contingent Valuation Method (CVM). A brief description of some of these methods is given in the following sections.
Table 2-4. Methods used for valuing goods and services of coral reef ecosystems
Generally Applicable Methods
>• Using the change in conventional market value of goods and services that results
from a change in the environmental resource
• Change in Productivity / Effect of Production (EoP)
• Change in Stock (houses, infrastructure, land) at Risk (SaR)
• Loss of earnings / Human capital approach (HC)
• Opportunity cost approach (OC)
>• Using the value of direct expenditures (cost based)
• Preventive expenditures (PE)
• Compensation payments (CP)
>• Using the observed market prices to analyze the current economic activity
generated (Financial Analysis)
Potentially Applicable Methods
Using implicit or surrogate market values - indirect approaches
• Properly-value and other land-value approaches (PV)
• Travel-cost approaches (YC)
Using the magnitude of potential expenditures (cost based)
• Replacement costs (RP)
• Shadow-project costs (SPC)
Survey-Based Methods
>• Using surveys of individuals to elicit values
• Contingent valuation method (CVM) - hypothetical markets and situations
(Willingness-to-Pay [WTP] and Willingness-to-Accept [WTA])
Source: Cesar 2000
2.4.1 Effect on production (EoP)
EoP estimates the difference in value of productive output before and after the impact of a threat or a management intervention.
The change in net profit (i.e., effect on production) can be calculated and used as a proxy for the loss in tourism value. One
challenge with this method is determining and modeling the relationship between the damage to an environmental resource and
its corresponding impact on the production of the specified good or service. An example of EoP is the previously cited coral
mortality in 1998 and loss of tourism revenues in Zanzibar, Sri Lanka, and the Philippines (Bruno 2008).
2.4.2 Financial analysis (FA)
FA uses observed current financial activities, revenues, costs, and financial flows in the economy from market-based uses of
the reef (such as diving and snorkeling) to analyze the economic activity generated by use of an ecosystem good or service.
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Data availability and quality vary considerably and have direct bearing on the statistical confidence of the resulting analysis. In
addition, this approach will underestimate the tourism value of reefs, because it omits consumer surplus (the additional welfare
a consumer enjoys beyond what he or she has paid for the service) (Cooper et al. 2009).
An example is the World Resources Institute (WRI) economic valuation of coral reefs in Tobago and St. Lucia (Burke et al.
2008). Using FA, Burke et al. (2008) valued direct and indirect economic impacts from visitor spending in 2006 associated
with coral reefs in Tobago to range from $101m to $130m and in St. Lucia from $160m to $194m.
For both EoP and FA, it is important to calculate secondary (i.e., indirect) impacts on the economy from spending by coral reef
associated visitors. Economists estimate the magnitude of these indirect impacts using a tourism multiplier. A multiplier of 1.6,
for example, represents 60 cents of additional impact for every $1 in direct tourist expenditure. The size of the multiplier is
influenced by the portion of goods and services used in the tourism sector that are produced domestically, such as linens,
beverages, food, dive equipment, and construction materials. (Cooper et al. 2009).
2.4.3 Travel costs (TC)
TC uses the travel time or travel costs as a proxy "total entry fee", and therefore, it is a measure of a person's willingness to pay
for visiting a particular tourist location. The further away people live from the location, the higher the costs. A demand curve
can be developed and the associated consumers' surplus can be determined. This surplus represents an estimate of the value of
the environmental good in question (e.g., the coral reefs).
An example of TC is given by Pendleton (1995), who used this method to estimate the economic value of the Bonaire Marine
Park. Pendleton used marine park permit data to estimate the number of visitors from each state and county. This number was
then divided by the population of the state or county to determine a visitation rate that was then regressed on travel costs,
providing the demand curve for coral reef associated vacations to Bonaire. Pendleton was then able to calculate the annual
value of Bonaire Marine Park at approximately $19.2m.
2.4.4 Contingent valuation method (CVM)
CVM is used to obtain information on consumers' preferences by asking people what they are willing to pay for a benefit
(willingness to pay or WTP), or what they are willing to accept by way of compensation to tolerate a loss (willingness to accept
or WTA). Analysts may use either a direct questionnaire/survey or experimental techniques in which subjects respond to
different stimuli under controlled conditions. Analysts now use a combination of conjoint analysis (developed in the social
psychology field) and multi-attribute utility theory (Adamowicz et al. 1998).
An example of a CVM is given by Spash (2000), who surveyed visitors to Montego Bay (Jamaica) and Curacao (Netherlands
Antilles) to estimate the benefits of maintaining and improving coral reef biodiversity (a nonuse benefit). Respondents were
willing to pay $3.24 per person for Montego Bay and $2.08 per person for Curacao to preserve coral reef biodiversity (Spash
2000). A weakness in applying CVM in this context is whether respondents believe that coral reefs possess inherent rights or
that humans have a duty to protect coral reefs. Such preferences could increase WTP by up to a factor of three (Spash 2000).
Economists are now combining various approaches to achieve a more robust valuation. Park et al. (2002) developed a TC-CV
model of demand for trips to the Florida Keys focusing on willingness to pay to preserve the current water quality and coral
reef condition. The integrated model "incorporates key factors for establishing baseline amenity values for tourist dive sites,
including perceptions of reef quality and dive conditions" (Park et al. 2002).
2.4.5 National scale valuation of tourism
Most countries maintain National Account Systems (NAS), which provide a complete and consistent conceptual framework for
measuring the economic activity of a nation. NAS are derived from a wide variety of source data including surveys,
administrative and census data, and regulatory data. Most countries have a national statistical office or central bank that
compiles, integrates, harmonizes, and publishes the data. NAS include a number of aggregate measures (e.g., gross domestic
product [GDP], disposable income, savings and investment) and other information (e.g., input-output tables that show how
industries interact with each other in the production process).
In the United States, the Bureau of Economic Analysis (BEA) is responsible for the national account. BEA prepares and
publishes a variety of economic statistics on U.S. industries, including the annual industry accounts and the benchmark input-
output accounts. U.S. industries are defined according to the North American Industry Classification System (NAICS) (BEA
2010).
The North American Industry Classification System (NAICS) was cooperatively developed by the United States, Canada, and
Mexico, and it uses a production-oriented conceptual framework that groups establishments into industries based on the activity
in which they are primarily engaged. Corporate entities using similar raw material inputs, similar capital equipment, and similar
labor are classified in the same industry. NAICS is based on the product that is being produced (e.g., accommodation services)
rather than who is consuming the product (e.g., a tourist or a local resident).
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Tourism is not designated as an industry in NAICS. Measuring the economic contributions of tourism presents a twofold
challenge:
1. the "Tourism Industry" is actually made up of parts of many industries; and,
2. tourism is traditionally measured and understood from the demand side (i.e., what are visitors spending?),
while industries are properly measured from the supply side (i.e., what is being produced?).
To properly value tourism in a manner consistent with the other economic accounts, the World Tourism Organization and the
United Nations developed the Tourism Satellite Account (TSA) standard (Sacks 2004). "Satellite accounts provide a framework
linked to the central accounts and which enables attention to be focused on a certain field or aspect of economic and social life
in the context of national accounts; common examples are satellite accounts for the environment, or tourism, or unpaid
household work" (OECD 2010).
The BEA develops the U.S. Travel and Tourism Satellite Accounts (TTSAs), based on the benchmark input-output accounts
and consistent with the integrated annual industry accounts. BEA methods used to prepare the TTSAs are consistent with the
methods used to estimate GDP, national income, and other national economic measures. The BEA (2010) characterizes the
TTSAs in the following way:
The TTSAs present a detailed picture of travel and tourism activity and its role in the U.S. economy. These
accounts present estimates of expenditures by tourists, or visitors, on 24 types of goods and services. The
accounts also present estimates of the income generated by travel and tourism and estimates of output and
employment generated by travel and tourism-related industries. The accounts are updated annually and have
been expanded to provide quarterly estimates of the sales of goods and services to travelers and employment
attributable to those tourism sales.
Several states are now developing their own TTSAs (Alaska, Delaware, Hawaii, New Jersey, North Carolina, Rhode Island,
South Carolina, and Virginia) (Aydin 2008). There has also been interest in developing a TTSA for Florida (Ayden 2008;
Florida Tax Watch News 2007).
2.4.6 National scale valuation of recreational fishing
The Sustainable Fisheries Act of 1996 (PL 104-297) and the Magnuson-Stevens Fishery Conservation and Management
Reauthorization Act of 2007 (PL 109-479) both amend the Fishery Conservation and Management Act of 1976 and are
incorporated into 16 U.S.C. 1801. Together, they mandate collection of detailed information on marine recreational fishing.
Since 1981, the National Marine Fisheries Service (NMFS) has conducted an annual survey of marine recreational fisheries
covering all fishing modes (private/rental boat, party/charter boat, and shore) and including estuarine and brackish water. A
variety of survey methods are used, including:
• a coastal household telephone survey (CHTS);
• a telephone survey of for-hire fishing vessel operators (FHS); and,
• a field intercept survey of angler fishing trips.
Additional information is also obtained from State or regional logbook programs and is used to supplement survey
data to produce more robust catch and effort estimates (NOAA 2009).
An additional source of information on recreational fishing can be obtained from the U.S. Fish and Wildlife Service's National
Survey of Fishing and Hunting (NSFH), which collects economic information about recreational saltwater fishing at five-year
intervals. The NSFH canvasses the U.S. population by telephone and conducts personal interviews with a subsample to obtain
statistically reliable results at the State level.
2.4.7 Noneconomic human dimensions measures
Economic values are contingent on income and wealth and, therefore, cannot capture the full value of ecosystem services.
Human values (e.g., social, political, cultural, spiritual) can also be measured. Several key concepts (importance, satisfaction,
and expectation) can be measured using survey techniques.
• Importance refers to how a consumer would rate various attributes of the service. Important attributes would presumably
figure heavily in choices among alternatives (Alpert 1980).
• Satisfaction is the consumer's fulfillment response received from a service (Myers & Alpert 1968). Satisfaction is
influenced by the consumer's perceptions of experienced quality, service quality, price, and other factors (Loomis et al.
2008a).
• Expectation is what the consumer believes is most likely to happen. An expectation may or may not be realistic.
Expectations may significantly condition perceptions of experiences or services.
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2.4.7.1 Importance-satisfaction ratings.
Martilla and James (1977) first described the concept of importance-satisfaction when considering people's demands for goods
and services. Importance-satisfaction involves rating certain attributes of the service on their importance to the rater and on
satisfaction with the service. Connecting importance-satisfaction ratings to a conceptual model of economic demand and value
allows for interpretation of the satisfaction ratings as "indicators" of demand and value for particular ecosystem
attributes/ecosystem services. Importance-satisfaction ratings are widely used to assess the demand for outdoor recreation
(Guadagnolo 1985; Richardson 1987; Joppe et al. 2001; Tonge & Moore 2007) and have recently been used to assess demand
for tourism and recreation associated with coral reefs (Johns et al. 2003a, 2003b; Leeworthy & Bowker 1997; Leeworthy &
Wiley 1996/1997; Leeworthy et al 2004).
2.4.7.2 Expectancy-discrepancy analysis.
Expectancy-discrepancy theory suggests that satisfaction is a measure of how closely a consumer's desired experiential
outcome is to that consumer's perceived reality once the activity takes place (Vroom 1964; Porter & Lawler 1968). When
perceptions meet or exceed expectations, consumers tend to be more satisfied (Manning 1999). Two consumers may receive the
same service at the same time and place, but experience very different satisfaction levels due to their expectations. Expectancy-
discrepancy analysis was used by Loomis et al. (2008a, 2008b, 2008c) in their valuation of recreational fishing, diving, and
snorkeling in the Florida Keys.
2.4.7.3 Norm curves.
Social norms are the rules of behavior that coordinate interactions within a society or group. Normative theory involves
identifying the standards that individuals and groups use to evaluate behavior and social/environmental conditions. Jackson
(1965) developed a methodology — return-potential curves — to measure norms. The methodology involves using stakeholder
surveys to measure respondent normative evaluations of varying levels of indicators. The personal norms of individuals can
then be aggregated to test for the existence of social norms. Social norm curves (Manning et al. 1999) can be used to determine
the level at which indicator values shift from acceptable to unacceptable conditions.
Normative theory and methods have been used to formulate standards of quality for outdoor recreation, including issues related
to crowding (Shelby 1981; Heberlein et al. 1986; Whitaker & Shelby 1988; Patterson & Hammitt 1990; Williams et al. 1991;
Vaske et al. 1986; Manning et al. 1996; Jacobi & Manning 1999), ecological impacts at campsites (Shelby et al. 1988), and
wildlife management practices (Vaske & Donnelly 1988). Researchers have begun using photographs and videos to represent
levels of impacts (Vaske et al. 1996; Manning et al. 1999). Visual representations can be used to effectively "tell a story" and
resonate well with stakeholders.
2.4.7.4 Entergy.
Emergy analysis is another way to quantify ecosystem services. Odum (1996) defined emergy as "the available solar energy
used up directly and indirectly to make a service or product". After converting energies of different kinds to the same kind of
energy (i.e., solar joules, sej) by multiplying each by the appropriate emergy per unit factor (sej/J), the value of an ecological or
economic product or service is determined by summing the inputs. Emergy has units of solar emjoules denoting that it is an
accumulation of available energy used in the past. The emergy of an ecosystem product, such as fish, can be converted to a
monetary value by dividing by the emergy to dollar ratio of the economy in which the item was sold. The emergy to money
ratio is the total emergy flows supporting an economy divided by the dollar flow of the GDP of the economy. Emdollars then
redistribute the monetary flow of the system in proportion to the emergy flows, thereby assigning each item the portion of the
total buying power in the system attributable to it based on its emergy relative to the emergy of all other products and services
in the system's economy including those not counted by economic measures. Emdollar values can be compared to dollar values
of the same item to determine the unvalued work of nature that was required for an item compared to the human services
required for the same item.
2.5 Reflections
There is considerable variation in the way coral reef based tourism and recreation is defined, measured, and valued. Generally,
existing definitions are incomplete and not rigorously developed. Linking the attributes to the ecosystem service should be a
step towards standardization. While there are regional differences as to which sectors are operative and at what level, if we
develop an appropriate classification framework, that variability should not matter.
We have a fundamental understanding of the factors that affect delivery of the tourism and recreation ecosystem service. We
need to improve our understanding of the cumulative and synergistic effects of multiple stressors in both a spatial and temporal
context. The application of landscape ecology approaches and metrics to understand coral reef ecosystem functions and to
assess the impacts of a variety of management activities (both terrestrial and aquatic) may contribute significantly to our
understanding.
We also need to develop, test, and refine indicators that are sufficiently sensitive to distinguish the effects of human disturbance
from those of natural variability and that can serve as quantitative estimates of services provided. Ecological models can help to
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illustrate and quantify relationships among environmental and ecological reef elements and can be used to investigate
thresholds for reef persistence and sustainable delivery of services.
Better understanding of consumer preferences vis-a-vis the attributes would be helpful.
• The colloquial literature, including marketing materials, may reflect what people are actually looking for from their
tourism experience but does not quantify its value.
• NOAA has done a fairly comprehensive analysis of consumer preferences for the Florida Keys National Marine
Sanctuary, although the survey questionnaire could be refined somewhat based upon the attributes work.
• We need the same type of statistically valid information for USVI and Puerto Rico.
From a valuation perspective, the Tourism Satellite Accounts could contribute significantly towards the valuation of coral reef
ecosystem services, if refined both from a scale perspective (need to have state or finer scale TSAs), and in a manner that can
link them directly to reef-based tourism (need to be able to separate out tourism associated with the coral reef from nonreef-
based tourism). If the consumer preference surveys are properly designed, they could help quantify the latter and fill in the gaps
with nonmarket values.
Throughout this chapter, we have been using the commonly used term "tourism and recreation" as the ecosystem service.
However, more precise terminology is needed to facilitate the interaction between ecological assessment and economic
valuation of changes in ecosystem goods and services (Munns personal communication). One approach would be to distinguish
between final and intermediate ecosystem services (Boyd & Banzhaf 2007; Daily & Matson 2008). Final ecosystem services
are the components of nature, directly enjoyed, consumed, or used to yield human well-being. These are described in units upon
which accounting systems and valuation can be based. Intermediate ecosystem services are the components of nature that are
not directly enjoyed, consumed or used to yield human well-being, but that are important for the production of final ecosystem
services. Final ecosystem services are the units upon which valuation will be based. It is also important to understand
intermediate services, because their relationship to final services is of great importance in understanding, assessing, predicting
and managing final services and the human well-being provided (Ringold et al. 2009). Clearly differentiating between final
ecosystem services and intermediate ecosystem services precludes "double counting" (Boyd 2008).
Within "tourism and recreation" there is a suite of final ecosystem services. We are defining these as "opportunities", since the
ecosystem does not actually provide tourism and recreation, but rather provides the opportunity for humans to enjoy
recreational experiences. We have identified the following final ecosystem services:
• Recreational fishing opportunity
• Recreational diving/snorkeling opportunity
• Recreational underwater photography opportunity
• Recreational surfing opportunity
• Opportunity to view nature and wildlife
• Opportunity to sunbath and swim at the beach
• Opportunity to collect objects (beachcombing)
Ecosystem production functions describe the relationship between intermediate ecosystem services and final ecosystem
services. The intermediate ecosystem services that support tourism and recreation include:
• Production of benthic and aquatic prey for consumption by recreational fish
• Coral reef formation and maintenance
• Maintenance of water quality
• Maintenance of reef breaks
• Maintenance of biological integrity and biodiversity
• Sand production
Table 2-5 illustrates the ecosystem services (final and intermediate), as well as those features that support them. The table
serves to help address the following question: What biophysical metrics directly facilitate the integration of biophysical
measurement, analysis, and models with analyses of the social and economic benefits derived from ecosystem goods and
services?
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Table 2-5. Final ecosystem services and supporting features for tourism and recreation
Ecosystem Service(s)
Final (FES)
Intermediate
Natural Features
Social Values
_ , . Ecosystem- Potential
Complementary Derived Indicators of Final
Goods & Services
Benefits Ecosystem
Service(s)
Tourism & Recreation
P d t' f
_ .. , „. , . benthic and aquatic
Recreational Fishing ,,
_ prey tor
Opportunity .
consumption by
Recreational
Diving/Snorkeling
Opportunity
Recreational
Underwater
Photography
Opportunity
recreational fish
Coral reef formation
& maintenance;
maintenance of
water clarity;
production of ben-
thic and aquatic prey
for consumption by
recreational fish
Coral reef formation
& maintenance;
water clarity;
production of
benthic and aquatic
consumption by
recreational fish
Recreational Surfing „ „, .
_ . Reef breaks
Opportunity
Opportunity to View „. . ...
." , .,,., „.„ Biological integrity
Nature and Wildlife ° 6 J
Opportunity to
Sunbath and Swim at
the Beach
Water quality,
shoreline protection,
sand production
Opportunity to
Collect Objects Water quality
(Beachcombing)
Fish diversity and
abundance
Coral diversity,
abundance and
health; fish
diversity and
abundance; water
clarity
Coral diversity,
abundance and
health; fish
diversity and
abundance; water
clarity
3-D reef structure
Biodiversity
(birds, marine
mammals, turtles)
White coralline
sands; calm
waters
Wide sandy
beaches,
biodiversity,
occasional storms
Desirability offish , .
, . Adequate
species and size . ,, .,
,, , , , infrastructure (boats,
for rod-and-reel . .
, marinas, etc.)
catches
Revenues from , . . -
, Abundance ot
tourism and . . .
catchable snappers
recreation .
. . . and groupers
activities
Desirability of Access to reef, safe
coral reef for swimming conditions, Revenues from
recreation based adequate tourism and
on physical infrastructure (hotels, recreation
appearance (color, dive boat operators, activities
visibility, etc.)
etc.)
Desirability of Access to reef, safe
Taxa richness, size
and density of reef
organisms
coral reef for swimming conditions, Revenues from „ . ,
... . , Taxa richness, size
recreation based adequate tourism and , , ,, ,,
... . „ . ,. . . , . and density ot reet
on physical infrastructure (hotels, recreation .
. , ,. , , ,. ... organisms
appearance (color, dive boat operators, activities
visibility, etc.) etc.)
Desirability based
on wave size and
Desirability of
species (rarity,
size)
Desirability of
coralline sand
beach for
sunbathing (size,
cleanliness,
appearance)
Access to reef,
adequate
infrastructure (hotels,
board shops, etc.)
Revenues from
tourism and
recreation
activities
3-D structure and
proximity to deep
Access to reef and Revenues from „ . .
, , 1 axa richness,
adequate tourism and -
. „ * ., .. presence ot
infrastructure (boats, recreation . _
. v specific species
tour guides) | activities |
Access to beach
Areal extent of
Revenues from beach, color of
tourism and beach, water
recreation temperature, days
activities of sunshine, beach
trash
Areal extent of
Desirability of _ ,, . . „
„ . , . Revenues trom beach, frequency
walking on beach , . , ,, ,
, ° .. . . . tourism and of storms,
and ot finding Access to beach , . . ., „ „
. . - , „ recreation proximity of reef,
beautiful & ..... r . ,
, , . . activities taxa richness ot
unusual obiects . .
invertebrates
Definitions (proposed by the Ecosystem Services Research Program and currently under discussion by the Program)
• Final Ecosystem Service - Output of ecological functions or processes that directly contributes to social welfare or has the potential to do so
in the future (broadly based on Boyd & Banzhaff [2007]).
• Intermediate Ecosystem Service - Output of ecological functions or processes that indirectly contributes to social welfare or has the potential
to do so in the future.
• Natural Features - The biological, chemical, and physical attributes of an ecosystem or environment.
• Social Values - The social attributes that influence economic demand for an ecosystem service.
• Complementary Goods & Services - Inputs (usually built infrastructure or location characteristics) that allow a good or service to be used by
complementing the ecological condition. For example, complementary goods and services that allow the presence of fishable fish to become
an opportunity for recreational fishing will include aspects of site accessibility, such as road access, available parking and the presence of a
fishing pier, all of which make fishing at the site possible and enhance enjoyment of the activity.
• Ecosystem-Derived Benefits - The contribution to social welfare of ecosystem goods and services. In the ESRP, the term applies specifically
to net improvements in social welfare that result from changes in the quantity or quality of ecosystem goods and services attributable to
policy or environmental decisions.
• Indicator of Final Ecosystem Service - Biophysical feature, quantity, or quality that requires little further translation to make clear its
relevance to human well-being (i.e., "public-friendly" measurement)
-------
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-------
Appendix 2-A
Studies quantifying tourism and recreation
The following table lists studies and the endpoint they used to quantify tourism and recreation. Also included are the biological
reef attributes and the physical and socioeconomic variables used to estimate that endpoint. The method and location of the
studies are also given.
Reef
attributes
Physical Socioeconomic
variables variables
Method
Location
Citation
Final Ecosystem Service: Recreational Diving/Snorkeling Opportunity
Coral cover
Coral bleaching; coral
cover; fish species
composition
(preference for more
colorful reef fish)
Coral cover,
abundance offish
Coral biodiversity,
coral cover, fish
biodiversity, fish stock,
algse cover
"Pristine" coral reef
condition; coral
diversity; fish diversity
and abundance
Fish abundance, coral
health
Coral reef and coral
health
Coral quality
Coral diversity
Diversity of colorful
marine life
Amount of live coral,
fish and sea life
diversity, fish
abundance
Coastline development
on shoreline, water number of boats,
clarity, plastic debris, presence of MPA,
risk of contracting ear number of snorkelers,
infection by swimming fee
in polluted water
. . Swimming restrictions;
Water clarity 6, ...
payment vehicle
Water clarity
Perceptions of
crowding (social
carrying capacity),
ease of access
, . Number of divers per
Water clarity .
J site per year
Clean and odor-free
Water clarity water; crowd-free
experience
, . Access and availability
Water clarity „„ .....
J ot facilities
Survey
questionnaire
Survey
questionnaire
Survey
questionnaire
Model
Two survey questionnaires -
tourist exit survey and tourist
operator survey; face-to-face
interviews of 400 households
for recreational fishing;
three focus groups
Survey questionnaire and
pictures of alternative scenarios
Survey
questionnaire
Survey
questionnaire
Summary
article
Survey questionnaire
& photo analysis
Survey
questionnaire
Survey
questionnaire
Bolinao,
Philippines
Zanzibar
&
Mafia
Tobago
Kihei Coast,
Hawaii
Bermuda
Florida
Keys
Seychelles
Phi Phi Islands,
Thailand
Bonaire
Great Barrier
Reef,
Australia
Florida
Keys
Ahmed et al.
(2007)
Andersson
(2007)
Beharry-Borg &
Scarpa(2010)
van Beukering
& Cesar (2004)
van Beukering
etal. (2010)
Bhat (2003)
Cesar et al.
(2004)
Christiernsson
(2003)
Davis & Tisdell
(1996)
Dixon et al.
(2000)
Hajkowicz
(2006)
Leeworthy &
Wiley (1996)
-------
Reef
attributes
Physical
variables
Socioeconomic
variables
Method
Final Ecosystem Service: Recreational Diving/Snorkeling Opportunity
Amount of live coral,
fish and sea life
diversity, fish
abundance
Amount of live coral,
fish and sea life
diversity, fish
abundance
Coral health, fish
abundance, large fish,
coral cover
Species richness,
density and average
size of organisms
Health of corals
Species diversity (fish
and corals), percent
coral cover
Percent coral cover
Coral reef biodiversity
(inferred from other
study)
Amount of marine life
Coral biodiversity
Coral growth
Aesthetic beauty of
dive site (not defined
further)
Coral diversity,
abundance and health;
fish diversity and
abundance
Coral bleaching,
"marine life"
Abundance and
diversity of corals
and fish
Water clarity
Water clarity
(visibility)
Current WQ
Visibility
(water clarity)
Access to reef, safe
swimming conditions,
water quality
Warm temperatures,
clear waters, beach
characteristics
Water clarity
(visibility)
Access and availability
of facilities
Access and availability
of facilities
Perceptions of
crowding (social
carrying capacity),
knowledge of
ecosystem, experience
snorkeling
Dive tag price
number of divers at a
site at any one time
Adequate infra-
structure (hotels, dive
boat operators, etc.)
Low health risks
Entrance fee
Survey
questionnaire
Survey
questionnaire
Self-administered
questionnaires
Survey
questionnaire
Survey
questionnaire
Model
Interviews (survey) of
municipal fishers, gleaners,
seaweed farmers, and tourism
business operators
Survey
questionnaire
In-person interviews using
open-ended elicitation
questions
Survey
questionnaire
Survey
questionnaire
Questionnaire;
four different surveys,
secondary data sources
Video of alternative diving
sites; survey questionnaire
Location
(con 't)
Florida
Keys
Florida
Keys
South Sinai,
Egypt
Jervis Bay,
Australia
Florida
Keys
Bonaire
National
Marine Park
Roatan,
Honduras
Bohol Marine
Triangle,
Philippines
Worldwide ?
Jamaica
&
Curacao
American
Samoa
Similan Islands,
Thailand
Bonaire
&
Barbados
Indian Ocean
(Tanzania/Kenya
& Maldives/
Sri Lanka)
Eilat, Israeli
Red Sea
Citation
Leeworthy &
Wiley (1997)
Leeworthy et al.
(2004)
Leujak &
Ormond (2007)
Lynch et al.
(2004)
Park et al.
(2002)
Parsons & Thur
(2007)
Pendleton
(1994)
Samonte-Tan et
al. (2007)
Sorice et al.
(2007)
Spash (2000)
Spurgeon et al.
(2004)
Tapsuwan &
Asafu-Adjaye
(2008)
Uyarra et al.
(2005)
Westmacott et
al. (2000)
Wielgus et al.
(2003)
-------
Reef
attributes
Physical Socioeconomic
variables variables
Method
Location
Citation
Final Ecosystem Service: Recreational Dlvlng/Snorkellng Opportunity (con 't)
Abundance and variety
of fishes, number of
"unusual", and number
of "large" fish
Interview
Jamaica
Williams &
Polunin (2002)
Final Ecosystem Service: Recreational Fishing Opportunity
Target sportfishing
species (bonefish,
permit, tarpon)
Target sportfishing
species abundance and
diversity
Target sportfishing
species (80 fish and
invertebrate species)
Clean and odor-free
water; crowd-free
experience
Survey
questionnaire
Belize
_ Great Barrier
S"rV£y. Reef,
questionnaire
Australia
Jervis Bay,
Australia
Fedler & Hayes
(2008)
Hajkowicz
(2006)
Lynch et al.
(2004)
Final Ecosystem Service: Beach Recreation Opportunity
Quality of beaches
Quality of beaches
Quality of beaches
Large wildlife
(manatees, whales,
dolphins, sea turtles)
Final Ecosystem Service: Wildlife
Survey
questionnaire
Survey
questionnaire
Survey
questionnaire
Viewing Opportunity
Survey
questionnaire
Florida
Keys
Florida
Keys
Florida
Keys
Florida
Keys
Leeworthy &
Wiley (1996)
Leeworthy &
Wiley (1997)
Leeworthy et al.
(2004)
Leeworthy &
Wiley (1997)
-------
-------
3
Coral Reefs: Fish & Coral for
Food, Aquaria, & Ornaments
All oceans are affected by humans to various degrees, with overfishing
having the most widespread and dominant direct impact on food
provisioning services that will affect future generations
(MEA 2005)
3.1 Describing the service
3.1.1 Definitions
A fishery is an entity engaged in harvesting fish, which is typically defined in terms of the people involved, species or type of
fish, area of water or seabed, method of fishing, class of boats, purpose of the activities or a combination of these identifiers
(FAO2010).
Although a fish in the strictest sense is an aquatic vertebrate with fins, the term fisheries is used more broadly, to include
mollusks, crustaceans, or any other aquatic animals (and in a few cases, aquatic plants) that are harvested. These include, but
are not limited to, oysters, scallops, conch, squid, octopus, lobster, shrimp, and kelp.
Fisheries are most often associated with food production, but harvest of aquatic organisms can be for other purposes, such as
for sale as live aquarium fish, bait, or fish meal for agriculture. Other uses include harvest of shells and skeletons for curios and
jewelry (Table 3-1).
Food production from coral reefs includes both fish and invertebrate organisms that are harvested on the reef or spend at least a
portion of their life cycle on the reef and are harvested elsewhere.
Food products are derived from commercial fishing (for profit), subsistence fishing (for survival), or recreational fishing (for
pleasure). Because the benefit of recreational (sport) fishing is pleasure, it is discussed in the Tourism and Recreation section.
Artisanal fishing is a term that usually describes traditional harvesting techniques (e.g., rod and tackle, spear, throw-net) that
are more likely to represent a small-scale, low-intensity (usually subsistence) fishery. Although coastal fisheries in many parts
of the world are mostly artisanal, some dominate the catches of some species or particular year/size classes.
Table 3-1. Benefits or amenities derived from fish production
Seafood for human consumption (fish & invertebrates)
Live fish and coral for aquariums (Chan & Sadovy 2000)
Shells and skeletons for ornamental art and jewelry
Recreational fishing for pleasure (Brander et al. 2007; NOAA 2009)
Human health and well-being (Olsen et al. 1984; WHO 2010)
Fish meal and oil for livestock and aquaculture feed
3.1.2 Benefits of coral reef fisheries
3.1.2.1 Seafood.
The most commonly described and most highly valued benefit of coral reef fisheries is food for human sustenance. In general,
humans are obtaining their protein from fish, both marine and freshwater, in ever-increasing numbers. After the remarkable
increase in both marine and inland capture of fish during the 1950s and 1960s, world fisheries production has leveled off since
the 1970s. This leveling of the total catch follows the general trend of most of the world's fishing areas, which have apparently
reached their maximum potential for fisheries production, because the majority of stocks have been fully exploited. Therefore,
it is very unlikely that substantial increases in total catch will be possible in the future.
The total food fish supply and hence consumption has been growing at a rate of 3.6% per year since 1961, while the world's
population has been expanding at 1.8% per year. The proteins derived from fish, crustaceans, and mollusks account for between
13.8% and 16.5% of the animal protein intake of the human population. The average apparent global per capita consumption
increased from about 9 kg per year in the early 1960s to 16 kg per year in 1997, nearly doubling in 40 years (FAO 2002).
Currently, two-thirds of the total food fish supply is obtained from capture fisheries in marine and inland waters, while the
remaining one-third is derived from aquaculture.
-------
Typically, fish provide about 20-30 kcal per person per day. In a few countries, such as Iceland, Japan, and some small island
states, where there are few alternative proteins or the people have a strong preference for fish, fish can contribute up to 180 kcal
per person per day. Fish proteins are essential in the diet of some densely populated countries where the total protein intake
level is low. Worldwide, about a billion people rely on fish as their main source of animal protein. Dependence on fish is
usually higher in coastal than in inland areas. About 20% of the world's population derives at least 20% of its animal protein
intake from fish, and some small island states depend almost exclusively on fish. Artisanal coral reef fisheries provide an
inexpensive source of protein and employment where few alternatives exist (Burke & Maidens 2004). For example, Seychelles
has one of the highest per capita consumption rates offish in the world (65 kg per year) with 900-1,000 artisanal fishers fishing
full-time at 35 landing sites (Cesar et al. 2004).
The current annual global harvest from tropical reef fisheries has been estimated at six million metric tons (Polunin & Roberts
1996). Some scientists have estimated that worldwide coral reefs could produce a sustainable fisheries yield of 20m-35m
metric tons peryear (Crossland et al. 1991; Hatcher et al. 1987); however, Birkelund (1997) argues that coral reefs cannot or
should not sustain such large fisheries yields.
3.1.2.2 Live fish and coral for aquaria.
Another specialized coral reef fishery is the collection of live fish and coral for aquaria (Livengood & Chapman 2007). The
United States is the single largest importer of ornamental fish in the world, but the European Union is the largest market for
ornamental fish (FAO 1996-2005; Chapman 2000). Estimates of the magnitude and value of the aquaria fishery vary widely.
Livengood and Chapman (2007) estimate the value of ornamental fish and invertebrates imported into different countries
worldwide at $278m. Chan and Sadovy (2000) conducted a survey of marine aquarium shops in Hong Kong and estimated that
close to a million individual coral reef fish enter the aquarium trade annually with an average value of about HK$60 (~$8) per
fish. Bruckner (2005) estimated the market to be 14m-30m fish per year, with an import value of $28m-$44m.
3.1.2.3 Ornaments and jewelry.
Species of "precious coral" such as red and pink corals (Family Coralliidas), black corals (Order Antipatharia) and gold corals
(Family Parazoanthidas) have historically been harvested from many parts of the world for high end jewelry and beads
(Hourigan 2008; Grigg 1984, 1989). It has been estimated that precious coral catch reached roughly 450 tons per year in the
1980s and has now declined to roughly 50 tons per year (Oceana 2010). The United States and the European Union proposed
thirty-one species of the family Coralliidas (Corallium spp. and Paracorallium spp.) for inclusion in the Convention on
International Trade in Endangered Species (CITES) Appendix II (Oceana 2010). However, the measure failed to reach the two-
thirds majority at the 2010 CITES conference in Doha, Qatar (ENS 2010).
3.2 Providing the service
3.2.1 Linkages among reef condition, reef structures, and reef functions with respect to delivery of fishes for
aquaria and food
Almost a third of the world's marine fish species are found on coral reefs (Moberg & Folke 1999). Reefs provide essential
habitat for adult fish, and their physical structure creates quiet water areas necessary for seagrass and mangrove nurseries.
Valued open water commercial fish such as groupers, snappers, grunts, and barracuda spend critical life stages on the reef and
in the reef-seagrass-mangrove system (Mumby et al. 2004, 2008; Mumby 2004; Dorenbosch et al. 2004). Some reefs in the
Pacific and Indian Oceans are extremely rich in species (Tibbetts 2004); for example, Pereira (2000) lists 794 species in 93
families of reef-associated fishes that live in Mozambique waters.
3.2.1.1 Scleractinian corals.
Coral reefs are composed of a physical infrastructure that provides essential fish habitat. The infrastructure is constructed by
reef-building scleractinians, or "stony corals". These corals are distinguished by their ability to secrete an extracellular calcium
carbonate (aragonite) skeleton, which in most cases forms a solid, relatively permanent reef structure. To form the skeleton,
scleractinians rely on symbiotic algas to produce energy for skeletal growth through photosynthesis. Capturing the energy from
sunlight is one reason that stony corals are usually found in shallow, transparent waters. The durability of stony coral skeletons
provides an enduring reef habitat that allows the evolution of complex reef communities that include harvestable fish and
invertebrates. Dahl (1973, p. 240) stated:
The production, occupation, and destruction of surface area are, therefore, basic reef processes, and the balance
between them is an essential aspect of the reef ecosystem. The efficient production of surface is a primary function of
many reef organisms, and the control of surface by secondary occupants is a basic competitive force and a major
determinant of reef communities.
Scleractinians provide enormous reef surface area. Although slow-growing by most standards, scleractinian corals can live for
hundreds of years, and some species can grow to the size of an automobile (WS Fisher, personal communication).
In the Caribbean, the dominant, large reef-building corals are in the Montastraea genus, which includes M. cavernosa and three
closely related species ((M. annularis, M. faveolata, and M. franksii) that are often referred to as the Montastraea complex.
-------
Montastraea colonies are often found on the fore-reef and appear particularly critical to the biodiversity of fish and
invertebrates and for maintaining the structure, function, and flow of reef services (Mumby et al. 2008; Beets & Friedlander
1998). The Caribbean acroporid species, Acropora palmata and A. cervicornis are also relatively large and are highly branched,
which provides additional surface area and greater reef complexity. Unfortunately, these species are vulnerable to elevated
water temperatures that would result from climate warming and white-band disease (Aronson & Precht 2001). Because of
enormous losses suffered by acroporids in the last three decades, both of these critical species are listed as threatened. As might
be expected, healthy stony corals appear to be critical to productive fisheries. Fish productivity, species richness, fish biomass,
and potential yield have all been reported to decrease with a decline in stony coral health (Warren-Rhodes et al. 2003).
Mumby et al. (2008) used 11 classes of reef habitat as surrogates for species, functions, and ecosystem services and found that
one-fourth to one-third of benthic invertebrates and fish occurred in the Montastraea-dominated fore-reefs, which consistently
had the highest richness, the highest number of processes, and the most services. Yet only 10% offish species functional
classes were unique to any one habitat. Functional classes of fish were an effective surrogate for total fish and benthic species
richness, and the representation of species or functional classes ensured inclusion of all processes and services in the design of a
reserve network. This research suggests that using the number of fish functional groups as a proxy indicator for benthic
richness may be helpful in managing reef functions, services, and biodiversity for maintaining the resilience of reefs.
3.2.1.2 Reef structural complexity.
The physical structure of reef habitat influences the biodiversity and ecosystem functions of a reef community (Alvarez-Filip et
al. 2009). In general, more complex habitats facilitate species coexistence through niche partitioning and provision of spatially
delineated refuge from predators (Beets & Friedlander 1998; Bruno & Bertness 2001). Physical structures influence the spaces
that are inhabited by organisms by defining volume, orientation, accessibility, water residence time, and food availability,
among other factors (Scheffers et al. 2003). The rich diversity of coral reefs rests partly in the provision of habitable surface
area and partly in the variability of that surface area.
There has been a widespread decline in the health of coral reefs that has reduced the amount and complexity of the available
habitat for fish. A potential consequence of this decline, referred to as "reef flattening" (Alvarez-Filip et al. 2009), is the loss of
species richness and abundance of reef fishes and invertebrates (Gratwicke & Speight 2005; Idjadi & Edmunds 2006; Wilson et
al. 2007).
For coral reefs, there appears to be a strong positive correlation of habitat complexity to fish species richness (Walker et al.
2009; Pittman et a/.2007). Rugosity (an indicator of habitat complexity) has, therefore, been used successfully in the Virgin
Islands as an index offish diversity and in data-poor areas may be used to spatially assess where areas of high fish species
richness may occur. Studies on the recruitment behavior of epibenthic communities have also shown that substrate irregularity
may encourage the diversity of initial substrate colonizers, which may result in higher diversities later in succession (Breitburg
1985). Habitat complexity, especially appropriately-sized holes or cover for a particular species, provide shelter from predators
(Hixon & Beets 1993; Roberts & Ormond 1987; Friedlander & Parrish 1998; Aguilar-Perera & Appeldoorn 2008).
3.2.1.3 Seascape connectivity.
Along with coral reefs, seagrass meadows and mangrove forests combine in a complex and dynamic mosaic that provides
critical foraging areas, nurseries, and refugia for fish and invertebrates (Christensen et al. 2003; Aguilar-Perera & Appeldoorn
2007; McField & Richards Kramer 2007). Some commercially important species and threatened species, such as the rainbow
parrot fish, utilize mangroves exclusively as nursery habitat, and the biomass of other fish is significantly increased when
mangrove habitat is available (Mumby et al. 2004, 2008; Meynecke et al. 2008). Functional dependency of some fish on
specific habitats, like the mangrove-dependent rainbow parrotfish, can also make them more vulnerable to extinction (Mumby
et al. 2004). Proximity to seagrass and mangrove nursery habitat and the connectivity of reefs with nursery habitat has been
measured using a variety of landscape connectivity metrics (Dorenbosch et al. 2004; Meynecke et al. 2008; Mumby 2006;
Edwards et al. 2010).
Recent studies have directly compared the value of seagrass and mangrove habitats with the value of other possible shallow
water habitats (Dorenbosch et al. 2006). Nagelkerken et al. (2000) used a visual survey technique to evaluate the importance of
mangroves, coral reefs, and seagrasses as habitat for juvenile fishes in Bonaire. Their study showed that all three habitats were
important nursery areas, but for different species. A similar study in the Indo-Pacific (Dorenbosch et al. 2006) documented
ontogenetic shifts from juvenile habitats (seagrasses and mangroves) towards adult habitats (coral reefs). In a multi-year study
in La Parguera, Puerto Rico, Pittman et al. (2010) found a high degree of multi-habitat use, with size-dependent ontogenetic
habitat shifts. The importance of different habitats for juveniles and adult fishes is shown in Table 3-2.
-------
Table 3-2. Important (*) habitats for juveniles and adults of selected fish species
H indicates the most important habitat for juveniles
Acanthurus chirurgus
(doctorfish)
Chcetodon capistratus
(foureye butterflyfish)
Cheilinus undulatas
(humphead wrasse)
Hcemulonflavolineatum
(French grunt)
Hcemulon plumierii
(white grunt)
Hcemulon sciurus
(bluestriped grunt)
Lutjanus apodus
(schoolmaster snapper)
Lutjanus griseus
(gray snapper)
Lutjanus mahogoni
(mahogany snapper)
Lutjanus synagris
(lane snapper)
Ocyurus chrysurus
(yellowtail snapper)
Scarus guacamaia
(rainbow parrotfish)
Sparisoma radians
(bucktooth parrotfish)
Sparisoma viride
(stoplight parrotfish)
Sphyrcena barracuda
(great barracuda)
Seagrass
H
X.
H
H
X.
H
H
X.
X.
X.
X.
X.
X.
X.
H
X.
-
X.
H
X.
Juveniles
Mangrove
-
H
-
X.
H
X.
X.
H
H
H
H
H
H
H
-
-
H
X.
-
H
Coral Reefs
X.
-
-
X.
X.
X.
-
-
-
X.
-
-
X.
H
-
X.
-
-
X.
-
Seagrass
-
-
-
-
-
-
-
-
-
X.
-
-
-
-
-
X.
-
X.
-
X.
Adults
Mangrove
-
-
-
-
-
-
-
X.
X.
X.
X.
X.
X.
-
-
-
-
X.
-
X.
Coral Reefs
X.
X.
X.
X.
X.
X.
X.
X.
X.
X.
-
-
X.
-
X.
X.
X.
-
X.
X.
Study
Location
Bonaire
Bonaire
Indian Ocean
Bonaire
Puerto Rico
Puerto Rico
Bonaire
Puerto Rico
Bonaire
Puerto Rico
Bonaire
Puerto Rico
Puerto Rico
Puerto Rico
Bonaire
USVI
Indian Ocean
Puerto Rico
Bonaire
Bonaire
D— f
N
N
D
N
P
P
N
P
N
P
N
P
P
P
N
P
D
P
N
N
References: D: Dorenbosch et al. (2006) N: Nagelkerken et al. (2000) P: Pittman et al. (2010)
Source: Partially adapted from Nagelkerken et al. (2000).
3.2.2 Linkages among reef condition, reef structures, and reef functions with respect to provision of stony corals,
black corals and precious corals for aquaria stock and jewelry
Corals collected for the aquarium and jewelry industries generally are rare, slow-growing, long-lived species (USFWS 2011).
According to the Global Marine Aquarium Database (GMAD), there are 61 species of soft corals and 140 species of stony
corals collected for aquaria (Wabnitz et al. 2003). Of the approximately 220 species of precious corals (those used in the
jewelry industry), only about 16 are commercially important (Hanfee 1997). Precious corals are found in deeper water (250-
1600 feet depth) and include red and pink corals, black corals, gold corals and bamboo corals (Tsounis et al. 2010).
Coral condition is an important characteristic of corals collected for both the aquarium and jewelry industries. Corals collected
for the aquaria trade must be healthy enough to survive collection and transport. Precious corals must be healthy enough that
their skeletons can be formed into jewelry. It is not known how structural complexity or seascape connectivity relate to corals
for aquaria stock and jewelry.
-------
3.3 Measuring the service
3.3.1 Fish
A number of indicators of fisheries production are directly monitored as attributes of reef condition, including fish abundance,
fish size, conch abundance, lobster abundance, and the biomass of commercially important species (McField & Richards
Kramer 2007; Healthy Reefs Initiative 2010). Reef fish surveys are one method of measuring the available fish biomass. A
variety of standardized methods has been developed and implemented (Bohnsack & Bannerol 1986; McField & Richards
Kramer 2007; Schmitt & Sullivan 1996; Pattengill-Semens & Semens 2003). Divers swim along transects, estimating the
number and size ranges of fish species. Several metrics that can be used as indicators of stock status can be derived from the
data (e.g., species abundance, density, size structure, and frequency-of-occurrence; total fish biomass; commercially significant
fish biomass) (Ault et al. 1998; Paddack et al. 2009; McField & Richards Kramer 2007). Paddack et al. (2009) conducted a
meta-analysis of reef fish density obtained from 48 studies covering 318 reefs across the Caribbean and found that overall reef
fish density has been declining significantly for more than a decade, at rates that are consistent across all subregions of the
Caribbean basin (6%-22.7% per year and in three of six trophic groups). There appears to be a considerable lag-time between
degradation of coral reef habitat and the decline of fish populations; however, a consistent significant decline across several
trophic groups and among both fished and nonfished species indicates that Caribbean fishes have begun to respond negatively
to habitat degradation (Paddack et al. 2009).
Fishery-dependent population estimates can also be developed using catch data (e.g., the National Marine Fisheries Service
(NMFS) headboat catch and effort data). The NMFS data provide total numbers of individual fish in the catch as well as total
weight in the catch by species by year (Ault et al. 1998).
3.3.2 Stony corals
Despite the fact that more stony corals (greater abundance) and larger corals provide greater habitat for fish and invertebrates
(Beets & Friedlander 1998), few studies are available that directly measure coral surface area. Most commonly, stony coral
studies measure "live coral cover", which reflects only a 2-dimensional planar area viewed from above the coral. The planar
approach is convenient but does not account for colony height, which can vary widely, and cannot be used to estimate colony
surface area (quantity of habitat). The concept of 3-dimensional (3D) colony surface area has been explored (Dahl 1973;
Szmant-Froehch 1985; Roberts & Ormond 1987; Babcock 1991; Alcala & Vogt 1997; Bak & Meesters 1998), but only recently
have 3D colony surface area methods been developed for use in applied field studies (Fisher 2007; Fisher et al. 2007; Fisher et
al. 2008; The Nature Conservancy 2010).
3.3.3 Reef structural complexity
The structural complexity of reefs is determined by size, shape, and juxtaposition. Complexity refers not only to the surface
area but also to the size variability of spaces, which provide different habitats for organisms with different sizes and behaviors.
A complexity index (Aronson et al. 1994) and a similar rugosity index (NOAA 2008) are calculated from comparison of the
length of a chain to the distance covered by the chain when draped over a coral reef (Risk 1972; Rogers et al. 1982; Connell &
Jones 1991). Despite the intention to measure multiple aspects of reef complexity, this approach only measures the cumulative
height of coral colonies on a reef, which is only one component of complexity. Since reef height reflects greater surface area,
this is a useful measurement for predicting habitat availability. Another component of complexity, however, is variability in
spaces, which is created by corals of different sizes and holes or caves in coral structures. In a recent study, Fisher et al. (2008)
calculated the coefficient of variation of colony size to reflect this component of complexity. Others have developed tools to
estimate holes and caves in coral structures (Scheffers et al. 2003). While all of these approaches are useful, no truly
comprehensive method has yet been presented to indicate reef complexity.
3.3.4 Seascape mapping
Despite the fact that the earliest landscape ecology studies of marine systems had their beginnings in the classic works of
Levins (1969) and Levin and Paine (1974), it is only within the last decade or so that we have seen a significant increase in the
application of landscape ecology principles to seascapes. Early work concentrated on mapping habitats and understanding reef
functions and processes (Dierssen et al. 2003; Kvernevik et al. 2002). Even though this work continues, there has been recent
growth in the availability of spatial data from GIS (geographic information system) and remote sensing technologies, which are
necessary to map coral reef habitats (Phinn et al. 2008). This, in combination with survey and monitoring data of reef attributes,
has led to methods that increase the data resolution needed to make meaningful observations at more local scales (Harbourne et
al. 2006).
These developments have also inspired work on spatially explicit modeling and mapping offish distributions and fish
production (and service provisioning) on reefs (Pittman et al. 2010; Pittman et al. 2007; Mumby et al. 2008; Purkis et al. 2008).
Although mapping of services is in its infancy, trade-offs in managing services result in changes in the location or scale of the
beneficiaries (e.g., local fisheries or jobs versus global tourists), which we are only beginning to understand (Hodgson & Dixon
2000). The connectivity of coral reef fish habitat and nursery habitat is important to fish production, so methods for mapping
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these habitats at finer scales and methods to measure the connectivity of habitat used throughout the life cycles of fishes is
increasingly important in making land use and other management decisions.
3.3.5 Fisheries modeling
Ecopath is a trophic structure model that simulates ecological processes in complex food webs, while Ecosim allows the
simulation of scenarios to maximize benefits under different management regimes and allows evaluation of tradeoffs for
different decisions. Combined socioeconomic and ecologic process models such as EwE (Ecopath with Ecosim) have been used
to evaluate fisheries management alternatives. Examples include managing a resource to optimize biodiversity or habitat for at-
risk species, or optimizing social values in terms of fisheries jobs (Cheung & Sumaila 2008). Ecopath was used by Pauly et al.
(2000) to document the structural changes in global fisheries. By looking at all historic data and estimating the trophic level of
each species, Pauly et al. (2000) estimated that top predators are being lost and that the average catch is lower in the food chain
today than in the past. This approach might be useful at a more local level to determine sustainable levels of harvest.
Another program, Marxan, is a model used to design marine protected areas (MPAs). The model incorporates an optimization
algorithm for finding spatially cohesive sites that meet specified criteria, such as biodiversity (Smith et al. 2002).
Jordan et al. (2008) demonstrated an approach that links production at the scale of habitat patches to large-scale delivery of the
ecosystem service (edible fish). This framework may be used to model habitat effects for use in predicting and managing coral
reefs and other coastal habitats, to identify sources of uncertainty and data gaps to improve the precision and accuracy of
predictions, and to demonstrate the potential for large-scale effects of multiple small-scale decisions on delivery of ecosystem
services.
3.4 Valuing the service
Coral reef associated fisheries encompass both direct and indirect values (van Beukering et al. 2010). Most of the research on
the value of ecosystem services has focused on direct use values (e.g., consumption for food, marine ornamentals) or indirect
use values (e.g., habitat provisioning, cultural and recreational importance).
Fisheries agencies often use the annual ex-vessel value (i.e., the gross value paid to commercial fishermen for their harvest).
For example, in 2000 the annual ex-vessel value of commercial fisheries associated with U.S. coral reefs was estimated at over
$137.1m (NOAA 2001). The ex-vessel value is an incomplete value of the fishery: it does not include the value added by
processors and vendors; it does not reflect the value of future catches; and, it does not reflect the uncertainly surrounding the
ability to obtain a comparable income in the future. For more complete fisheries valuation, economists estimate either the
present value (P V) or the net present value (NP V) of the fishery (Cheung & Sumaila 2008; Costanza et al. 1989). P V represents
a series of future cash flows expressed in today's dollars. NPV is a method used in evaluating investments; the NPV of all cash
outflows (such as the cost of the investment) and cash inflows (returns) is expressed in today's dollars. Both PV and NPV use a
discount rate (i.e., the rate at which society as a whole is willing to trade off present for future benefits) to calculate the value.
Data on the trade of marine ornamentals is, at best, qualitative . Collectors are generally small-scale fishermen working alone or
in small groups, using artisanal equipment (Wabnitz et al. 2003). CITES covers some marine ornamentals (including all species
of stony corals) and provides some trade data. National governments also produce statistics regarding the export or import of
marine ornamentals. A few countries report the actual number of specimens exported (e.g., Singapore and the Maldives). More
precise data can be obtained by interviewing collectors (Cesar et al. 2002), but this has not been done at a global scale.
For indirect use values (habitat provisioning) researchers may have derived a value for the resource by first identifying a
management scenario (MPA, No-Take Areas, incentive programs, etc.) from which they determined what the expected increase
in fish production and its value would be (opportunity cost methods). The value of fisheries has also included attempts to
calculate the Total Economic Value (TEV) of reefs, which includes fish production (Spurgeon 1992; Cesar 2002), but
difficulties arise when trying to sum nonuse and use values. Random utility models are most often used in valuation of
recreational fishing (Bockstael et al. 1989; McConnell et al. 1995). Many of these methods, except for determination of the
present monetized value of fish for food consumption, use contingent valuation methods to determine what people are willing
to pay, accept, or volunteer for the service (Table 3-3).
Cesar (2000), Moberg and Folke (1999), and Spurgeon (1992) provide excellent reviews on problems coral reefs face and on
what types of values and valuation methods should be used for valuing different benefits (Table 3-4). Stressors have been
translated into "disservices" for the purposes of valuation. For example, Cesar et al. (1997) used a quasi-option value approach
to evaluate the costs (potential losses due to the threat) and benefits of overfishing.
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Table 3-3. Valuation methods
1. Stated preference
Uses surveys to determine WTP (willingness to pay) or WTA (willingness to accept) or WTV (willingness to volunteer
one's services for fundraising, etc.)
a. Discrete or Dichotomous Choice Method
A good or service (or change in a good or service) is presented to samples of individuals. Randomly assigned dollar
amounts are presented, and individuals in the sample group choose their preferred amount. This is consistent with
how choices are made in markets and helps the analyst to derive estimates of economic value.
b. Choice Methods - Multi-Attribute Utility Theory
This method uses conjoint-type analysis to yield changes in total economic value or changes in preference rankings
for changes in attributes.
2. Revealed preference
a. Travel Cost Method
Uses the distance one has to go and the cost involved to assess value (e.g., bird watching)
b. The value of time spent traveling, or the opportunity cost of travel time
c. Opportunity cost
The value of the best alternative to a given choice, or the value of resources in their next best use. In regard to time,
the opportunity cost of time spent on one activity is the value of the best alternative activity that the person might
engage in at that time.
d. Random utility modeling
A version of the travel cost method that is often used for recreational fishing.
3. Cost/benefit analysis
4. Damage cost avoided, replacement costs, substitutes
5. Hedonic pricing
The hedonic pricing method is used to estimate economic values for ecosystem or environmental services that directly
affect market prices. It is most commonly applied to variations in housing prices that reflect the value of local
environmental attributes (e.g., housing values of Wisconsin lake front property increases with greater water clarity). It
can be used for estimating the economic benefit of environmental quality, including air pollution, water pollution, or
noise environmental amenities, such as aesthetic views or proximity to recreational sites.
6. Costless choice method
Dixon & Sherman (1990)
7. Total consumer surplus
Consumer surplus is the difference between the total amount that consumers are willing and able to pay for a good or
service (indicated by the demand curve) and the total amount that they actually do pay (i.e., the market price for the
product). An example might be the cost savings in property tax relief to individuals who place conservation easements
on their properties (these are deed restrictions that landowners voluntarily place on their land to protect important
resources). The total consumer surplus is simply the sum of all the consumer surpluses for each individual good
purchased.
8. Direct cost
For example, research expenditures by the Smithsonian Institution surveying coral reefs in Belize
9. Change in productivity approach
Difference in value of the biologically supported economic activity in situations with and without the reef (can use fish
production and yield estimates with and without MPA, or production on degraded versus undegraded reefs)
10. Percentage dependence technique
The value of the supported activity multiplied by an estimate of the percentage dependence of the activity on the reefs
presence
11. Replacement cost
For example, cost of installing artificial coastal defenses to replace reef protection function or fish production
(e.g., artificial reefs)
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12. Option price
Option value (benefit received by retaining the option of using a resource in the future by protection or preserving it
today) plus expected consumer surplus (the measure of the welfare that people gain from the consumption of goods and
services, or a measure of the benefits they derive from the exchange of goods).
13. Bioeconomic models
In this approach, both supply and demand are estimated. Production functions relating natural system attribute inputs
along with capital, labor and energy inputs are estimated and cost curves and the supply curve are estimated. Demand
curves for the good or service are also estimated. The outputs are changes in consumer's surplus, producer's surplus, and
a special part of producer's surplus—economic rent (ER) or the amount of profit over and above a normal return to
investment. ER is a measure of welfare for the fisheries where no one pays a price for the fish.
14. Total economic value (TEV)
TEV is an aggregation of consumer's surplus, producer's surplus/economic rent (CS, PS/ER). This includes use and
nonuse/passive economic use values (both direct and indirect). These values are used in Benefit-Cost Analysis (BCA),
damage assessments and restorations.
15. Value transfer
From meta-analysis of valuation studies (Brander et al. 2007)
16. Noneconomic human dimensions measures
(A more extensive discussion of these measures is provided in section 2.4.7 of this report.)
a. Importance-satisfaction ratings
b. The value of time spent traveling, or the opportunity cost of travel time
c. Opportunity cost
The value of the best alternative to a given choice, or the value of resources in their next best use. In regard to time,
the opportunity cost of time spent on one activity is the value of the best alternative activity that the person might
engage in at that time.
d. Random utility modeling
A version of the travel cost method that is often used for recreational fishing.
Source: except where noted, from Spurgeon (1992), Moberg & Folke (1999), and Cesar (2000).
Table 3-4. Categories of values
Direct use value
^ Extractive value (e.g., fisheries, coral for jewelry)
^ Nonextractive value (e.g., scuba diving)
Indirect use value
^ Habitat that supports fish
^ Nutrients
^ Shoreline protection
^ Global life support
Nonuse values
^ Existence value (i.e., value attributable to the presence of the reef, whether used or not)
^ Option value (i.e., potential future direct or indirect used, such as bioprospecting)
^ Bequest value (i.e., value of preserving for future generations)
^ Intrinsic value (i.e., innate value without reference to humans)
Source: Spurgeon (1992); Cesar (2000).
Income derived from fishing is another important benefit. Surveys conducted by Cinner et al. (2008) found that fishers from
poorer households would be less likely to exit a severely declining fishery. They suggest that wealth generation and
employment opportunities targeted at the poorest fishers would help reduce fishing effort in overfished areas (Cinner et al.
2008).
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3.5 Reflections
3.5.1 Threats to coral reef fisheries
Global landings offish have been in serious decline since the mid-1980s (Pauly et ol. 2005). Overfishing has been identified as
one of the major causes of reef ecosystem decline in recent decades, because the loss of harvested fish species changes the
structure and functioning of reef systems (Jackson et al. 2001; MEA 2005; Pauly et al. 2000; Burke & Maidens 2004) and
reduces reef resilience to natural disturbances (Hughes et al. 2003). While the loss or reduction of specific functional groups
(e.g., herbivores) through overfishing may reduce the size of harvestable fish stocks, overfishing of keystone herbivores like
parrot fish and surgeon fish completely alter reef dynamics. Their loss substantially increases growth of macroalgas and can
cause a phase shift in the reef system from coral to algal dominance (Hughes et al. 2003; Pandolfi et al. 2003; Burke &
Maidens 2004).
Many tropical fish species (e.g., groupers and snappers) form large spawning aggregations once or twice a year. The fish that
come to these aggregations are the oldest and largest individuals in the population (Coleman et al. 2000; Domeier & Colin
1997; Beets & Friedlander 1998; Smith 1972). These aggregations are easily targeted by fishermen, and intensive fishing
during spawning can quickly deplete a population (Burke & Maidens 2004; Beets & Friedlander 1998). Removing the largest
individuals from the population decreases spawning potential and reduces larval recruitment, since larger animals produce more
eggs (McField & Richards Kramer 2007; Roberts & Hawkins 2000). There is also evidence that spawning-site fidelity is a
learned behavior, and when heavy fishing at aggregation sites removes the experienced fish, new recruits are unable to locate
the aggregation site (Coleman et al. 2000; Warner 1990; Clark & Tracey 1993; Sadovy & Eklund 1999).
Destructive fishing methods pose another threat to coral reefs and to sustainable fish populations. Cyanide fishing (using
cyanide to stun reef fish for collection of live fish for the aquarium trade); blast fishing (using explosives to kill or stun reef
fish); muroami netting (nets that are weighted and dropped repeatedly onto coral); and gleaning (digging through reefs with
steel tools in search of abalone and invertebrates) have significant impacts on the reef structure, connectivity of the reef with
other habitats, and community structure of nontarget species (Pauly et al. 2000; Cesar 2002).
3.5.2 Management options
The American Fisheries Society (AFS) supports conservative management of reef fishes to avoid rapid overfishing and stock
collapse (Coleman et al. 2000). Two management approaches (essential fish habitat and marine protected areas) can be
combined to help maintain fish populations at sustainable levels.
3.5.2.7 Essential fish habitat (EFH).
The EFH provision of the 1996 Sustainable Fisheries Act (PL 104-297 1996) amended the habitat provisions of the Magnuson
Act (now called the Magnuson-Stevens Act) to require the National Marine Fisheries Service (NMFS), the Fishery
Management Councils, and Federal agencies to protect, conserve, and enhance essential fish habitat. Congress defined EFH as
"those waters and substrate necessary to fish for spawning, breeding, feeding or growth to maturity." Additionally, EFH that is
determined to be particularly important to the long-term productivity of populations of one or more managed species, or to be
particularly vulnerable to degradation, can be identified as "habitat areas of particular concern" (HAPC) to help provide
additional focus for conservation efforts (Duval et al. 2004).
EFH can account for spatial and temporal variation in the distribution of life history stage, seasonal and geographic
distributions, abundance, and interactions with other species. However, the regulations governing EFH designation do not
provide for temporal designation (i.e., a habitat is EFH all year long). Fisheries management plans now include a description
and identification of EFH, a description of potential threats (including how different fishing methods affect EFH), and actions
to conserve and enhance habitat (EPA 2005).
Spawning aggregations are potential EFH, since they are concentrated production sites and can be predictable in space and time
(Lindemann et al. 2000). EFH can also be identified by correlating benthic habitat variables with the distribution, abundance,
and size of reef fishes (Pattengill-Semmens & Semmens 2003). NOAA's Biogeography Program has produced benthic habitat
maps of most U. S. coral reefs. These data have been used to identify and map structural habitats used by fish species at
different life stages (Recksiek et al. 2001). The goal is to develop predictive habitat affinity models for selected fish species that
will support location of essential fish habitat.
3.5.2.2 Marine protected areas (MPAs).
MPAs or no-take areas (NTAs) may provide the best protection against overfishing (Hughes et al. 2003), so it is not
unexpected that a large body of literature is focused on optimization of MPA system design and the ecosystem service benefits
that they provide (Roberts & Polunin 1993; Trexler & Davis 2000; Cesar 2000; Pendleton 1995; Arias-Gonzalez et al. 2004;
Roncin et al. 2008; Stelzenmilller et al. 2008). Protected areas are an effective management tool, because "if well enforced they
change human behavior", and in actuality, it is human behavior and not the resource that is managed (Hughes et al. 2003).
Reserves in the Mexican Caribbean generally have a greater number of species, higher organism density, and larger-sized
herbivores than unprotected reefs (Nunez-Lara et al. 2003; Arias-Gonzalez et al. 2004).
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But estimates of the no-take area needed to be effective may be economically prohibitive in the short term despite the
increasing probability of the future collapse of the industry and the system as a whole. For example, Hughes et al. (2003) point
out that ecological modeling studies indicate that at least 30% of the world's coral reefs should be NTAs to ensure the
sustainability of exploited fish stocks. While this may seem extreme, Moberg and Ronnback (2003) point out that it is most
likely more cost effective to try to preserve ecosystem functioning than to restore or replace ecosystems when they have been
degraded or lost.
The quantitative associations between fish populations and their habitats is a key issue in determining EFH and establishing
MPAs and NTAs that protect fish and the fisheries. These must incorporate concepts of targeted fish species, life history stages,
age structure, genetic diversity of the stock, community structure, and physical habitats (Recksiek et al. 2001; Roberts et al.
1993; Coleman et al. 2000; Murray et al. 1999). While precise corridors of connectivity between habitats are not yet fully
understood, MPAs that encompass areas of connected habitats (e.g., coral reefs, seagrass meadows, mangroves forests) may
best protect fisheries (Pittman et al. 2010; Mumby et al. 2004; Lindemann et al. 2000).
MPAs may also address other management concerns, such as how to allocate ecosystem services among user groups. The
Florida Keys National Marine Sanctuary (FKNMS) is using no-take areas and other marine zoning methods to resolve conflicts
between users.
3.5.3 Conclusions
Fisheries are fairly well defined, but measurements and valuation are incomplete and vary widely. Attributes associated with
fishing have been identified, and scientists have begun to link those with fish production. Spatial and temporal analysis of fish
populations and their use of various habitats has advanced considerably in recent years.
Linking the attributes to ecosystem services (not just fish production) (Table 3-5) and conducting valuation studies will further
advance the science. Fish production is an ecological endpoint, a necessary part of the realization of benefits (amenities, goods,
or services) to people. The benefits can be thought of as ecosystem service endpoints for which there are values. Valuation
places a monetary or nonmonetary value on these benefits by establishing a relationship between the inputs (physical,
biological, socioeconomic factors) and the output (benefit). This relationship may be defined by a model, statistical equation,
function, or conceptual model.
Often the methods available to value a service, benefit, amenity, or stock, dictate how the benefits are described. The same
benefit may have very different inputs depending on the scale of analysis, the area under investigation, the focus of the research
(individual type of fish, what people in a particular area value most highly, the availability of certain tradeoffs, etc.). As we
suggest in Chapter 1, it may be helpful to distinguish between ecological services (which are expressed in physical units) and
economic benefits (which are expressed in monetary units). Some researchers have derived ways to combine the monetary and
nonmonetary benefit values of a system to compare purely economic values with the value of the system as a whole (Odum
1996).
Ecological models can be used to illustrate and quantify relationships among environmental and ecological reef elements and to
investigate thresholds for reef persistence and sustainable delivery of services. This information can be applied in the
development of EFH and MPAs, with the ultimate goal of a sustainable fishery.
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Table 3-5. Final ecosystem services and supporting features for fish production
Ecosystem Service(s)
Final (FES)
Intermediate
Natural Features
Social Values Complementary «-
Goods & Services Benefjts
Potential
Indicators of Final
Ecosystem
Service(s)
Fishing
Seafood Products
(fish, shellfish, algse
harvested)
Aquarium Products
(live fish & coral
taken)
Material Removed
for Curios and
Jewelry
Fish diversity and
_. , . , abundance; coral
Biological ...
. . .. health; seascape
integrity F
connectivity; and
structural complexity
Biological
integrity
Biological
integrity
Coral diversity,
abundance and health;
fish diversity and
abundance; seascape
connectivity; and
structural complexity
Coral diversity,
abundance and health;
water clarity
Desirability of
species based on
taste
Desirability of
species for aquaria
based on physical
appearance (color,
size, etc.), rarity
Aesthetic values
and artistic
inspiration
Adequate
infrastructure (boats,
marinas, etc.)
Revenue from
commercial
seafood fisheries
Revenue from
sales of
aquarium fish
and coral
_ . . . . ,. Revenue from
Diving and boating . ,,
. „ ° . sales ot curios
infrastructure , . .
and jewelry
Abundance of
commercially
desirable fish
species
Species abundance
and diversity of
target populations
Species abundance
and diversity of
target populations
Definitions (proposed by the Ecosystem Services Research Program and currently under discussion by the Program)
• Final Ecosystem Service - Output of ecological functions or processes that directly contributes to social welfare or has the potential to do so
in the future (broadly based on Boyd & Banzhaff [2007]).
• Intermediate Ecosystem Service - Output of ecological functions or processes that indirectly contributes to social welfare or has the potential
to do so in the future.
• Natural Features - The biological, chemical, and physical attributes of an ecosystem or environment.
• Social Values - The social attributes that influence economic demand for an ecosystem service.
• Complementary Goods & Services - Inputs (usually built infrastructure or location characteristics) that allow a good or service to be used by
complementing the ecological condition. For example, complementary goods and services that allow the presence of fishable fish to become
an opportunity for recreational fishing will include aspects of site accessibility, such as road access, available parking and the presence of a
fishing pier, all of which make fishing at the site possible and enhance enjoyment of the activity.
• Ecosystem-Derived Benefits - The contribution to social welfare of ecosystem goods and services. In the ESRP, the term applies specifically
to net improvements in social welfare that result from changes in the quantity or quality of ecosystem goods and services attributable to
policy or environmental decisions.
• Indicator of Final Ecosystem Service - Biophysical feature, quantity, or quality that requires little further translation to make clear its
relevance to human well-being (i.e., "public-friendly" measurement)
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Coral Reefs: Shoreline Protection
4.1 Defining the service
Coral reefs form natural barriers along the coast, protecting coastlines from erosion, flooding, and storm damage (UNEP-
WCMC 2006; WRI 2009). In general, the term "shoreline protection" refers to the ability of reefs to attenuate offshore wave
energy, providing sheltered nearshore waters, and protecting coastlines from erosion, flooding, and storm damage. Shoreline
protection has been defined in various ways, including intermediate biophysical ecosystem services endpoints that are indirectly
relevant to human well-being, final ecosystem services endpoints that are directly relevant to human well-being, or ecosystem-
derived benefits that provide social value (Table 4-1; Wainger & Boyd 2009). However, far greater progress has been made in
advancing our understanding of wave energy attenuation than in linking that attenuation to the provision of ecosystem services.
The key to quantifying shoreline protection is to understand the links between reef attributes, physical processes, and benefits
relevant to society.
Table 4-1. Measures that have been used to quantify shoreline protection
Ecological processes
Physical processes
• Reduction in wave energy, velocity, or height
Biological processes and structures
• Damage to coral reefs by hurricanes and storm events
• Presence of seagrasses or mangroves
• Fish density and species composition
Ecosystem services
• Rates of beach or shoreline erosion
• Shoreline geography
• Wave set-up during extreme events
• Coastal inundation during extreme events
Socioeconomic benefits
• Reduced properly damage or loss of life during extreme events
• Dollar value of avoided damages during extreme events
• Dollar value to build artificial wave breaks
Categories derived from Wainger & Boyd 2009
The physical properties of wave attenuation have been measured and modeled in studies of numerous locales worldwide,
including the Caribbean (Lugo-Fernandez et al. 1998), Australia (Hardy & Young 1996), and Hawaii (Gerritsen 1981). Ocean
waves traveling over coral reefs experience significant attenuation of energy, height, and velocity (Wolanski 1994; Gourlay &
Colleter 2005; Lowe et al. 2005). Wave attenuation by coral reefs has biological consequences for the reef itself as well as
inshore ecosystems:
• shelter is provided to nearshore coral from damaging effects of hurricanes
(Woodley et al. 1981);
• low-energy environments are created that are favorable to the growth of highly
valued wetlands, including seagrasses and mangroves (Birkeland 1985); and,
• recruitment and nursery habitats (mangroves and seagrasses) for fish are protected (DeMartini et al. 2009).
Shoreline protection has also been quantified using metrics that are more directly relevant to humans. Wave energy moderation
provided by reefs can greatly influence the geography of the coastline (Black & Andrews 2001) and reduce the rate of shoreline
erosion (Frihy et al. 2004). The value of such ecosystem services is often underappreciated until expensive beach restoration
(Riopelle 1995) or artificial breakwaters are needed to protect eroding beaches (Berg et al. 1998; Talbot & Wilkinson 2001).
Reefs also play an important role in mitigating coastal flooding caused by natural hazards, such as large storms, hurricanes or
cyclones, and tsunamis, which can cause immense properly damage and loss of human life (Sudmeier-Rieux et al. 2006).
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Quantifying shoreline protection in terms of hazard mitigation is challenging and often anecdotal. Although hurricanes or
cyclones occur worldwide, much of the information on coastal flooding, properly damage, and loss of life comes from
examining the effects of reefs in mitigating tsunami damage in Asian countries (UNEP-WCMC 2006). Recent extreme events
undoubtedly color the interpretations of shoreline protection, as do regional differences in geography, coastal development, and
frequency of storm events (WRI 2009).
4.2 Providing the service
4.2.1 Presence of the reef
The scientific evidence for shoreline protection by coral reefs is largely anecdotal and observational, quantified by various
physical, biological, and social endpoints with and without the presence of reefs (Table 4-1). The presence of coral reefs has
long been known to provide wave-sheltering and protection to the coastline, with locals utilizing calm waters for navigation
routes, fishing, and recreation (UNEP-WCMC 2006). The presence of reefs is associated with:
• the inshore presence of seagrasses and mangroves (Birkeland 1985; Ogden & Gladfelter 1983; Short et al. 2007);
• a reduction in offshore wave energy reaching the shoreline (Lugo-Fernandez et al. 1998);
• reduced rates of shoreline erosion (Hayden et al. 1978); and,
• reduced properly damage and loss of life during extreme events (EJF 2005; UNEP-WCMC 2006).
Furthermore, the presence of certain reef types or species (e.g., patch reef, Acropora spp., or Montastraea spp.) has also been
used to characterize the relative magnitude of wave energy attenuation (Mumby et al. 2008).
4.2.2 Reef attributes
Beyond the presence or absence of a reef, key attributes such as reef height, width, and topography (see Table 4-2) need to be
monitored to provide a more detailed understanding of wave energy attenuation.
The contribution of reef attributes to the hydrodynamic processes governing wave energy attenuation by coral reefs have been
extensively modeled and studied with field observations (Monismith 2007). The general idea is that incoming waves break at
the face of the reef, causing an initial increase in water level within the surf zone, which pushes waves over the reef flat as they
travel to shore (Figure 4-1). Offshore wave height and propagation over the reef will determine the height of waves reaching
the shore, typically quantified by the wave set-up (the increase in water level above the still water level) and wave run-up (the
height along the beach that water reaches due to incoming waves).
Table 4-2. Reef attributes that contribute to wave attenuation. Definitions are a synthesis of those used in literature
(see Appendix 4-A; also Figure 4-1 and Glossary)
Attribute
Definition
Presence of reef indicates whether or not an offshore reef is present near the coastal area of interest.
Reef continuity the extent to which the reef is uninterrupted or unfragmented in distribution; namely, the absence of large gaps
such as those due to degradation or coral mining.
Reef depth the distance from the ocean surface to the top of the reef; may be an assumed or fixed value in simulation models,
or an average value from field observations for the reef in question.
Reef distance the distance between the reef crest at the seaward edge of the reef and the edge of the shoreline; essentially, the
width of the lagoon; may be an assumed or fixed value in simulation models, or an average value from field
observations for the reef in question.
Reef height the distance from the top of the reef to its base; may be an assumed or fixed value in simulation models, or an
average value from field observations for the reef in question.
Reef roughness the bottom drag coefficient (which characterizes friction); may be approximated in field studies by variability in
colony height, or other measures of topography, along the reef flat; may be estimated indirectly by fitting models
to data on wave energy attenuation.
Reef slope the angle, from gradual to steep, of the reef front where offshore waves are first encountered; may be an assumed
or fixed value in simulation models, or an average value from field observations for the reef in question.
Reef type describes the general structure of the reef and its relationship to the shoreline, including fringing reefs that border
the shoreline, barrier reefs that are separated from shore by a deep lagoon, atoll reefs that form a circular barrier
around an island, and patch reefs that are small, isolated reef outcrops.
Reef width the length of the reef flat, the flat expanse of reef from where offshore waves first crest over the reef to the edge
closest to the shoreline; may be an assumed or fixed value in simulation models, or an average value from field
observations for the reef in question.
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Figure 4-1. Illustration of wave set-up and attenuation over a reef (redrawn from Monismith [2007]).
Coral reefs reduce offshore wave energy primarily in two ways: (1) the steep gradient from deep water to shallow causes the
wave to break at the reef crest; and, perhaps more importantly, (2) the increased bottom friction along the reef flat creates drag
(Wolanski 1994; Lowe et al. 2005). Reef depth is perhaps the simplest single attribute that can be related to wave attenuation
(Gourlay 1994; Barbier et al. 2008). Reef area, reef slope, and distance from shoreline are additional key attributes derived
from field (Young 1989; Hardy & Young 1996; Brander et al. 2004) and modeling studies (Tait 1972; Gourlay & Colleter
2005). In general, wave energy attenuation decreases with increasing reef depth across the reef flat, with greater energy being
attenuated as reef width increases (Figure 4-2; Kunkel et al. 2006), such that broad, shallow reefs provide the greatest
attenuation. Tidal variations in water depth and initial offshore wave height can also influence the degree of wave attenuation
(Madin et al. 2006).
Modeling studies indicate that friction across the reef surface is a strong determinant of the degree of wave energy attenuation,
as can be seen in Figure 4-2 (Sheppard et al. 2005; Kunkel et al. 2006). Drag coefficients used to quantify reef friction in
simulation models are often calibrated by comparing model outcomes to field data. These coefficients indicate that coral reefs
exert ten times more drag than sandy bottoms (Tait 1972; Roberts et al. 1975; Lugo-Fernandez 1998; Reidenbach et al. 2006).
Measurements of in situ reef friction are challenging to obtain. Field estimates of reef roughness based on the standard
deviation of reef height have provided values similar to model-calibrated estimates of reef friction(Lowe et al. 2005). Other
methods for estimating reef roughness have been used, including draping chains over reefs along a transect to generate a linear
measure of surface topography. There are as yet no standard methods for deriving wave energy attenuation from field
measurements of reef roughness (Monismith 2007).
a)
1 -,
u> 2 0.5
SI
<1) -s
0)
Reef Width
Narrow
Moderate
Wide
b)
Reef Depth
c «
§ 0)
IE Of-
i- O
;s 0.5 -
<
Figure 4-2. Relationship of wave energy with increasing
reef depth and width (top) and with
increasing reef friction (bottom) (derived from
Sheppard et al. [2005] and Kunkel et al. [2006])
Reef Friction
-------
4.2.3 Reef health
The health of a coral reef is an important determinant of wave attenuation. Degraded reefs have diminished roughness, and
model simulations indicate the reduction in reef friction reduces the reef's attenuation of wave energy (Figure 4-2; Kunkel et al.
2006; Sheppard et al. 2005). A reduction in reef friction of approximately 50% could produce a doubling of wave energy
reaching the shores behind those reefs. Dead corals on degraded reefs break or disintegrate easily under wave action, resulting
in reduced reef height and roughness and, consequently, in reduced attenuation of wave energy. In general, the degree of
shoreline protection afforded by reefs during storms reflects a balance between the reef's attenuation of wave energy and the
damage inflicted on the reef itself by that storm event (Lacambra et al. 2008).
Reef continuity is another important indicator of shoreline protection (WRI 2009). Damage to reefs from coral mining , in
which large sections of coral are harvested to provide blocks for construction, can accelerate rates of beach erosion and require
expensive beach restoration (Riopelle 1995). Fragmented reefs can allow high energy tsunami waves to reach the shoreline
(Nott 1997) and may intensify flooding (Fernando et al. 2005; Chatenoux & Peduzzi 2005). Furthermore, sections of degraded
reef that have been invaded by macroalgas may afford less protection than reefs with a high abundance of large stony coral
species (Mumby et al. 2008).
Shoreline protection may be devalued by as much as 80%-90% when reefs are degraded (Burke & Maidens 2004). For
valuation purposes, reefs are assumed to retain their protective capacity until coral cover (living tissue) loss exceeds 25%, after
which the value of coastal protection declines linearly with increasing loss of coral (Cesar 1996; Pet-Soede et al. 1999). The
most extreme degradation leads to loss of reefs, which can have severe consequences for property damage and human life in the
case of extreme hazard events (Sudmeier-Rieux et al. 2006).
4.3 Measuring the service
Shoreline protection has been quantified in numerous ways that vary in their relevance to human well-being (Table 4-3).
Natural features, including reef attributes and physical variables, have been translated into metrics of shoreline protection using
anecdotal information, statistical relationships, and mechanistic models. Estimates of coastal protection also depend on
socioeconomic variables.
4.3.1 Physical processes
Incoming waves encountering the steep gradient at the reef front causes the waves to break, producing a sudden increase in
wave amplitude that diminishes (or attenuates) as the residual wave energy moves toward shore (Figure 4-1). Numerous field,
laboratory, and modeling studies have looked at physical endpoints of shoreline protection in terms of reductions in wave
energy, wave height, or wave velocity from offshore to the shoreline (reviewed in Gourlay & Colleter [2005], Sheppard et al.
[2005], and Monismith [2007]; see Table 4-3). Field studies have measured as much as a 68%-95% reduction in wave energy
as waves travel over the reef flat (Roberts et al. 1975; Gerritsen 1981; Young 1989; Lugo-Fernandez et al. 1998; Brander et al.
2004). Tidal variations in reef depth will produce variations in the degree of wave energy attenuation. At high tide, waves
within the normal range may be reduced only slightly by the reef, whereas at low tide all waves may effectively be blocked
(Madin et al. 2006). Attenuation of wave energy and height are essential for understanding the height of wave inundation on the
shore, referred to as wave run-up, which also has been modeled (Kunkel et al. 2006). Wave run-up may be a potential indicator
of damage from flooding, and physical models can be used to translate reductions in wave height into flood hazard maps,
depending on the slope and porosity of the shoreline (FEMA 2007).
4.3.2 Biological processes
Shoreline protection by coral reefs has also been quantified through the health and survival of highly valued biological
components in near-shore areas and along the coast. For example, the presence of outer reefs can shelter nearshore coral from
damaging effects of hurricanes (Woodley et al. 1981). The protection afforded by coral reefs also creates low-energy
environments favorable to the growth of seagrasses or mangroves in coastal wetlands (Ogden & Gladfelter 1983; Birkeland
1985; Short et al. 2007; Sheaves 2009). Reefs reduce the vulnerability of wetlands to damage and vegetative loss during
hurricane events (Fourqurean & Rutten 2004). Wetlands themselves provide numerous ecosystem services (Mumby et al. 2008)
and in many regions are more important for coastal protection than the presence of reefs (EJF 2005; Cochard et al. 2008). Reefs
appear to serve as a first line of defense by diffusing wave energy and protecting coastal wetlands, thereby enhancing the
protective value of those wetlands. Whether there exists a synergistic relationship between wave energy attenuation by reefs
and shoreline protection by mangrove and seagrass wetlands has not been studied.
-------
Table 4-3. Measures used to quantify shoreline protection, and the major reef attributes, physical parameters,
and socioeconomic parameters used to estimate that protection's value (see Appendix 4-A for details)
Measures
Physical processes
Reduction in wave energy, velocity,
or height in appropriate units
Biological processes
Damage to coral by hurricanes
(e.g., prevalence of broken coral)
Presence of seagrasses or
mangroves (e.g., area)
Fish density & species
composition
Ecosystem services
Rates of shoreline erosion (e.g.,
distance or volume lost per year)
Shoreline geography (e.g., presence
of salients & tombolos, change in
shoreline position)
Wave set-up during extreme events
Coastal inundation during extreme
events (e.g., decrease in area
inundated)
Reef Attribute
Reef depth, width, slope,
roughness, distance to shore
Presence of outer reef;
Reef depth, width, slope
Presence of reef;
Gap in reef
Presence of reef
Presence of reef;
Reef depth
Reef width;
Distance from shoreline
Presence of reef; Reef depth,
width, slope, & roughness
Presence or area of reef;
Gap in reef;
Physical Variables Socioeconomic
Variables
Offshore wave energy & height;
Tidal depth
Hurricane path
Water motion;
Wind or wave exposure
Wave energy; Beach elevation &
sediment grain size
Wave height and period
Offshore wave energy, height,
& amplitude
Distance from hazard event;
Slope of coastline;
Socioeconomic benefits
Decrease in property damage or loss
of life during extreme events
Value of avoided damages
Replaces need for costly artificial
breakwaters or beach replenishment
Presence of intact reefs
Presence of reef; Reef type,
continuity, & distance from shore
Presence of reef
Coastal geography; ,, ,
„, ,° ,° „ ,- Property values
Storm height & trequency
Breakwater costs;
Restoration costs
Reductions in wave energy by reefs may also provide prime wave-sheltered habitat for larval fish and enhance local
recruitment. Lower water velocities and reduced wave exposure are associated with high juvenile fish densities (Burgess et al.
2007; DeMartini et al. 2009). Fish assemblages in wave-sheltered reefs often have different species than exposed habitats,
including small fish species whose locomotion and foraging activities may be inhibited in fast-moving water (Fulton &
Bellwood 2005). Increased fish abundance would likely increase fishing and tourism services, something that has been
discussed anecdotally (UNEP-WCMC 2006) but not linked directly to the physical processes of wave energy attenuation.
4.3.3 Ecosystem service measures
Physical hydrodynamic processes associated with coral reefs affect the geography, appearance, and stability of the shoreline.
The strength and pattern of waves reaching the beach determine rates of beach erosion and shapes the contours of the shoreline
(Hayden et al. 1978). Models have been developed to predict rates and patterns of shoreline changes under a variety of wave
conditions (Frihy et al. 2004). Assuming the edge of the shoreline may change by an average of 0.4 meters per year in
unprotected areas, an example of a quantitative measure of an ecosystem service is given in Berg et al. (1998), who estimated
that 1 km2 of reef, protecting 5 km of shoreline along the coast of Sri Lanka could prevent a loss of 2,000 m2 of land per year.
The presence of reefs can also influence the contour of the shoreline through the creation of salients (bell-shaped extensions of
the shoreline toward the reef) and tombolos (shoreline extensions connecting to offshore sandbars) (Figure 4-3). The size of
-------
these shoreline extensions depends on the width of nearby reefs and reef distance offshore (Black & Andrews 2001). Coral
reefs also play a role in replenishing sandy beaches and islands as corals and other calcified organisms break down after death
(UNEP-WCMC 2006).
Figure 4-3. Reef influences on shorelines: salient (left) and tombolo (right).
(photo credits: salient - California State Parks [2011]; tombolo - Tim Bekaert [2005])
Reductions in wave energy or wave height by coral reefs have clearer socioeconomic relevance when connected to wave run-up
or coastal flooding during storm or extreme hazard events. Numerous studies that have examined wave run-up during tsunami
events suggest that, while there is some evidence of reduced flooding in areas behind reefs (Fernando et al. 2005), channelized
or fragmented reefs may actually accelerate movement of tsunami waves to the coastline (Nott 1997; Chatenoux & Peduzzi
2007). Modeling studies indicate the buffering ability of reefs is largely dependent on the state of reef health (Kunkel et al.
2006). In other observational studies, however, reefs appeared to have little protective value, and tsunami inundation was
largely determined by wave height and coastal topography (Baird et al. 2005). Coral reefs may be more effective at buffering
normal wave action or storm events than tsunamis, which have longer wavelengths and larger wave amplitudes (Cochard et al.
2008).
4.3.4 Socioeconomic benefits
The value of reefs in shoreline protection is often underappreciated until quantified in terms of human lives, properly damage,
or economic costs. Anecdotally, the presence of reefs is linked to diminished properly damage and loss of life during hurricanes
(Whittmgham et al. 2003) or tsunamis (Liu et al. 2005; WI 2005; UNEP 2005; EJF 2005; UNEP-WCMC 2006). In one study
of a tsunami's effects, areas protected by reefs experienced wave heights of only 2-3 m with inundation extending only 50 m
inland with no loss human life (Fernando et al. 2005). In contrast, just 3 km to the north where there was no coral reef to
protect the shore, the same tsunami resulted in a 10 m wave that flooded 1.5 km inland and killed 1,700 people. This evidence
is largely anecdotal, however, and there is some question as to whether reefs substantially reduce coastal inundation during
tsunami events (Baird et al. 2005; Cochard et al. 2008).
Although shoreline protection can be a significant contributor to the total economic value of coral reefs, it is largely
underestimated by decision-makers, except during times of crisis. A few studies (see below) have attempted to place a dollar
value on coastal protection by reefs, either in terms of expected damages or beach/shoreline replacement costs due to reef
degradation (Chong 2005).
4.4 Valuing the service
There are many coral reef studies estimating costs of reef restoration, fisheries value, or recreational value, but only a few
studies have looked at the economic value of shoreline protection (Chong 2005; Table 4-4). One approach to valuing coastal
protection is to estimate defensive expenditures required to replace the loss of the reef. When reefs are severely degraded, they
may need to be replaced with artificial breakwaters that may cost $10m per linear kilometer to construct (Wells & Edwards
1989; Weber 1993; Berg et al. 1998; Talbot & Wilkinson 2001). In other cases, expensive beach and shoreline restoration is
needed as a consequence of lost coral reefs (Riopelle 1995).
Another approach to estimating the economic value of coastal protection is to estimate the damages avoided due to the presence
of the reef. In a study of Indonesian Reefs, Cesar (1996) estimated the monetary value of damage avoided based on the value of
three types of coastal development: (1) the value of agricultural production ($820 per km coastline); (2) the cost of replacing
homes and road infrastructure ($50,000 per km coastline); and, (3) hotel expenditures toward maintaining beaches ($1,000,000
per km coastline). These values, or similarly derived values, have been used to value damages avoided due to the presence of
reefs in Bermuda (Beukering et al. 2010), the Philippines (White et al. 2000), throughout the Caribbean (Burke & Maidens
2004), southeast Asia (Burke et al. 2002), and worldwide (Cesar et al. 2003).
The economic value of coral reefs depends on more than just the presence of the reef. Reliable economic estimates require
knowledge of biological, physical, and socioeconomic factors that influence the provision of and need for coastal protection
-------
(WRI 2009). First, the value of shoreline protection will depend on whether the coastal area is vulnerable to erosion or storm
damage, namely low-lying lands near the coast. Second, coastal vulnerability is determined by the physical stability of the
shoreline (including coastal geomorphology and geology, elevation, and vegetation) and the potential for storm surges
(including offshore wave energy and hurricane frequency). Third, reef attributes such as distance from shore, reef continuity,
and type of reef determine the protective capability of the reef. Fourth, the amount and value of coastal development will
determine the potential value of protection. All of these factors must be considered when estimating the potential economic
damages resulting from the loss of reefs. This approach has been used to estimate the value reef coastal protection throughout
the world as ranging from $0.3 billion to $2.2 billion (Burke et al. 2002; Burke & Maidens 2004). Coastal development,
sedimentation, pollution, overfishing, and climate change can severely degrade reefs, reducing their protective ability by an
estimated 80-90%, with a potential net loss of benefits associated with shoreline protection on the order of $ 140m to $420m
per year in the Caribbean (Burke & Maidens 2004).
Table 4-4. Estimated values of shoreline protection (modified from Chong 2005)
Location
Americas
Bermuda
Caribbean
Caribbean
USA
Asia
Australia
Indian Ocean
Indonesia
Japan
Philippines
SEAsia
Pacific
Sri Lanka
World
Values
Caribbean
Low development
Medium development
High development
Indonesia
Remote areas
Some construction
Major infrastructure
Philippines
Maldives
Total Value
$266m
$720m
$700m-$2,200m
$172m
$629m
$l,595m
$314m
$268m
$326m
$5,047m
$579m
$30m
$9,009m
Study
Beukering et al. 2010
Cesar et al. 2003
Burke & Maidens 2004
Cesar et al. 2003
Cesar et al. 2003
Cesar et al. 2003
Burke et al. 2002
Cesar et al. 2003
Burke et al. 2002
Cesar et al. 2003
Cesar et al. 2003
Bergetal. 1998
Cesar et al. 2003
per kilometer of shoreline
$2,000-$20,000
$30,000-$60,000
$100,000-$ 1,000 ,000
$820
$50,000
$1,000,000
$5,000-$25,000
$10,000,000
Burke & Maidens 2004
Burke & Maidens 2004
Burke & Maidens 2004
Cesar 1996
Cesar 1996
Cesar 1996
White et al. 2000
Talbot & Wilkinson 2001
4.5 Reflections
Final ecosystem services for shoreline protection are summarized in Table 4-5. Natural features, including reef attributes and
physical variables, contribute to biophysical processes that provide ecosystem services, such as wave energy attenuation. These
natural processes directly benefit humans by reducing shoreline erosion and protecting coastlines from inundation during
extreme events. Social values, such as the desirability of coastal housing or the attractiveness of sandy beaches, influence the
demand for shoreline protection. Complementary goods and services, such as the availability, intensity, and location of coastal
development or the absence of constructed breakwaters, influence the opportunity to take advantage of shoreline protection.
The existence of constructed breakwaters, including rubble mounds and artificial reefs, can also diminish the demand for
shoreline protection by natural reefs.
4.5.1 Improving current knowledge
Physical models of wave energy attenuation are based on relatively simple physical assumptions, so field validation of
mathematical models of wave energy attenuation would improve our ability to characterize the degree of protection provided
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under different conditions (Monismith 2007). Furthermore, the reef is generally treated as a single object, rather than a living
community. Although some models have attempted to connect reef roughness to wave energy attenuation (Sheppard et al.
2005; Kunkel et al. 2006), there is little understanding of how reef attributes, such as species composition or skeletal
calcification rates, contribute to wave attenuation, or how growth, reproduction, and survival contribute to sustainable shoreline
protection.
Our understanding of the social benefits of shoreline protection is largely derived from anecdotal evidence, but it could be
improved with more quantitative or statistical studies relating properly damage or loss of life to reef attributes and physical
properties of the coastline. Furthermore, studies of flooding or potential damage during extreme events are largely tied to
tsunamis (UNEP-WCMC 2006). Less is known about the extent to which reefs reduce hurricane damage, but the protection
provided by reefs may result from buffering wave energy rather than reducing inundation (Cochard et al. 2008).
The presence of reefs is associated with providing wave-sheltered environments for mangroves and seagrasses (Birkeland
1985), yet there is little understanding of how subtle changes in reef integrity may impact these neighboring systems.
Furthermore, wetlands themselves are important for coastal protection and their protective ability may be enhanced by the
presence of reefs as a first line of defense (EJF 2005). However, the potential synergistic relationship between reef shoreline
protection and the provision of wetland ecosystem services is not well understood.
Few studies have attempted to quantify or value indirect consequences of shoreline protection by reefs (Chong 2005). Direct
consequences of storm events, hurricanes, and tsunamis include loss of lives, housing, and buildings. These direct losses can
have long-lasting reverberations as the affected society becomes vulnerable to disease epidemics or economic instability
(Cochard et al. 2008). The potential value of shoreline protection will depend on the likelihood of such indirect consequences,
and the potential for the local economy to rebound from disasters. Historical data on disease outbreaks, economic losses, and
time to economic recovery for post-hurricane or post-tsunami economies might suggest which societies are particularly
vulnerable to a loss of shoreline protection. The current lack of consideration of indirect consequences is an oversight that may
lead decision makers to underestimate the potential value of shoreline protection.
4.5.2 Connecting biophysical processes and ecosystem services to socioeconomic benefits
For biophysical endpoints to be relevant to humans, they must connect to an ecosystem service or social benefit (Wainger &
Boyd 2009). Ideally, our ability to quantify shoreline protection in socially relevant endpoints requires that we characterize two
relationships in quantitative terms: (1) that between reef attributes and the physical environment; and, (2) that between the
physical environment and ecosystem services, including their social benefits (Figure 4-4). Natural features of the environment,
including attributes of the reef and physical attributes of offshore waves, affect physical ecosystem processes, such as
attenuation of wave height or energy. Attenuation of wave energy by reefs leads to protection from shoreline erosion or
flooding, but the degree of protection will depend on physical variables, such as coastal geography, vegetation, and the
frequency of storm events. Humans derive benefits from shoreline protection through reductions in properly damage or loss of
life, but the value of protection will depend on socioeconomic factors, such as the degree of coastal development and properly
values (WRI 2009). Wave energy attenuation can also indirectly benefit humans by providing wave-sheltered habitats for fish
or protecting wetlands, which themselves provide ecosystem services.
The quantitative relationships between reef attributes and shoreline protection and between physical processes and
socioeconomic benefits are poorly understood. Models of wave energy attenuation are reasonably adept at accounting for the
impacts of reef attributes such as height, width, slope, and roughness (Lowe et al. 2005; Sheppard et al. 2005; Kunkel et al.
2006). However, our understanding of the socioeconomic benefits, such as the prevention of properly damage or loss of life, is
largely anecdotal, based solely on the presence or absence of a reef (UNEP-WCMC 2006). Further research and modeling
efforts are needed to provide useful quantifications of these relationships.
-------
Table 4-5. Final ecosystem services and supporting features for shoreline protection
Ecosystem Service(s)
Natural Features Social Values Complementary J
Final (FES) Intermediate 1 Goods & Services I Benefjts
Potential
Indicators of Final
Ecosystem
Service(s)
Shoreline Protection
Protection from Decreased erosion in Reef height, Reef continuity Beach elevation, Desirability of
shoreline erosion kg/ha/y width, slope, & Offshore wave sediment grain size housing near
Protection from
coastal inundation
during extreme
events area in
hectares protected
Reduction in wave
set-up, or storm
surge
Reduction in wave roughness energy & wave Attractiveness of water
energy, velocity, or
height
Presence of reef
Reef height, width,
slope, roughness
Reef continuity
Wave energy
height sandy beaches Absence of
Tidal depth
Distance from
hazard event
constructed
Slope of coastline
breakwaters
Higher property
values
Frequency &
intensity of
extreme events
Past history of
extreme events
Absence of
constructed
breakwaters
Presence of reef
Opportunity to use
beaches (see
Chapter 2)
% reduction in
rates of shoreline
erosion due to
presence of reef
Location,
intensity, and
value of coastal
development
Lower insurance
rates
Definitions (proposed by the Ecosystem Services Research Program and currently under discussion by the Program)
• Final Ecosystem Service - Output of ecological functions or processes that directly contributes to social welfare or has the potential to do so
in the future (broadly based on Boyd & Banzhaff [2007]).
• Intermediate Ecosystem Service - Output of ecological functions or processes that indirectly contributes to social welfare or has the potential
to do so in the future.
• Natural Features - The biological, chemical, and physical attributes of an ecosystem or environment.
• Social Values - The social attributes that influence economic demand for an ecosystem service.
• Complementary Goods & Services - Inputs (usually built infrastructure or location characteristics) that allow a good or service to be used by
complementing the ecological condition. For example, complementary goods and services that allow the presence of fishable fish to become
an opportunity for recreational fishing will include aspects of site accessibility, such as road access, available parking and the presence of a
fishing pier, all of which make fishing at the site possible and enhance enjoyment of the activity.
• Ecosystem-Derived Benefits - The contribution to social welfare of ecosystem goods and services. In the ESRP, the term applies specifically
to net improvements in social welfare that result from changes in the quantity or quality of ecosystem goods and services attributable to
policy or environmental decisions.
• Indicator of Final Ecosystem Service - Biophysical feature, quantity, or quality that requires little further translation to make clear its
relevance to human well-being (i.e., "public-friendly" measurement)
One possible approach for linking reef attributes with socioeconomic benefits of shoreline protection would be to integrate
socioeconomic metrics into physical process models. Wave attenuation by reefs has been fairly well characterized in laboratory
experiments, field observations, and models (as reviewed in Gourlay & Colleter [2005], Sheppard et al. [2005], and Monismith
[2007]), and it has been connected to shoreline changes using predictive models (Frihy et al. 2004). However, for all intents and
purposes, there are no studies connecting changes in wave energy or height to potential loss of life or properly damage. The
Reefs at Risk shoreline protection index (WRI 2009), which is used to calculate the damage prevention provided by reefs,
indirectly accounts for a number of physical processes by considering key reef attributes related to wave attenuation. In
addition to reef distance from shore, Reefs at Risk includes reef type as a proxy for reef depth and slope, and reef degradation
as a proxy for bottom friction. However, these factors are combined into a single index reflecting shoreline stability and fail to
capture more subtle changes in reef topography or species composition that may influence wave energy attenuation.
Bayesian probabilistic networks could be useful for characterizing the complex relationships between reef attributes and
probabilistic characterizations of exposure, damage, and cost (Cochard et al. 2008). Physical models of wave energy
attenuation that permit subtle changes in reef height, slope, width, and friction, combined with the physical properties of the
coastline and local climate, could be the basis for developing predictive models for the probability of flooding or shoreline
erosion. These could then be tied to socioeconomic factors, such as the degree of coastal development, to predict the probability
of properly damage, loss of life, or a variety of indirect consequences. Such models could reflect the uncertainly associated with
largely anecdotal information and could be updated as probabilistic characterizations of relationships or physical process
models are improved.
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4.6 Bibliography
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Pratchett MS, Rudi E and Siregar AM. 2005. Acehnese reefs in the wake of the Asian Tsunami. Current Biology, 15:1926-1930.
Barbier EB, Koch EW, Silliman BR, Hacker SD, Wolanski E, Primavera J, Granek EF, Polasky S, Aswani S, Cramer LA, Stoms DM,
Kennedy CJ, Bael D, Kappel CV, Perillo GME and Reed DJ. 2008. Coastal ecosystem-based management with nonlinear
ecological functions and values. Science, 319:321-323.
Bekaert T. 2005. Accessed 2011.08.09.
Berg H, Ohman M, Troeng S and Linden O. 1998. Environmental Economics of Coral Reef Destruction in Sri Lanka, Ambio,
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van Beukering PJH, Sarkis S, McKenzie E, Hess S, Brander L, Roelfsema M, Looijenstijn-van der Putten L & Bervoets T. 2010.
Total Economic Value of Bermuda's Coral Reefs. Valuation of Ecosystem Services, van Beukering Consulting, Amsterdam.
Birkeland C and Grosenbaugh D. 1985. Ecological interactions between tropical coastal ecosystems. Nairobi: United Nations
Environment Programme (UNEP), Regional Seas Reports and Studies, vol. 73, 71 pp.
Black K and Andrews C. 2001. Sandy shoreline response to offshore obstacles: Part 1. Salient and tombolo geometry and shape.
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Appendix 4-A
Studies quantifying shoreline protection
The following table lists studies and the endpoints they used to quantify shoreline protection. Also included are the biological
reef attributes and the physical and socioeconomic variables used to estimate that endpoint. The method and location of the
studies are also given.
Ecosystem
Service Endpoint
Reef
Attributes
Physical
Variables
Socioeconomic .. .. .
., . . . Method
Variables
Location
Citation
Physical processes
wave height
% change in wave
height
change in wave height
wave energy reaching
shore
shore wave energy;
wave "set-up";
wave velocity
wave height and
period; wave spectra;
wave breaking
wave attenuation,
spectra
change in wave
height; rate of wave
dissipation
wave set-up;
wave spectra
loss of wave height,
velocity, and
acceleration
wave energy
dissipation
wave energy
dissipation spectra
over range of
frequencies
% offshore energy
reaching the shore
reef top water depth
distance to beach;
variation in reef
topography; reef depth
presence of reef
reef depth & slope
reef top water depth (reef
height); reef friction; reef
slope; width of reef flat
(distance from shore)
distance along reef flat
from reef front; depth
over reef flat
presence of reef flat
reef roughness (standard
deviation in reef height);
presence of reef flat;
presence of fore reef
presence of reef crest,
reef flat, lagoon
distance from reef crest;
model-calibrated reef
friction parameter
presence of mangroves,
sea grass, patch reef,
Acropora, algal,
gorgonians, or
Montastraea
presence of reef crest,
reef flat
reef top water depth (reef
height); reef friction; reef
slope; width of reef flat
(distance from shore);
relative area of live,
dead, eroded coral, sand,
rubble, seagrass, or algal
turf
low or high tide
wave height, water
level
wave height, water
depth
offshore wave height
& period
wave height
tidal level
fore reef wave height,
wave frequency,
seawater density, etc.
tide level
wave height, period;
tide level
offshore wave height
& period
Statistical
analysis
Field study
Field study
Laboratory
study & model
Model
Field study
Field and
laboratory
studies
Model & field
study
Field Study
validation of
Tail's model
Field
validation of
model
Field study &
literature
review
Field study
Model
Australia
Australia
Hawaii
Australia
Australia
Australia
Japan
Hawaii
Caribbean
Australia
Caribbean
Caribbean
Indian Ocean
Barbier et al.
2008
Brander et al.
2004
Gerritsen 1981
Gourlay 1994
Gourlay &
Colleter 2005
Hardy &
Young 1996
Kono&
Tsukayama
1980
Lowe et al.
2005
Lugo-
Fernandez et
al. 1998
Madin et al.
2006
Mumby et al.
2008
Roberts et al.
1975
Sheppard et al.
2005
-------
Ecosystem
Service Endpoint
Reef
Attributes
Physical Socioeconomic
Variables Variables
Method
Location
Citation
Physical processes (con't)
reduction in wave
height
wave dynamics;
hydrodynamic
processes
wave attenuation;
wave spectra; area of
influence
presence of reef front,
reef flat; distance from
reef front;
depth at reef crest; fore
reef slope
reef slope, height, depth
presence of reef flat, fore
reef; reef flat water
depth; reef area
offshore wave height
& period; beach slope
offshore wave height
& period
incident wave height
Model
Model
Field study
Hawaii
Australia
Australia
Tail 1972
Wolanski 1994
Young 1989
Biological processes processes
damage to coral by
hurricane
low energy environ-
ment favorable for
growth of mangroves
and seagrasses
loss of seagrass
vegetation
low energy
environment
favorable for growth
of mangroves and
seagrasses
presence of
mangroves
presence of seagrasses
presettlement fishes
larval fish densities
fish assemblages
reef profile (depth, slope,
& shelf width); presence
of reef crest
presence of reef
presence of reef; gap in
reef
presence of reef
presence of reef flats
presence of reef
presence of reef
presence of reef
sheltered reef vs. wave-
exposed fore-reef
wave sheltered
tidally generated
eddies
wind/wave exposure
water motion (flow
velocity & rates of
direction change)
Model
Anecdotal
(review)
Field
observations
Workshop
summary
Review,
anecdotal
Mapping,
anecdotal
Field study
Field study
Field study
Jamaica
Caribbean &
Pacific
Florida
Caribbean
Global
Global
Australia
Hawaii
Australia
Woodley et al.
1981
Birkeland 1985
Fourqurean &
Rutten 2004
Ogden &
Gladfelter
1983
Sheaves 2009
Short et al.
2007
Burgess et al.
2007
DeMartini et
al. 2009
Fulton &
Bellwood 2005
Ecosystem services
shift in shoreline due
to erosion
beach sand erosion
formation of salients
and tombolos
hydrographic profile of
seabed composition [reef
face vs. sand],
reef depth
presence of reef
ratio of length of reef
along shoreline to
distance of reef from
undisturbed shoreline;
completely submerged
waves (height,
direction, period),
current, sediment
(grain size),
water depth
wave energy (height
& period of waves at
beach); beach
elevation; height of
sand deposition; beach
elevation; sediment
grain size
wave climate
Model
Field study
Statistical
analysis
Egypt
U.S. Virgin
Islands
Australia
Frihy et al.
2004
Hay den et al.
1978
Black &
Andrews 2001
-------
Ecosystem
Service Endpoint
Ecosystem services (con '
tsunami run-up
relative to no reef
reduction in wave
height during cyclone
coastal flooding
tsunami height &
inundation; tsunami
damage (structures
destroyed, deaths)
coastal flooding
tsunami run-up,
height
tsunami movement
to coast
tsunami wave run-up
Coastal inundation
during extreme events
probability of indirect
or direct
consequences due to
damage from
exposure
(p[CID|D,EX])
Reef
Attributes
t)
depth and width of reef
flat; width of lagoon
(offshore distance to the
reef); bottom drag
coefficient ("reef
health"); presence,
width, and location of a
gap
presence of reef
% coral protection; %
seagrass; % mangrove;
reef orientation
presence of reef or gap in
reef
reef profile (depth, slope,
& shelf width); reef
continuity; reef area; reef
topography; terrace
width; species
distribution, geometry, &
ecology; species at reef
front more resistant
gap in reef
gaps in reef
presence of reef
presence of reef
presence of reef;
fragmentation of reef
(size & orientation of
channels); seagrass or
mangrove vegetation
structure; orientation to
coastline; vegetation
relation to landscape
morphology; presence of
rivers
Physical
Variables
tsunami wavelength
and amplitude
distance from tsunami
fault line, sea bed
depth 10 km offshore,
length of slope
extreme storm events;
distance from
hurricane event
coastal bathymetry
source distance &
coastal
geomorphology; type
of coastline; profile of
coastline; seabed
depth; distance to
tsunami source;
coastline orientation;
probability of
geologic event
ntEST —
Model
Field study and
model
Statistical
analysis
Observational
Review
Observational
Review
Observational
location of human
inhabitants _ .
, . , , Framework
relative to hazard
& vegetation
Location
Indian Ocean
Australia
Indian Ocean
Sri Lanka
Global
Sri Lanka
Australia
Global
Indonesia
Indian Ocean
Citation
Kunkel et al.
2006
Young &
Hardy 1993
Chatenoux &
Peduzzi 2007
Fernando et al.
2005
Lacambra et
al. 2008
Liu et al. 2005
Nott 1997
UNEP-WCMC
2006
Baird et al.
2005
Cochard et al.
2008
Socioeconomic benefits
human deaths & loss
of property
damage to human
lives & livelihoods
damage to human
lives & livelihoods
loss of human life
loss of village
avoided damages
presence of intact reefs
presence of reefs
absence of reefs
presence of reefs
presence of reef
cyclone
Review
Review
Observational
Observational
Observational
Economic
analysis
Indian Ocean
Global
Indonesia
Indonesia
India
SE Asia
EJF 2005
Sudmeier-
Rieux et al.
UNEP 2005
WI 2005
Whittingham
et al. 2003
Burke et al.
2002
-------
Ecosystem
Service Endpoint
Reef
Attributes
Physical
Variables
ntEST —
Location
Citation
Socioeconomic benefits (con 't)
length of coastline
within 2 km of
mapped coral reef
avoided damages
avoided damages
avoided property
damage due to
presence of reef
dollar value of
coastal protection
dollar value of
coastal protection
relative contribution
of reefs to shoreline
stability
cost of replacement
breakwaters
cost to restore
eroded beach
cost to replace with
concrete breakwaters
cost of replacement
artificial reef
eroding coastline
presence of healthy coral
reef; presence of
degraded coral reef
reef loss
coral reef locations,
"role of coral reefs"
presence of reef
coral loss
reef type; reef continuity ;
reef distance from shore
damage to reef due to
mining
presence of reefs
reef loss
presence of undamaged
reefs
length of coastline
coastal profile,
susceptible flood
zones, shoreline
stability,
storm regime
wave energy;
hurricane frequency;
coastal geology &
elevation;
coastal vegetation
breakwaters
Economic
analysis
level of shoreline Economic
development analysis
Economic
analysis
historical property _
. Economic
damage, property , .
. analysis
values
Economic
analysis
Economic
analysis
Economic
analysis
Economic
analysis
Economic
analysis
Case Study
Observational
presence of low
lying homes, cost „, , . .
. Observational
to repair
breakwaters
Caribbean
Indonesia
Worldwide
Bermuda
Philippines
Indonesia
Caribbean
Sri Lanka
Indonesia
Maldives
Maldives
Maldives
Burke &
Maidens 2004
Cesar 1996
Cesar et al.
2003
Beukering et
al. 2010
White et al.
2000
Pet-Soede et
al. 1999
WRI 2009
Berg et al.
1998
Riopelle 1995
Talbot &
Wilkinson
2001
Weber 1993
Wells &
Edwards 1989
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5
Coral Reefs: Natural Products
The flora and fauna of coral reef ecosystems are the source for a large number of pharmaceuticals and biochemicals and the
inspiration for a wide variety of chemical and structural models. This chapter describes some of these natural products, the
ecological forces that create them, and how they can be quantified as an ecosystem service.
5.1 Natural products as sources and templates for pharmaceuticals,
biochemicals, and biomaterials
Relative to their terrestrial counterparts, marine ecosystems are latecomers as sources and templates of pharmaceuticals,
biochemicals, and other biomaterials. This tardiness is mainly due to their inaccessibility, especially as compared to the easy
availability of terrestrial flora and fauna. Consequently, there is no indigenous medicine tradition to draw upon, because there
are so few instances (southern China being one) where marine species were used for medicinal purposes (Fenical 1996).
However, advances in undersea technology in the past few decades have gradually opened up marine ecosystems to more
systematic exploration. This exploration has revealed marine ecosystems to be complex and species-rich with a vast array of
predator-prey relationships that, coupled with the challenges of living in an aqueous medium, have resulted in a myriad of
secondary metabolites with extraordinarily complex, and hitherto unseen, structures (Fenical 1997; Gerwick 2008). From
1977-1987, the first decade of intensive marine exploration, about 2,500 previously unknown metabolites were reported
(Newman et al. 2000). Among the relatively small percentage of marine biochemicals that have been isolated, identified, and
tested are pharmaceuticals, nutraceuticals, cosmetics, food additives, antifouling agents, adhesives, and physical and chemical
templates in a variety of fields.
5.1.1 Pharmaceutical uses of marine natural products
Natural products have long been used for medicinal purposes. India's use of plants for medicinal treatment dates back over
5,000 years and has become codified in the Ayurveda, which contains over 8,000 herbal remedies. This same system of
treatment is still used in over 14,000 dispensaries in India today. During the period in which the Ayurveda was created, a
Chinese emperor was describing 365 herbal remedies, including ginseng, opium (the source of codeine and morphine), and
ephedra (the source of ephedrine). During the subsequent millennium, the Assyrians listed 250 medicinal plants, and the
Sumerians recorded 1,000 plants with medicinal properties (Huxley 1984). A Chinese herbal pharmacopoeia written about
2,000 years ago describes the use of marine seaweeds for medicinal purposes (Bowling et al. 2007).
More recently, from 2000 to 2006, about 50% of small molecule new chemical entities were natural products or based on
natural products (Newman & Cragg 2007). From 1981 through mid-2006, 63% of all new chemical entities were natural
products or based on natural products (Newman & Cragg 2007; Cragg & Newman 2009). However, over 50% of marketable
pharmaceutical products are consistently natural products or based on natural products (NRC 1999, p. 73), and half of all cancer
drug research is devoted to marine natural products (Fenical 1996). Further, about 50% of the drugs introduced from 1994 to
the present are either natural products or based on natural products (Harvey et al. 2010).
There are over 10,000 marine biochemicals with potentially useful pharmacological properties, and Appendix 5-A lists over
200 of them, showing their biological source, geographic location, chemical name and structure, the nature of their biological
activity, and their approval status, where applicable. There were 36 marine-derived natural products that were in clinical trials
(Table 5-1) or approved for use as of 2006 (Table 5-2), and 20 or more that were in the preclinical stage of testing (Simmons
etal. 2005; Wijffels 2007).
A secondary metabolite is a substance produced by an organism that seemingly has no direct role in the organism's metabolism, though they
are often produced via pathways that are derived from primary metabolic pathways. It is believed that they are created because they confer
some evolutionary advantage, particularly in nonmotile organisms. Most often secondary metabolites are used by the organism in intraspecies
or interspecies interactions usually related to defense or signaling (e.g., for reproduction) (NRC 1999, pp. 74-75; Croteau et al. 2000,
pp.1250-1, 1316; Seigler 2002, p. 3; Wink 2003).
"In the fields of pharmacology and biochemistry, a small molecule is a low molecular weight organic compound which is by definition not a
polymer. The term small molecule, especially within the field of pharmacology, is usually restricted to a molecule that also binds with high
affinity to a biopolymer such as protein, nucleic acid, or polysaccharide and in addition alters the activity or function of the biopolymer. The
upper molecular weight limit for a small molecule is approximately 800 Daltons which allows for the possibility to rapidly diffuse across cell
membranes so that they can reach intracellular sites of action. In addition, this molecular weight cutoff is a necessary but insufficient
condition for oral bioavailability." (Wikipedia 2010)
A new chemical entity is a drug that contains no active moiety that has been approved by the FDA (21CFR314.108).
-------
Table 5-1. Testing stages for new pharmaceuticals
Trial Stage
Preclinical
Phase 0
Phase I
Phase II
Phase III
Phase IV
Nature of Testing
in vitro tests and in vivo animal model tests for preliminary dose ranging, efficacy,
toxicity, and pharmacokinetic evaluation
instituted by FDA in 2006; single, subtherapeutic dose; 10-15 human subjects; on-
going evaluation of phase's usefulness, ethics, and claimed benefits (to save
money and to speed up the approval process)
normally, first stage of human testing; trials assess the drug's safety, tolerability
(including maximum tolerable dose), pharmacokinetics (effects of drug), and
pharmacodynamics (metabolism of drug); 20-100 healthy volunteers or patients
with the target disease
continues evaluation of the drug's safety in larger groups of subjects (200-300
patients with the target disease); evaluation of the drug's efficacy, optimal dosing
regimen, and side effects; often double-blind tests versus placebo
multi center trials involving hundreds or thousands of subjects; trials attempt to
establish the drug's efficacy vis-a-vis current best practice and an overall risk-
benefit ratio in a demographically diverse sample of patients with the target
disease; usually drug versus standard treatment
post-approval monitoring of large populations of patients taking the drug; may
evaluate drug's usefulness in treating diseases other than the original target
Source: University of Pittsburgh (2002); Wikipedia (2011).
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Table 5-2. Status of marine-derived natural products in clinical trials or clinical use
Compound Name
Abyssomlcin C
ACV1 (aka a-conotoxin)
Acyclovir (aka Zovirax®)
Aplidine |
(aka plitidepsin.
Aplldin®)
Ara-A
(aka vidarabine, Vira-A®)
Ara-C f
(aka cytarabine,
Cytosar-U®, DepoCyt®,
Tarabine PFS®)
AZT (aka Retrovir®,
zidovudine)
Bryostatin 1 f
Cematodin f
(aka LU103793)
Contulakln-G
(aka CGX-1160)
Diazepinoinicin
(aka ECO-4601)
Discodennollde f
DMXB
(aka GTS-21, DMXB-A)
Ecteinascidin 743 f
(aka trabectedin.
Yondells®)
Eribulln f &
Eribulln inesylate
(aka E7389, Halaven®)
Hemlasterlln f
(aka E7974)
IPL-576,092
(aka HMR-4011A)
Source
actinobacterium
(Vermcosispora marts)
mollusk (cone snail)
(Conus victoriae)
sponge
(Cryptothetya crypto)
tunicate
(Aplidium albicans)
sponge
(Cryptothetya crypto)
sponge
(Cryptothetya crypto)
sponge
(Cryptothetya crypto)
bryozoan
(Bugula neritina)
mollusk (sea hare)
(Dolabella auricularia\
cyanobacterium (Symploca sp.)
mollusk (cone snail)
(Conus geographus)
actinobacterium
(Micromonospora sp.)
sponge
(Discodermia dissoluta)
marine worm
tunicate
(Ecteinascidia turbinata)
sponges (Halichondria okadai,
Axinella sp., Phakellia carteri, &
Lissodendoryx sp.) (or possibly
their symbiotic4 bacteria)
sponges (Hemiasterella minor,
Auletta sp., Cymbastela sp.
& Siphonochalina sp.)
sponge
(Petrosia contignata)
Status (Disease)
Phase I (antibiotic)
Phase I (analgesic)
Clinically available
(antiviral)
Phase II (cancer)
FDA approved in 1976
(antiviral)
Clinically available;
FDA approved in 1969;
Phase I/II (cancer)
Clinically available
(antiviral)
Phase I/II (cancer);
Phase II (Alzheimer's)
Phase I/II (cancer)
Phase I (analgesic);
Phase II late 2005
Phase I (antibiotic, cancer)
Phase I (cancer)
Phase II (Alzheimer's.
schizophrenia)
Phase II/III (cancer) in 2003-
2005; approved by EMA* for
treatment of soft tissue sarcoma
Phase II/III (cancer);
FDA approved for late-stage
breast cancer in 2010.
Phase I (cancer)
Phase II (anti-asthmatic)
successfully completed
Comment
Metabolic Pharma (Australia)(06/2006);
conotoxin Vc 1 . 1
Synthetic analog of arabinose nucleotides;
antiviral used to treat herpes infections; King
Pharmaceuticals discontinued marketing in June
2001, possibly due to superior alternatives
Dehydrodidemnin B; synthetic analog
Synthetic analog of arabinose nucleotides;
antiviral used primarily for ophthalmic infections
Approved by FDA in 1969; first marine anticancer
drug; synthetic analog of arabinose nucleotides;
sold by Pharmacia & Upjohn
Synthetic analog of arabinose nucleotides; first
drug licensed for treatment of HIV; sold by
Glaxo SmithKline
Now in combination therapy trials; licensed to
GPC Biotech by Arizona State Univ.
Synthetic analog of dolastatin 15; some positive
effects in melanoma; studies discontinued in 2004
Cognetix and Elan Corporation (Ireland)
Ecopia BioSciences (Canada)
Licensed to Novartis by Harbor Branch Oceano-
graphic Institution; studies may have been discon-
tinued in 2005
Licensed to Taiho by the Univ. of Florida
Licensed to Ortho Biotech (J&J/Janssen Pharma-
ceuticals); PharmaMar currently makes and sells
Yondelis® in Europe
Eisai's synthetic halichondrin B derivative;
breast, prostate, & nonsmall cell lung cancer
(NSCLC) cancers
Eisai's synthetic analog of hemiasterlin; being
tested against colorectal cancer
Derived from contignasterol; Inflazyme Pharma
Symbiosis is a close relationship between two or more organisms of different species. There are four forms: amensalism, commensalism,
mutualism, and parasitism: amensalism is when one species is harmed while the other is unaffected; commensalism is when one species
benefits while causing little or no harm to the other; mutualism is when both species benefit from the relationship; and, parasitism is when
one species benefits and the other is harmed. We use symbiosis throughout this chapter, because the exact nature of the relationship is often
unknown or unspecified.
-------
Table 5-2. (continued)
Compound Name
IPL-512,602
(aka AVE 0547)
IPL-550,260
Irvalec® f
(aka ellsidepsin,
PM02734)
Source
Synthetic analog
Synthetic analog
mollusk (sea slug)
(Elysia rufescens)
green algae (Bryopsis sp.)
KRN-7000 f (aka a-GalCer, sponge
a-galactosylceramlde) (Agelas mauritianus)
LAF-389 f
LBH-589 (aka Faridak®,
panobinostat)
Marizomlb f
(aka salinosporamide A
& NPI-0052)
Neovastat f
(aka M-941)
NVP-LAQ824 f
(aka dacinostat)
Pllnabulin f
(aka NPI-2358)
Pseudopterosins
Soblldotin f
(aka auiistatin PE
& TZT-1027)
Spisulosine
(aka ES-285)
Squalamlne f
(aka Evizon™)
Synthatodin f
(aka ILX651, tasidotin)
Taltobulin f
(aka HTI-286)
Trodusqueinine
(aka MSI-1436)
Zalypsis® f
(aka PM1004)
Ziconotide
(aka Prialt®)
sponge
(Jaspis sp.)
Psamaplysilla spp.
(sponge)
actinobacterium
(Salinispora tropica)
shark
Synthetic combination
of three natural products
fungus
(Aspergillus sp.)
gorgonian (sea whip)
(Pseudopterogorgia elisabethae)
mollusk (sea hare)
(Dolabella auricularia)
cyanobacteria
(Symploca hydnoides
Lyngbya majuscula)
mollusk (arctic surf clam)
Spisula polynyma
(aka Mactromeris polynyma)
shark (spiny dogfish)
(Squalus acanthias)
mollusk (sea hare)
(Dolabella auricularia)
cyanobacteria (Symploca sp.)
sponges (Hemiasterella minor,
Auletta sp., Cymbastela sp.
& Siphonochalina sp.)
shark (spiny dogfish)
(Squalus acanthias)
mollusk (nudibranch)
(Jorunnafunebris)
mollusk (cone snail)
(Conus magus)
Status (Disease)
Phase II (anti-asthmatic)
Phase I (anti-asthmatic)
Phase II (cancer)
Phase I/II (cancer)
Phase I (cancer)
Phase III (cancer)
Phase I (cancer)
Phase II/III (cancer)
Phase I (cancer)
Phase I/II (cancer)
Phase II (anti-inflammatory)
Phase III (cancer)
Phase I (cancer)
Phase II (cancer &
macular degeneration)
Phase I/II (cancer)
Phase I/II (cancer)
Phase I (diabetes treatment;
weight loss)
Phase II (cancer)
Clinically available
(neuropathic pain)
Comment
Derived from IPL576,092; with Aventis;.
no further data as of 08/2005
Derived from IPL576,092; with Aventis;.
no further data as 08/2005
Synthetic analog of kahalalide F created to insure
sufficient supply; licensed to PharmaMar by Univ.
of Hawaii
An agelasphin derivative
Synthetic analog of bengamide B; may have been
withdrawn in 2006
Synthetic analog of psammaplin; with Novartis
Proteasome inhibitor; Nereus Pharma
Defined mixture of <500 kDa from cartilage; anti-
angiogenic; possibly withdrawn March 2007
Derived from psammaplin, trichostatin, and
trapoxin structures; possibly withdrawn in 2006
Synthetic analog of marizomib; selective tumor
vascular disrupting agent (VDA)
Used in Estee Lauder's Resilience skin cream
Synthetic derivative of dolastatin 10; no positive
effects found in Phase II trials, but appears to be
effective in combination therapy with vinca alka-
loids and bryostatin
Rho-GTP inhibitor
Anti-angiogenic activity is basis for its use to treat
both cancer and wet form age-related macular
degeneration; Evizon™ is the name used for the
ophthalamic formulation
Synthetic analog of dolastatin 15;
for melanoma, breast, and nonsmall cell lung
cancer (NSCLC)
A synthetic analog of hemiasterlin; studies may
have been discontinued in 2005
Genaera started Phase I in 2007 and reported
promising results in 2009. Shortly after, Genaera
was dissolved, and trodusquemine was sold to Ohr
Pharmaceuticals; current status unknown
A synthetic analog of jorumycin, safracin B, &
saframycin B; made by PharmaMar
Licensed by Elan to Warner Lambert; approved by
FDA in Dec 2004; also approved by EMA*;
chronic use does not result in tolerance
* EMA: European Medicines Agency
•f The 20 anticancer products used as a basis for the charts in Figure 5-3.
Source: primarily Fenical (2006), Simmons & Gerwick (2008), and Mayer et al (2010). Also Baerga-Ortiz (2009), Butler (2005), Dumez et
al. (2007), Glaser (2007), Gross & Konig (2006), Gullo et al (2006), Hunt & Vincent (2006), Lam (2006), Nereus (2010), Newman
& Hill (2006), Sashidhara et al (2009), UN (2007, pp. 26-27), and Yuan et al (2006).
-------
Identifying the optimum process of drug discovery has been the subject of a fairly contentious debate over the past three
decades. Although natural products have long been the primary source of new drugs, the difficulty of finding biologically
active chemicals, isolating and testing them, and then maintaining a steady supply of the chemical's source created an impetus
in the pharmaceutical industry to turn to synthetic combinatorial chemistry, which was later coupled with high-throughput
screening methods. The allure of de novo synthesis of new drugs induced most pharmaceutical firms to shift resources from
natural products research to combinatorial chemistry, and many firms shuttered their natural products research entirely.
However, despite large investments in combinatorial chemistry, it has yielded only one new drug that has been approved for
use (Nexavar® [aka sorafenib] in 2005) (Newman & Cragg 2007). Milller et al. (2004) show that natural products are much
more likely than combinatorial chemistry products to yield a drug approved for clinical use (Table 5-3).
Table 5-3. Number of candidate drugs reaching different stages of clinical trials
Natural Products
(Secondary Metabolites)
Available for study 200
Preclinical 200
Phase I trials »10
Combinatorial Chemistry
(Synthesized Chemicals)
5,000-10,000
200
10
Clinical use »1 ~1
Source: Miiller et al. (2004).
The situation is nicely explained by Firn (2003):
In contrast to the chemists, organisms use enzymes instead of chemical reagents to bring about chemical transformations.
The crucial advantage of using enzymes in biosynthetic sequences is that enzymes can bring about specific structural
changes to very specific sites in a complex molecule. This facility of microbes and plants to make structurally complex
molecules with relative ease means that humans inevitably find it hard to manufacture natural products.
The difficulty of creating new pharmaceuticals using only combinatorial chemistry was made abundantly clear by
GlaxoSmithKline's announcement that a six-year effort to discover broad-spectrum antibiotics failed because of the limited
chemical diversity of their synthetic screening libraries (Williams 2008). The best strategy may be to combine these two
approaches by seeking bioactive chemical structures from natural sources and then optimizing those structuresS via
combinatorial chemistry. By acknowledging the prominent role of natural products in drug discovery, this joint disco very-then-
optimization strategy may highlight the importance of preserving marine biochemicals.
Despite its potential, marine drug discovery faces some difficult challenges, not the least of which is acquiring a sufficient
quantity of the marine source material to allow extraction of a testable quantity. While the quantities of end product may seem
small, steps in the development of the end product require much more raw material, and the quantity of source material needed
rapidly escalates as the chemical proceeds through the discovery and testing process (Table 5-4). A vivid example of how much
source material is required is that of bryostatin. For the initial clinical trials, 13,000 kg of Bugula neritina were collected and
processed using large-scale chromatographic techniques, yielding 18 g of bryostatin 1 (about 1.4 mg per kg or 1.4 ppm)
(Newman & Cragg 2004). Such small yields are not uncommon: the concentration of halichondrin B in Lissodendoryx spp. is
~0.4 mg per kg, and of halistatin in sponges is 8.8 (ig per kg (Molinski et al. 2009). It seems likely that these secondary
metabolites appear in such minute quantities because of their extremely potent biological activity (Gerwick 2008).
In some rare instances, a bioactive chemical can be used directly as a drug and can be obtained in quantities sufficient for
therapeutic use. Ziconotide (Prialt®), a toxin from Conus snails, is such a rarity; it was the first "direct from the sea" approved
drug (Donia & Hamann 2003; Newman & Cragg 2007). For terrestrial sources of pharmaceutical natural products, cultivation
of the drug's source to produce marketable quantities is usually feasible, but there has been little success in attempts to cultivate
marine sources to produce such quantities (Donia & Hamann 2003). As a result, chemical synthesis of the natural product is
often the only available option for producing sufficient quantities of the bioactive chemical. Even so, synthesis is not always an
option, as succinctly stated by Donia and Hamann (2003):
Unfortunately, the structural complexity of marine molecules, which suggests novel mechanisms of action and high
selectivity, has also resulted in few economically feasible strategies for total chemical synthesis.
It is often the case that an effective dose of a bioactive natural product is either too toxic or produces unacceptable side effects. These
undesirable effects can usually be reduced by selectively modifying the chemical structure of the natural product (Newman et al. 2000). For
example, salicylic acid from willow trees was acylated to form acetylsalicylic acid (aspirin), which is less irritating to the gastrointestinal tract
than the natural product.
-------
Table 5-4. Timeline of drug development and amount of raw material and pure product needed at each stage
Stages
Collection of field samples & bioactivity screening
Identification, isolation, & purification of bioactive compound
Determination of chemical structure
Identification of mechanism of action and potential for synthesis
Preclinical trials
Clinical trials
Length of Stage
(approx.)
1-2 years
1-2 years
2-A, years
4-6 years
Raw Material
Needed
-0.1-1 kg
-1-5 kg
>5kg
>10,000 kg)
Pure Chemical
Needed
—
1-10 ug
1-10 mg
1-10 g
Source: Koehn & Carter (2005); Hunt & Vincent (2006).
A steady progression of advances in chemical synthesis methods combined with the joint discovery-then-optimization strategy
described above may make more of these syntheses economically feasible. It is worth noting that even when synthesis is
achieved, the synthetic product may not have the same structure and biological activity as the natural product. Pettit and Taylor
(1996) report an instance where the natural product (stylopeptide 1), despite seeming to be pure based on all physicochemical
measurements, was in fact in association with a halistatin-like poly ether compound that was a highly active anticancer agent.
The presence of this poly ether eluded the usual physical, chromatographic, and NMR tests and could only be detected using
biological methods. As a result, synthetic stylopeptide 1 had none of the biological activity attributed to the natural product
even though the two were structurally identical (Newman & Cragg 2004).
Although marine microbes6 have long been considered a likely and potentially significant source of bioactive chemicals
(Fenical 1982; Kaul & Daftari 1986; Franco & Coutinho 1991; Fenical 1993), recent discoveries suggest that their importance
could exceed expectations. Initially, marine microbes were viewed as likely counterparts to terrestrial microbes, which are the
source of many antibiotics. However, in the past decade, it has become apparent that many of the bioactive chemicals attributed
to higher order flora and fauna (e.g., sponges and nudibranchs) are in fact created by symbiotic microorganisms (often algae,
actinomycetes, cyanobacteria, or fungi) (Donia & Hamann 2003; Leeds et al. 2006; Wase & Wright 2008).
Marine microbes represent a surprisingly large amount of biomass: they can account for more than 60% of a sponge's wet
weight (Wilkinson [1978] as cited by Bowling et al. [2007]). In total, marine prokaryotes (bacteria and archaea) outnumber
their terrestrial counterparts; Whitman et al. have estimated that there are about 3.67 x 1030 prokaryotic cells in marine
ecosystems (give or take a few billion), and that there are about 305 Pg of carbon in these cells.
Marine microbes demonstrate a surprising degree of host specificity, both with respect to different species in the same location
and to the same species in different locations. Of the 100 bacterial species found on three nearby sessile organisms, only two
were common to all three (Longford et al. [2007] as cited by Penesyan et al. [2010]). Further, the microbial community found
in the coral Montastraea franksi had almost no overlap with the microbes found in the surrounding seawater (Rohwer et al.
[2001] as cited by Penesyan et al. [2010]).
Despite their great potential as sources of new drugs or structural templates, marine microbes pose a considerable challenge,
because fewer than five percent of them can be grown in standard laboratory or industrial conditions. To overcome this
problem, it may be possible to use metagenomic techniques to move the section of the microbe's genome responsible for
creating the bioactive chemical into the genome of a microbe that is already used in large-scale fermentation processes (Donia
& Hamann 2003). However, many microbes will not produce the bioactive compounds of interest if they are not in contact with
their symbiont host (Wijffels 2007), so a means must be devised to replace the biochemical signaling between microbe and host
that activates the transferred genetic sequence. It seems likely that metagenomic techniques could solve this problem, but it has
yet to be demonstrated. Although it has great promise and has already been successfully used, metagenomics is still early in its
development, and it would be imprudent to depend on it exclusively. As pointed out by Udwary et al. (2008, p. 521),
metagenomics has its drawbacks (including cost and complexity) and has had its failures (discodermolide). Metagenomics is
probably best seen as one of the arrows in the quiver rather than the only arrow.
The term "microbe" is used throughout this chapter to refer collectively to unicellular or colonial microorganisms, including bacteria, fungi,
archaea, or protists.
-------
5.1.2 Marine natural products as structural templates for synthesis
The many unique chemical structures found in marine species provide templates that can be used in the synthesis of new drugs
and for insight into structural possibilities. One of the more significant contributions to medicinal chemistry was the discovery
that sugars other than ribose or deoxyribose were constituents of naturally occurring nucleosides. This discovery resulted from
the isolation of spongouridine and spongothymidine from marine sponges in the early 1950s, and it heavily influenced drug
development for the next 30 years. According to Newman et al. (2000):
These two compounds can be thought of as the prototypes of all of the modified nucleoside analogues made by chemists
that have crossed the antiviral and anti-tumor stages since then. Once it was realized that biological systems would
recognize the base and not pay too much attention to the sugar moiety, chemists began to substitute the 'regular pentoses'
with acyclic entities, and with cyclic sugars with unusual substituents. These experiments led to a vast number of
derivatives that were tested extensively as antiviral and anti-tumor agents over the next thirty plus years... such structures
evolved in the (then) Wellcome laboratories, leading to AZT and, incidentally, to Nobel Prizes for Kitchens and Elion,
though no direct mention was made of the original arabinose-containing leads from natural sources.
5.1.3 Marine natural products as molecular probes
Molecular probes are chemicals that are used to explore and elucidate biochemical structures and processes at the cellular and
molecular levels. A great many marine biochemicals that are biologically active but unusable as drugs are used extensively as
molecular probes (some examples are shown in Table 5-5). Marine neurotoxins, including tetrodotoxin, saxitoxin, conotoxin,
and lophotoxin, have been used with great success to advance our understanding of a wide variety of receptors and ion channels
in the operation of nervous systems. "The importance of molecular probes in resolving the complexities of diseases and cellular
processes has often outweighed any value that they would have as commercial drugs." (NRC 1999, p. 79)
Table 5-5. Marine biochemicals used as molecular probes
Chemical
adociasulfate-2
brevitoxin
conotoxins
jaspamide
(jasplakinolide)
latrunculin A
lophotoxin
manoalide
okadaic acid
saxitoxin
swinholide A
tetrodotoxin
Use
selectively inhibits the intracellular molecular motor protein kinesin
sodium channel inhibitor in nerves and muscle
calcium channel inhibitor (co-conotoxins);
block voltage-gated sodium channels (u- & uO-conotoxins);
delay inactivation of sodium channels (5-conotoxins);
potassium channel inhibitor (K-conotoxins);
inhibits norepinephrine transporter (/-conopeptides);
nicotinic acetylcholine receptor (a-conotoxins);
blocks type 3 serotonin receptors (a-conotoxins);
inhibits aradrenergic receptor (p-conopeptides);
inhibits NMDA receptor (conantokins);
vasopressin receptor agonist (conopressins);
neurotensin receptor agonist (contulakins)
selective binding agent to the intracellular actin network
selective binding agent to the intracellular actin network;
used to explore the role of phospholipase A2 in inflammation
irreversible nicotinic receptor antagonist
selective inhibitor of the inflammation enzyme
phospholipase A2
potent and selective inhibition of phosphatases
inhibits calcium, potassium, & sodium channels in nerves and muscles
selective binding agent to the intracellular actin network;
severs F-actin filaments; binds G-actin filaments
inhibits calcium, potassium, & sodium channels in nerves and muscles
Reference
NRC 1 999; Bri ^ et al. 2006
Al-Sabi et al. 2006;
Karunasagar & Karunasagar 2008
NRC 1999;
Layer & Mclntosh 2006;
Lewis 2009
Senderowicz et al. 1995;
Saito 2009; Robinson et al. 2010
Matthews et al. 1997;
Amagata et al. 2008
Karunasagar & Karunasagar 2008
Fusetani & Kern 2009
Glaser & Jacobs 1986;
Yasuhara-Bell et al. 2006
NRC 1999
Hay & Fenical 1996; NRC 1999;
Al-Sabi et al. 2006
Bubb et al. 1995; Saito 2009
Hay & Fenical 1996; NRC 1999;
Al-Sabi et al. 2006;
Fusetani & Kern 2009
-------
5.1.4 Nonpharmaceutical uses of marine natural products
Marine natural products have a wide variety of nonpharmaceutical uses, both as products and as templates or sources of insight
that may lead to innovative new products (Table 5-6). Perhaps the largest and most economically significant use at this time is
in antifouling coatings. In a review of the literature, Chambers et ol. (2006) found that potential antifouling products had been
found in 160 marine species from a wide variety of phyla (Figure 5-1).
Table 5-6. Current and potential nonpharmaceutical uses for marine natural products
Source Use Reference
algae
algae (green)
(Chlamydomonas
reinhardtii)
algae (green, cryptophyte)
annelid (sandcastle worms)
(Phragmatopoma californica)
cnidarian (corals)
(Porites spp.)
echinoderm (various)
echinoderm (brittle stars)
(Ophiocoma wendtii)
fish (sharks)
fish (sharks)
fish (zebrafish)
(Danio rerio)
fish (zebrafish)
(Danio rerio)
mammal (dolphin)
(Tursiops truncatus)
mollusk (abalone)
(Haliotis spp.)
mollusk (mussels)
(Mytilus californianus,
M. galloprovincialis)
mollusk (mussels)
(Mytilus edulis)
mollusk (scaly-foot snail)
(Crysomallon squamiferum)
porifera (sponge)
(Agelasspp.)
tunicate
tunciate (ascidian)
(Pegea confoederata)
cultured for production of jet fuel
surrogate production of drugs
more efficient light-harvesting in photosynthesis
medicinal glues and adhesives that function under water
highly interconnected microporous structures of calcium carbonate
(aka aragonite from corals) or calcite (from echinoderms) used as hard
tissue prostheses, cardiovascular material, and tracheal prostheses
development of biomimetic compound lenses that minimize optical
aberrations while maximizing focal length or field of view
shark-skin textured material stops bacterial growth
shark-skin textured surface improves fuel economy by reducing drag
gene expression activation to repair and replace damaged cardiac cells
damaged
use of zebrafish model to identify genes and proteins that promote
melanoma
genetic pathway for controlling blood sugar levels
nacre (the inside lining of abalone shells) is a template for tough,
lightweight structural coatings for buildings and airplanes
medicinal glues and adhesives that function under water
medicinal glues that bonds to living tissue and adheres in wet environ-
ments; used to repair human fetal membranes
unique shell structure (fortified with iron sulfide) in a deep-sea snail
found near hydrothermal vents will improve helmets & body armor
ageliferin renders previously resistant bacterial biofilms susceptible to
antibiotics
cellulose from ascidians has a nanoscale structure that can be used to
structurally align skeletal muscle tissue grown in the laboratory
the most efficient filter feeders may help remove carbon from ocean
surface water thereby limited CO2 to atmosphere
Guardian 20 10
Rasalaetal. 2010
Collinietal. 2010
Fountain 20 10
White & White 2002
Fee & Szema 2005
Sharklet Fechnologies
2010
Bhushan 2009
Joplingetal. 2010;
Kikuchirfa/. 2010
Ceol et a\. 2008;
Ceo\etal. 2011;
White et al. 2011
Venn-Watson &
Ridgway 2007
Mayer 2005
Messersmith 2010
Harrington etal. 2010
Benedict 2002
BiYicetal. 2010
Yaoetal. 2010
Huigens et al. 2008
Dugan etal. 2010
Sutherland et al. 2010
-------
Cnlorophyla,
Rhodophyta, &
Heterokontophyta
1)9.5%)
Figure 5-1. Phyletic distribution of 160 reviewed marine species from which potential antifouling
natural products have been extracted (figure adapted from Chambers et al. 2006)
5.2 Sustaining the presence of natural products in coral reef ecosystems
Maintaining the richly diverse cornucopia of marine natural products requires some understanding of the dynamic ecosystems
that drive their creation. Two of the contributing factors are the great diversity of marine life and the density of that life in the
marine environment. Of the approximately 35 phyla, every phylum but one has marine species (NRC 1999, p. 74; Davidson &
Erwin 2006). In 1999, the NRC (loc. cit.) stated that the 200,000 marine species that had been described to that point
represented "a small percentage of the total number of species that have yet to be discovered and described." Bouchet (2006)
gives a range of 230,000-275,000 marine species, with 1,300-1,500 new species being identified every year. The First Census
of Marine Life (CoML 2010, p. 11) states that as of 2010, experts believe that there are about 244,000 cataloged marine species
and that this number will rise to about 250,000 in the next few years. However, the CoML goes on to state that the consensus of
the CoML scientists is that at least a million marine species are likely to exist. In other words, for every marine species that has
been identified, three more species are yet to be discovered.
Coral reefs are thought to have the highest species density of any marine ecosystem, with some areas having about 1,000
species per square meter (Donia & Hamann 2003). This combination of high species diversity and high species density leads to
the profusion of secondary metabolites, with over 18,000 unique chemical structures having been identified (Gerwick 2008,
p. 428). This may be just the tip of the iceberg. A metagenomic analysis of 1,800 species (primarily unicellular) found in
seawater samples collected from the Sargasso Sea near Bermuda predicts more than six million proteins, more than double the
number that have been identified (Yooseph et al. 2007; Cragg & Newman 2009). Hunt and Vincent (2006) have attempted to
map the global distribution of previously unknown marine biochemicals, but the resulting map may better represent the location
and frequency of natural product explorations than where novel chemicals are to be found.
The species numbers just described do not include microorganisms7. This is practical but may be somewhat misleading,
because many of the natural products of pharmaceutical interest apparently originate in microorganisms. With respect to marine
microorganism diversity, the NRC (1999, p. 78) stated, "Most of the Earth's microbial diversity is found in the ocean." While
some can be found in both terrestrial and marine ecosystems, many taxonomic classes of microorganisms exist only in the sea,
including ones that have adapted to a variety of extreme environments, such as hypersaline conditions, enormous hydrostatic
pressures, hydrothermal vent temperatures, and high-sulfur environments. Based on what is known from the small areal extent
that has been surveyed, it is estimated that several million marine microorganism species exist (Gerwick 2008, p. 427). The
CoML (2010, p. 12) research has led to a hundredfold increase in estimates of the number of marine microbe genera and to an
estimate that there may be as many as a billion types of marine microbes.
The adaptability of microorganisms may be the reason why different populations of the nearly ubiquitous bryozoan Bugula
neritina yield very different amounts of bryostatins, a family of potential anticancer drugs. The few B. neritina populations that
It should be noted that only microbes that are protists (eukaryotes) are organized into species. Prokaryotes (including bacteria and archaea)
are organized by kind, or phylotype (CoML 2010, p. 12).
-------
produce detectable concentrations of the bryostatins are spatially scattered and are at depths greater than 9 m. A metagenomic
analysis showed that the symbiotic microorganisms (Candidatus Endobugula sertula) from B. neritina harvested at greater than
9 m varied by 8% in its mitochondrial carboxylase I sequences from those harvested at less than 9 m (Newman & Cragg 2004).
This suggests a pitfall for simplistic or insufficiently informed conservation strategies: implementing a conservation strategy for
B. neritina (had one been needed) that preserved the wrong populations would have resulted in losing the potential anticancer
benefits of the bryostatins (Hay & Fenical 1996).
A healthy reef with high biodiversity may increase the probability that any given species could be the source of a marketable
product. As such, the probability of a bioprospecting discovery may be represented as directly proportional to the state of the
reef (van Beukering & Cesar 2004). Certain benthic habitats may foster greater sponge diversity and abundance (Mumby et al.
2008), which, in turn, may foster the development of secondary metabolites with pharmaceutical potential by the sponges and
their symbiont microorganisms.
5.2.1 Ecological role and sources of secondary metabolites
The flora and fauna inhabiting marine ecosystems are confronted with a very complex and stressful environment, and the
biochemical adaptations made in response to those stresses have resulted in a vast trove of natural products with unprecedented
structural complexity. Marine ecosystems have existed far longer that terrestrial ecosystems—thereby providing greater
opportunity for evolutionary adaptation (e.g., cyanobacteria have existed for about 3.5 billion years [Gerwick 2008, p. 428]).
The immersion of marine species exposes them to attack by predators and pathogens to a much greater degree than in terrestrial
ecosystems. Space is also a significant stressor, though it might not seem so given the oceans' expanse. The shallow coastal
zones and infrequent seamounts (places where sunlight can be a useful source of energy) constitute a relatively small area,
within which competition can be intense. In particular, sessile organisms not only need room to grow, but they need to prevent
other sessile organisms from growing on top of them (Pawlik 1993).
Marine flora and fauna have adapted to the stresses inherent in their complex ecosystems by creating highly unusual and
complex secondary metabolites. Faulkner (2000) provides a concise explanation of how these complex biochemicals came to
be:
It is probable that chemical defense mechanisms evolved with the most primitive microorganisms but have been
replaced in many more advanced organisms by physical defenses and/or the ability to run or swim away and hide.
Sessile, soft-bodied marine invertebrates that lack obvious physical defenses are therefore prime candidates to possess
bioactive metabolites. If it is assumed that secondary metabolites evolved from primary metabolites in a random
manner, any newly produced secondary metabolite that offered an evolutionary advantage to the producing organism
would contribute to the survival of the new strain. The specific evolutionary pressures that led to chemically rich
organisms need not be defined but the longer the period of evolution, the more time the surviving organism has had to
perfect its chemical arsenal. Sessile marine invertebrates have a very long evolutionary history and have had ample
opportunity to perfect their chemical defenses.
In creating these chemical defenses, marine organisms use chemicals and pathways that are distinctly different than their
terrestrial counterparts (Hay & Fenical 1996). Secondary metabolites are usually associated with immobility, so in terrestrial
ecosystems they are most often found in plants. In marine ecosystems, secondary metabolites occur more widely both because
both plants and animals (along with their symbiotic microbes) may be sessile and because of the high rate of predation (Hay &
Fenical 1996). Interestingly, it is usually the case that more is known about the effect of a secondary metabolite on human
cellular biochemistry than its function in the species from which it was isolated (NRC 1999, pp. 74-75).
The very nature of secondary metabolites (i.e., as biochemicals produced specifically to interact with biological processes)
makes them more attractive to natural product chemists and far more likely than a randomly chosen organic structure to be
biologically active. As Faulkner (2000) points out,
Chemical defense mechanisms cannot be directly equated with potential biomedical activity, but it is remarkable how
well the two correlate in reality. This could be explained by the fact that targets of the chemical defenses, primary
metabolites such as enzymes and receptors, are highly conserved compared with secondary metabolites.
Simmons & Gerwick (2008, p. 433) describe four distinctive characteristics of marine secondary metabolites:
From the species studied to date, it is clear that marine organisms have been subject to unique adaptive pressures and
utilize rather different strategies for producing secondary metabolites compared to their terrestrial counterparts. In some
cases, seasoned organic chemists look at the structures of metabolites produced by marine life and characterize them as
bizarre, unlike anything found from the land environment. Alternatively, some marine metabolites are of exceptional
complexity representing true milestones of human achievement in the characterization of their convoluted multicyclic,
and three-dimensional structures, such as maito toxin... Coupled to the uniqueness of their physical structure are their
biological properties, which can be exquisitely potent against some cellular targets. Indeed, some of the most potent
natural toxins on the planet derive from marine life...Perhaps even more important than potency is the fact that some of
these marine metabolites exert their pharmacological activities through interactions at novel drug sites, such as
-------
enzymes or receptors not targeted by any current pharmaceutical agent. Hence, the real possibility exists that entirely
new drug classes will be discovered that have novel structures and new sites of action...
The potency of marine natural products is probably attributable to their inevitable dilution in seawater; that is, the chemical
must be sufficiently powerful to overcome the dilution that will take place en route to its target and still have the desired effect
(Newman & Cragg 2004). A fifth distinctive characteristic of marine secondary metabolites is the relatively high frequency
with which they incorporate chlorine or bromine in their chemical structure, probably resulting from the easy availability of the
halogens in seawater (Gerwick 2008, p. 428). It may also be that the toxicity often conferred by halogenation served to provide
an evolutionary advantage.
Historically, marine macroorganisms , particularly sponges and seaweeds from tropical waters, were thought to be the greatest
source of these secondary metabolites (Gerwick 2008, p. 428). However, several different lines of inquiry have led to the
understanding that the source is often one of the multitude of symbiotic microorganisms associated with the macroorganism.
Probably the first evidence of this was the discovery of identical secondary metabolites in different species of macroorganisms.
In some cases, this was a matter of macroorganisms ingesting the same species of microorganism. However, in other cases the
association is symbiotic. Metagenomic analyses have confirmed that the symbiotic microorganisms, not the macroorganisms,
possessed the genetic sequences capable of producing the biologically active secondary metabolites in question. It is fascinating
that these microorganisms produce secondary metabolites not for their protection, per se, but for the protection of their
symbiont macroorganisms, suggesting a lengthy co-evolution that may also include sophisticated biochemical signaling
between the species (e.g., the bacteria that grow on fish eggs protect the eggs from fungal infection) (Fisher 1983a; Fisher
1983b; Fisher & Clark 1983).
This state of affairs creates several difficulties. The first is that the identification of threatened or endangered ecosystems or
species routinely focuses on macroorganisms. The obvious importance of the symbiotic relationship between marine
macroorganisms and microorganisms clearly emphasizes the necessity of adopting a holistic perspective towards ecosystem
protection, since ecosystems are similar to organisms in that their well-being requires preservation of the whole, not merely a
few of the attractive parts (e.g., charismatic megafauna).
The second difficulty is that a macroorganism species growing in different geographic locations may well have quite different
symbiont microorganism populations. This possibility complicates strategies for both marine bioprospecting and ecosystem
protection. A third difficulty is that culturing symbiont microorganisms apart from their hosts has been very difficult, and even
when such cultures do succeed, they often do not produce the desired active natural products. Pharmaceutical screening is
consequently more complicated, because it is necessary to analyze the microbial genome, identify and extract the likely
sequences used to create the secondary metabolites, and insert those sequences into a culturable microorganism. If a sufficient
amount of the secondary metabolite has been isolated from natural sources, another alternative would be to identify the
chemical structure for synthesis. Unfortunately, synthesizing these complex secondary metabolites is unlikely to be feasible
either with respect to time or cost early in the screening process.
The macroorganisms that have been found with potentially useful secondary metabolites (whether originating from the
macroorganism or its symbiotic microorganisms) are predominantly from the phylum Porifera (sponges) (Figure 5-2 and
Table 5-7). This observation results from an examination of the U.S. National Cancer Institute's Developmental Therapeutics
Program and its Natural Product Extract Cancer Screening Database , which appears to have been last updated in 2003. The
database contains the results of 236,335 cancer screens for 4,335 marine natural products, 472 from marine plants and 3,863
from marine invertebrates. The predominance of sponges most likely results from several factors. Sponges tend to be fairly
prominent and discrete, making them visible and easy to collect. Also, from the earliest marine biochemical research, sponges
had developed a reputation for being excellent sources of biologically active chemicals. This may have led researchers to
preferentially collect sponges for evaluation. Sponges appear to be a good starting point for investigating symbiotic
microorganisms, because those microorganisms can account for greater than 60% of a sponge's wet weight (Bowling et al.
2007).
Simmons and Gerwick (2008) used metagenomic analysis to clarify the sources of 20 marine anticancer products (marked in
Table 5-2 with a f) currently in clinical trials (Figure 5-3). The current view is that only 20% of these anticancer products (or
the natural templates on which they are based) are produced by macroorganisms and that fully 80% are produced by symbiotic
microorganisms. Of course, as noted above, if the macroorganisms are not present to host the microorganisms, the
microorganisms will not produce the biologically active secondary metabolites.
A macroorganism is an organism that can be seen with the naked eye.
Cf
available at http://dtp.nci.nih.gov/docs/cancer/natural_products/natural_products_data.html (accessed 2010.08.19)
-------
Figure 5-2. Marine phyla contributing secondary Table 5-7. Percentage of secondary metabolites with
metabolites with pharmaceutical potential pharmaceutical potential from marine phyla
(percentages estimated from Figure 1 in Hunt &
Vincent [2006])
(adapted from Hunt & Vincent [2006])
Nemertea
& Annelida
Arthropoda
Phylum
Porifera
Cnidaria
Tunicata
Echinodermata
Heterokontophyta
Mollusca
Chlorophyta
Bryozoa
Arthropoda
Nemertea
& Annelida
PhylumMembers
sponges
corals, jellyfish, sea anemones
tunicates (ascidians)
starfish, sea urchins
brown algae, seaweeds
nudibranchs, snails, squid
green algae
colonial filter feeders
barnacles, crabs, shrimp
worms
Metabolites
64.7%
10.1%
6.0%
4.7%
3.4%
3.1%
1.8%
1.7%
0.3%
0.2%
Figure 5-3. Sources of 20 marine anticancer drugs in clinical trials or recently in clinical trials (see Table 5-2):
reported (left) versus predicted based on subsequent metagenomic analysis (right)
(adapted from Simmons & Gerwick [2008])
5.3 Measuring and valuing the service
The provision of natural products by marine ecosystems has not been assessed in the same manner as the other ecosystem
services discussed in this document, probably because this ecosystem service, being of a distinctly different nature, does not
lend itself to commonly employed assessment and valuation methods. For example, there is no easy or straightforward method
for estimating either the number of natural products that could be developed or their value. Developing an ecosystem service
assessment, particularly for pharmaceutical products, would be greatly impeded by the reticence of companies to discuss their
methods, the status of their investigations and clinical trials, or even the specific products being investigated. Given these
significant hurdles, it is not especially surprising that there appear to be no reported estimates of the value, or potential value, of
marine natural products. There are rare instances where a product's sales may be given for a single year, but such examples
-------
tend to be treated (even by authors) as throwaway factoids, rather than reliable data. Creating a list of potential products, such
as Table 5-2, requires a wide-ranging review of many journal articles, reports, and web pages, because the information has not
been compiled, and the lists that are available are inevitably incomplete and quickly outdated by new developments.
5.3.1 Specific examples
There are few well-documented market values for marine pharmaceutical products. Part of the problem is that there are not very
many marine pharmaceutical products currently in commercial production. Another aspect of the problem is that the
pharmaceutical industry tends not to make sales and profit figures available that are disaggregated by year or product. When
figures are made available, they tend to be provided in ways that make comparisons difficult, if not impossible. In 2005 in the
British MedicalJournal, the following estimates were given:
The annual profits from a sea sponge compound used to treat herpes, for example, are between $50m and $100m (£27m
and £55m; €41m and €81m), and cancer fighting agents derived from marine organisms are worth $lbn (Cole 2005).
The herpes drug is almost certainly acyclovir (aka Zovirax®) (see Table 5-2 and Appendix 5-A [listed as acycloguanosine in the
Anti-infectives section]).
Another approach is to consider the value of relatively new drugs not of marine origin to gain some sense of the market
potential for new marine drugs (Table 5-8).
Table 5-8. Market value of pharmaceutical products
Product
Avastin®
Herceptin®
Prezista®
all products
T • •. IB
Lipitor
Retrovir® (AZT)
^ • (8
Zovirax
Market Segment
cancer
cancer
HIV
antibiotics
cholesterol-lowering
HIV
antiviral (herpes)
Sales
$2.7billion/yr
$ 1.3 billion /yr
$25m (est. for 2006)
$1 81m (est. for 2007)
$5 billion
$13.6 billion (2006)
$23m (2005)
~$237m (2006)
Source: UN (2007)
5.3.2 Possible approach
One approach that could be used to approximate the potential value of marine natural products in the pharmaceutical industry
would be to rely on assumed values for the following parameters:
1. the number of marine species
for megafauna, the number of species is likely to be about 250,000 (as discussed above), but it will be more
difficult to estimate the number of species of microorganisms (the estimates discussed above come with great
uncertainly);
the probability that any given species could be the source of a marketable pharmaceutical product
estimates of this probability vary for terrestrial species, but it appears that the per species probability may be
considerably higher for marine species;
the expected value of a product's revenue
revenue estimates have been difficult to find (particularly revenue projections), but in the past decade, there
has been more independent research in this area;
the expected cost of a product's development
research costs have always been difficult to estimate, primarily because pharmaceutical companies consider
them to be competitively important and so rarely release them except in the most aggregated forms. However,
research on these costs has resulted in numerous journal articles in the past decade, particularly by DiMasi
and colleagues at the Tufts Center for the Study of Drug Development (e.g., DiMasi 2001a; DiMasi 2001b;
Grabowski et al. 2002; DiMasi et al. 2003; Adams & Brantner 2006; DiMasi & Grabowski 2007; DiMasi
etal. 2010); and,
the expected value of a product's nonmarket benefits
the nonmarket benefits are usually the most difficult to estimate and the most difficult on which to reach
consensus. These benefits will vary depending on: (a) the disease(s) a drug will be used to treat; (b) the
treatment regime (short-term vs. long-term consumption); (c) the number of expected users; and, (d) the age
distribution of the user population. Several difficult choices have to be made in arriving at an estimate,
2.
3.
4.
5.
-------
including the value of human well-being (including quality of life), the value of life extensions, the value of
palliative treatments, the value of a life saved, and whether the value of life depends on the age of the person
saved.
This approach has been used to estimate the potential pharmaceutical value of terrestrial plant biodiversity (Principe 1987;
Principe 1996). However, such global values are not very useful for assessments and management of specific marine locales
and do not provide the marginal values necessary for a proper economic assessment (Simpson et al. 1996; Simpson & Craft
1996; Craft & Simpson 2001).
5.4 Reflections
In considering the coral reef ecosystem and which of its attributes contribute to the creation of the ecosystem services provided
by natural products, it seems that one must inevitably conclude that these services are not so easily categorized. This results
from the services being exclusively created by the ecosystem as a whole; a single attribute is not responsible for the provision
of the service. The services provided by the natural products also do not lend themselves to a disaggregated perspective, as is
possible, for example, in shoreline protection. Consequently, the table of final ecosystem services and supporting features
(Table 5-9) is a brief one.
Marine natural products possess several characteristics that make them compellingly attractive for pharmaceutical purposes.
First, the structural complexity of marine secondary metabolites is far greater than from terrestrial sources. Second, marine-
derived compounds seem to possess a far higher probability of success than the one in ten thousand expected from traditional
sources of potential pharmaceuticals (Principe 1987; Principe 1996). Third, marine-derived compounds have demonstrated
biological activity against a wide variety of diseases, afflictions, and pathogens. Fourth, only a small percentage of marine
secondary metabolites have been investigated for pharmacological use to date, so there remains an extremely large pool of
marine biochemicals to investigate for pharmaceutical use. The utility for humans from marine-based pharmaceuticals is
potentially so large that its economic value could surpass that of all other coral reef benefits combined. Consequently, the
estimation of this utility will be a vital, if not determinative, element of analyses conducted to support policy decisions that
affect the health and integrity of coral reefs.
Even once one gets past the stage of gawping, touristic wonder when viewing the complex structures and strategies created by
the denizens of marine ecosystems, the tremendous potential of marine natural products to benefit humans remains mind-
numbingly large. Unfortunately, what also remains is the discouraging possibility that the degradation or destruction of coral
reef ecosystems will result in these being mind-numbingly large foregone benefits. This outcome could be forestalled if the
potential benefits of marine natural products could be better characterized and quantified, and this should be one of the goals of
the ESRP research program.
Table 5-9. Final ecosystem services and supporting features for natural products
Ecosystem Service(s)
Final (FES)
Intermediate
Natural Features
Social Values
Complementary
Goods & Services
Ecosystem-
Derived
Benefits
Potential
Indicators of Final
Ecosystem
Service(s)
Pharmaceuticals from natural products
Marketable natural
product or a template
that results in a
marketable product
Unique biologically
active secondary
metabolite
Shallow, marine
biodiverse,
species-dense
ecosystem
Desirability of
good health and
well-being
Pharmaceutical Increased
research programs for revenues from
both field collection pharmaceuticals;
and laboratory increased health
analysis and well-being
Species density,
biological
integrity, sponge
diversity, rare
species
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Appendix 5-A
Marine pharmaceutical products
This table contains a small subset of the marine biochemicals possessing biological activity that have been investigated as
potential pharmaceutical products or as molecular probes for exploring biochemical pathways and reactions. The biochemicals
are grouped into the following categories:
• Antiasthmatic drugs
• Anticancer drugs
• Antidiabetic drugs
• Anti-infective drugs
• Cardiac & circulatory system drugs; Anti-angiogenesis drugs
• Immunological drugs
• Molecular probes
• Neurological drugs
• Templates
Some of the chemicals have duplicate entries if they have multiple uses.
Analgesic drugs
Chemical
Source
Uses & Status
Structure
Citations
mollusk (cone snail)
(Conus geographus)
Indo-Pacific
conantokin G
analgesic & anti-seizure;
17-amino-acid competitive
antagonist of N-methyl-D-
aspartate (NMDA) receptors;
Phase I trials discontinued
precursor amino acid sequence
shown at (128, Figure 2, p. 275)
1,2,18
photo: 3
a-conotoxin
Vcl.l(ACVl)
mollusk (cone snail)
(Conus victories)
Australia, Japan
a-conotoxins inhibit the
postsynaptic acetylcholine
receptor; preclinical studies
Asp—Pro
Ser *
Cys--'
Gly-Cys
•Tyr
.-• :
N— Cys—lie—Glu-Pio
xln|ACV1)
(146)
18,146
X-conotoxin
MR1A/B
mollusk (cone snail)
(Conus sp.)
Indo-Pacific
X-conotoxins selectively inhibit
the activity of neuronal
noradrenaline transporter;
preclinical studies
4,18
mollusk (cone snail)
(Conus catus)
Indo-Pacific
oconotoxin
CVID (AM-336)
mollusk (cone snail)
(Conus geographus)
Indo-Pacific
contulakin G
(CGX-1160)
oconotoxins inhibit voltage-
activated entry of calcium into
the presynaptic membrane,
inhibiting the release of
acetylcholine; preclinical
studies
Cy*—Lys Sef—ly»—Gty—Al»—Ly»—Cy»—Sw—Ly*—
Gly Sef Cys Sw Gty Thr Cys CyB Aap—Tyr—M«t
Trw Val Gty—Arg-
w conoto.,n (AM-366)
(146)
a 16-amino acid peptide with
an O-glycosylated threonine
residue; a neurotensin
agonist; Phase I trials
18,146
photo: 5
6,18
photo: 3
Thr 10-contulakinG
(modified toxin)
mollusk (cone snail)
(Conus sp.)
Indo-Pacific
preclinical studies
7,8,18
-------
Analgesic drugs
Chemical
Source
Uses & Status
Structure
Citations
gorgonian (sea whip)
(Pseudopterogorgia elisabethce)
Caribbean
pseudopterosins
(including
methopterosin, a
derivative)
arthritis, anti-inflammation, &
analgesic; affect the arachidonic
acid cascade; inhibit synthesis of
I eicosanoids, (locally functioning
hormone-like substances) in specific
white blood cells; pharmaco-
logically distinct from other
SAIDs & their MOA seems novel;
Technical; sold as a cosmetic anti-
wrinkle cream by Estee Lauder
under the name
9,10,24,25
photo: 11
pseudopterosin A
NHa
spongosme
(2-
metoxyadenosine)
sponge
(Tectitethya crypto)
(synonym: Cryptotethya crypto)
Caribbean
analgesic; neuropathic &
inflammatory pain
146
spongosine
(146)
mollusk (cone snail)
(top '.Conus geographus,
bottom: Conus magus')
Indo-Pacific
H,N— Cys Lys Gly L<
ziconotide
(Prialt®)
inhibits N-type voltage-
dependent calcium channels to
short-circuit neurotransmitter
release in nerves that transmit
pain signals; the precisely
targeted MOA effectively
blocks pain while still allowing
the rest of the nervous system
to function properly; the effect
of (D-conotoxin M VII A is 100
to 1000 times that of morphine;
FDA approval in December
2004; licensed to Warner
Lambert
r
Ala Gly
H,N—Cys--- Cy«
' rs—Asp—Tyr
Lys Gly—Se/—flee—Arg —Leu—Mel
Gly —Ser Atg—Cys
(146)
V7
v>""i*K- - - -,
' _
'
3xr
.
12,13,14,
18,102,106,
146
photos:
3 (top)
15 (bottom)
Antiasthmatic drugs
Chemical Source Uses & Status Structure Citations
antihistamine & antiasthmatic; \
IPL-576,092
(HMR-4011A)
IPL-550,260
IPL-5 12,602
(contignasterol
derivatives)
contignasterol
sponge
(Neopetrosia contignatd)
(synonym: Petrosia contignata)
Indonesia
sponge
(Neopetrosia contignatd)
(synonym: Petrosia contignata)
Indonesia
IPL-576,092 completed Phase
II trials successfully (by
Inflazyme Pharma);
IPL-5 12,602 in Phase I trials
(by Avantis); IPL-5 12,602 in
Phase II trials (by Avantis)
antihistamine & antiasthmatic
rH
rfr
HO^-^OH
OH IPL-576,092 (18)
M C
/
--^O-"
~\^I.»
~JTL*
^j^fVTy
^vicxr
(16)
16,18
16,18
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
aplidine
(total synthetic of
dehydrodidemnin B)
(Aplidin®)
tunicate
(Aplidium albicans)
Mediterranean
anticancer; differs chemically
from didemnin B and the other
didemnins only in the structure
of its side chain; MOA is
unclear; a multifactorial
apoptosis inducer;
Phase II trials underway for
various cancers; PharmaMar
17,18
Ara-A
(vidarabine,
Vira-A®)
(derived from
spongouridine &
spongothymidine)
sponge
(Tectitethya crypto)
(synonym: Cryptotethya crypto)
Caribbean
antiviral & anticancer;
clinical use
18,19,102
0-,
HO/
Ara-C
(Cytosar-U*
DepoCyt0, ^ sponge
Tarabme PFS , (Tectitethya crypto)
cytarabine, 1-p-D- (synonym: Cryptotethya crypto)
arabinosylcytosine) Caribbean
(derived from
spongouridine &
spongothymidine)
antiviral & anticancer;
clinical use
18,19,
102,103
(102)
arenastatin A
(cryptophycins)
sponge
(Dysidea arenaria)
Palau Islands
anticancer; tubulin interactive
agent; Phase I trials; synthetic
derivative licensed to Lilly by
Univ. of Hawaii; withdrawn in
2002
18,20
ascididemnin
tunicate
(Leptoclinides sp.)
Central Pacific
anticancer; reductive DNA-
cleaving agents; preclinical
studies
18,21
azaspiracid-1
anticancer; shows toxicity to
lymphocytes & neuroblastoma
cells
sponge
(Jaspis sp.)
Fiji
bengamide
derivative;
LAF-389
anticancer & antihelminthic;
methionine aminopeptidase
(Met-API) inhibitor; licensed
to Novartis; withdrawn from
Phase I trials in 2002; LAF-389
is a synthetic bengamide B
derivative; Phase I trial may
have been suspended in 2006
18,22
photo: 23
bistratene
tunicate
anticancer; induces cell-cycle
arrest in G0/Gi & G2/M
128
bryozoan
(Bugula neritina [photo])
(chemical possibly from
commensal bacterium
\Endobugula sertuld\)
worldwide
bryostatin 1
anticancer; inhibits leukemia
via immunostimulation &
binding to the receptor, protein
kinease C (PKC), displacing
tumor promoting phorbol esters
that bind to the same place;
seems to enhance other drugs
but not effective by itself;
Phase II trials; in combination
therapy trials in 2004; licensed
by Arizona State Univ to GPC
Biotech., which stopped
development in 2003
(21:
18,24,25,26,
27,102
photo: 28
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
calyculin A
&
calyculin C
sponge
(Discodermia calyx)
anticancer; strong
serine/threonine protein
phosphatase inhibitors
OH OH OMo
(29)
cematodin
(synthetic derivative
of dolastatin 15)
(LU-103793)
sea hare
(Dolabella auricularia [photo])
(chemical possibly from
commensal cyanobacteria
[Symploca hydnoides &
Lyngbya majuscula])
Indian Ocean
anticancer; Phase II trials for
malignant melanoma, meta-
static breast cancer & non-
^mall-cell lung cancer; Phase II
trials for melanoma, breast, &
NSCLC; appears to stabilize
melanoma & breast cancers;
increase in QoL for NSCLS
18,44
photo: 31
chinikomycin A
&
chinikomycin B
chlorine-containing
manumycin
derivatives)
sediment actinobacterium
(Streptomyces sp.)
China
anticancer; antitumor activity
against several cancer lines
O diMkomychi • (32)
coscinosulfate
sponge
anticancer; cell cycle regulation
curacin A
cyanobacterium
bya majuscula)
Caribbean
anticancer; antimitotic activity;
tubulin interactive compound;
preclinical; synthetic
derivatives with better
solubility being evaluated
18,24,25,
33,34
(34)
diazepinomicin
(ECO-4601)
actinobacterium
(Micromonospora sp.)
antibiotic & anticancer;
Phase I trials (antibiotic &
anticancer)
diazepinomicin
OH ECO-4601
diazonamide
ascidian
(Diazona angulatd)
Phillipines
anticancer; inhibits microtubule
assembly, arresting the process
of cell division; preclinical
studies; synthesized & new
structure elucidated
dictyodendrins
sponge
(Dictyodendrilla verongiformis)
Japan
anticancer; telomerase
inhibitors; has shown 100%
inhibition; preclinical studies
06 J •',
dictyodendrin A (18)
dictyostatin
anticancer; inhibits the growth
of human cancer cells; active
against certain Taxol-resistant
tumors; MOA appears to be
prevention of the breakdown of
tubulin during mitosis (like
Taxol); preclinical studies
38
photo: 126
OH OH
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
didemnin B
tunicate
(Trididemium solidum)
Caribbean
anticancer & antiviral interrupts
protein synthesis in target cells by
binding noncompetitively to
palmitoyl protein thioesterase;
cytotoxic for lymphomas, some
leukemias & melanomas;
antiviral for herpes simplex and
several others; Phase II trials
showed significant toxicity at
efficacious doses; dropped in
middle 1990s
18,24,
39,40,41
(41)
discodermolide
deep-water sponge
(Discodermia dissoluta)
Caribbean
anticancer & immunosuppressive;
tubulin polymer stabilizer (like
;axol); essentially arresting cells at <
specific stage in the cell cycle and
halting cell division; Phase I trials;
licensed to Novartis by Woods
Hole; may be used in combination
with Taxol
-
24,25,42,43
(43)
sea hare
(Dolabella auricularid)
(chemical possibly from
commensal cyanobacteria
[Symploca hydnoides &
Lyngbya majuscula])
Indian Ocean
dolastatin 10
anticancer; mitotic inhibitor;
interferes with tubulin formation &
thereby disrupt cell division by
mitosis; binds to tubulin at the
vinca/peptide region, the target for
several structurally complex natural
products, including hemiasterlin;
cytotoxic for B-16 and LOX
melanomas; no positive effects in
Phase II trials, but may find use in
combination drug therapy with
vincas or bryostatin; many
derivatives made synthetically (see
next entries)
18,24,25,
44,45,46
photo: 31
tunicate (Caribbean sea
squirt, a mangrove ascidian)
(Ecteinascidia turbinata)
Caribbean
ecteinascidin 743
(trabectedin,
ET-743,
Yondelis®)
anticancer; cytotoxic for several
types of cancer; binds to target cell
DNA & inhibits cell division,
leading to apoptosis; induces
apoptosis only during active gene
transcription, which is much more
frequent in cancer cells; keeps
tumors from becoming resistant to
chemotherapy by interfering with
the gene that produces
P-glycoprotein, a membrane protein
.that enables drug resistance; Phase I
trials showed effectiveness against
advanced-stage breast, colon,
ovarian and lung cancers,
melanoma, mesothelioma and
several types of sarcoma; Phase
II/III trials in 2003; licensed by
Ortho Biotech (J&J); partial
synthesis from microbial
metabolite; approved by EC for soft
tissue sarcoma
18,24,25,
44,47,48,49,
102
photo: 50
(102)
eleutherobin
(related to
sarcodicytins)
alcyonarian
(Eleutherobia sp.)
gorgonian (Eunicella stricta)
coral (Erythropodium
caribceorum [photo])
W. Australia,
Caribbean (E. carib&o,
*•*>»'
anticancer; mimics pacilitaxel's
synthesized and derivative
structures created
18,51,102
photo: 52
-------
Anticancer drugs
Chemical Source
elisidepsin
(Irvalec®,
PM02734)
eribulin
&
eribulin mesylate
(E7389,
Halaven®)
farnesylhydro-
quinone
girolline
halenaquinone
halichondrin B
hectochlorin
hemiasterlin
(hemiasterlin analog
E-7974)
hermitamide A
&
hermitamide B
icadamides
sacoglossan sea slug
(Elysia rufescens) that grazed
on green macroalgse
(Bryopsis pennata)
Pacific
sponge
(Halichondria okadai,
Axinella sp., Phakellia carteri, &
Lissodendoryx sp.)
Japan, W. Pacific,
Eastern Indian Ocean, &
New Zealand
fungus
sponge
(Pseudaxinyssa c^ntharelld)
(synonym: Cymbastela
cantharella)
New Caladonia
sponge
Uses & Status Structure Citations
Synthetic analog of kahalalide
F created to insure sufficient
supply; licensed to PharmaMar
by Univ. of Hawaii;
Phase II trials
Eisai's synthetic halichondrin B
derivative; breast, prostate, &
non-small cell lung cancer
(NSCLC) cancers; Phase II/III
trials; FDA approved for late-
stage breast cancer in 2010
anticancer & antimalarial
anticancer; inhibits protein
synthesis at termination the
process rather than at the
initiation or chain elongation
steps like other known
inhibitors; Phase I trials
discontinued due to
hypertension
anticancer; induces apoptosis
sponge
(Halichondria okadai, anticancer; binds tubilin at a
Axinella sp., Phakellia carteri, & site close to the so-called vinca
Lissodendoryx sp.) s;te and altered tubulin
Japan, W. Pacific, depolymerization; in clinical
Eastern Indian Ocean, & trials; see eribulin
New Zealand
bacterium
sponge
(Hemii^^iell.: minor,
Auletta sp., Siphonochalina sp.,
Cymbastela sp.)
South Africa
cyanobacterium
(Lyngbj\; n;xula)
Papua New Guinea
sponge
(Leiosella sp.)
Phillipines
anticancer; inhibits cell growth;
induces actin polymerization
anticancer; cytotoxic,
anti-tubulin; mitotic inhibition
occurs through binding to tubulin at
the vinca/peptide region in a manner
similar to dolastatin and the vinca
alkaloids; Phase II trials; licenses to
Wyeth by Univ. of British
Columbia; a closely related
chemical, E7974, from Eisai is
currently in Phase I trials
anticancer;
anticancer, antiviral, &
immunostimulant
^vA^
.;,., C v^Cb
«, r*
Elisidepsin.
>N PM02734 (53)
Wtq
HO-/ 0 -: ° JV*0 M
Jr o-y p>v!cH
MlOO ^ U* t2?*'H
Eritxilm (mesylale sail)
(E7M9) (54)
OH
.^i Famesytiydroquinonc
^y^Y^vXV^~^^V'
OH (128)
HO ,
* /T^NH
»HX,
(55)
llal«n»numon» (128)
^Y'CpCtf? i>
*** * ' i'M'jf'"f~^r
(57)
^
s-Sr°<>^c1
Hectochtoiio O (128)
R,
K Hemimlnlln: R, R CH
Hgmimurlln A: R, = H: R, = CH,
Htmlaiterlin B: R. B H
E-?«4 /S (both 44)
?H 0 5QOH
•v OH ? 1^*
»sXfVt' HiNYNH
X^XX^ Icadamide B '-"
(106)
53
18,44,54
128
18,55
128
24,25,56,57
128
18,44,58
114
59,106
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
sacoglossan sea slug (Elysia
rufescens) that grazed on
green macroalgse (Bryopsis
pennatd)
Hawaii
kahalalide F
(revised structure)
(see elisidepsin,
above)
anticancer; disrupts lysosome [def]
membranes within certain target
cells, thereby initiating apoptosis
(programmed cell death); inhibits
gene expression related to DNA
replication & cell proliferation;
Phase II trials for NSCLC,
melanoma & androgen-independent
I prostate cancer; also being studied
for use on an drog en-resistant
prostate cancer, liver cancer, &
advanced solid tumors; licensed to
PharmaMar by U. Hawaii
18,44,60,61
photo: 60
KRN-7000
(a-GalCer,
a-galactosyl-
ceramide)
(agelasphin
derivative)
sponge
(Agf-'lay mauntiami)
Red Sea, Indian Ocean
anticancer; stimulates
lymphocytic proliferation under
certain conditions; appears to
stimulate the production of
natural killer T (NKT) cells in
the body; Phase I trial showed
effects with patients having
high levels of NKT cells. Phase
II trial ongoing
18.62
(62)
deep-water sponge
(Forcepia sp.)
Gulf of Mexico
lasonolides
anticancer & antifungal; kills
cancer cells in a different way
than most other cancer drugs;
the exact mode of action is not
yet fully understood, and is an
area of active research;
preclinical studies
latrunculins
sponge
(Latrunculia magnified}
Red Sea
cytotoxic actm-active agent;
disrupts actin polymerization,
microfilament organization, etc; cell
shape, cytokinesis, and
microfilament-mediated processes
such as fertilization and early
development are altered; preclinical;
early tests show low in vivo
activity; may require novel drug
delivery strategies; also used as a
probe for studying the role of actin
in maintaining cell shape
63
photo: 64
sponge
laulimalide (Cacospongia mycofijiemis)
Pacific
anticancer; microtubule
stabilizer; activity profiles are
clearly different than other
microtubule-binding agents
such as paclitaxel; preclinical
studies
18,66
(66)
commensal bacterium found
on tunicate (Lissodinum sp.)
Indo-West Pacific
'hose. LiKsocliniwi oaieJiam}
lissoclinamide 7
anticancer
67,106
photo: 68
-------
Anticancer drugs
Chemical
lomaiviticin A
makaluvamines
manzamine A
&
Ircinol A
(its likely biogenic
precursor, which
acks the p-carboline
moiety)
mechercharmycin A
micropeptins
namenamicin
Neovastat
OE-941)
(a derivative of
shark cartilage
extract; not a
Source
commensal bacterium
(Micromonospora
lomaivitiensis) found on
tunicate
Fiji
sponge
(Zyzzya fuliginosa)
Indo-West Pacific
sponges (Halidona sp.,
Pachypellina sp.) with
commensal bacterium
(Micromonospora sp.)
Uses & Status
anticancer; potent DNA
damaging activity
anticancer; cytotoxic through
inhibition of DNA
topoisomerase II
antitumor, antimalarial, anti-
Indo-Pacific infective, antituberculosis,
(photo: Halidona sarai) antitoxoplasmosis, &
^TT^
•
sediment actinobacterium
antineurogenic inflammation;
has shown activity against
malaria, tuberculosis, HIV, and
other inflammatory diseases;
preclinical studies
anticancer; active against lung
(Thermoactinomyces sp.) cancer & leukemia cell lines; a
Palau
bacterium
commensal bacterium found
on tunicate (Polysyncraton
Hthostrotuni)
Australia, New Zealand
, ,
(photo: Carcharhinus
patent claims cytotoxicity
anticancer; inhibition of trypsin
& chymotrypsin
antitumor & antibiotic; the
enediyne moiety is very
reactive with DNA, making
these chemicals extremely
cytotoxic for all cells & among
the most potent antitumor
agents known
anticancer; inhibits the binding of
Vascular Endothehal Growth Factor
(VEGF) to its receptors; normally,
specific mono- : | VEGF binds to target endothehal
molecular com- 1 Blreceptors & directs the profusion of
pound, JE-941 is a 1 new capillaries to nourish the
defined standardizedPJ 1 tumor; by blocking the receptor
liquid extract com- P sites, AE-941 preempts the
prising the <5 00 1 MIformation of the new blood supply;
kDa (kilodaltons, a 1 Phase II & III trials for renal
unit of mass)
fraction from shark
cartilage)
carcinoma and NSCLC
Structure Citations
jjj£^
C^XI^S5
r^7jj j
(no,!/ lonulviliclo A . .
/ — v /*>>*,
/ — { ^*\
v-««< \
\>/J~-^X^—i>^w
y
\ ''i>S
H CH2OH
V*"^ T?
Ircinol A \ /
(117) (103)
f~\
,/"*r!'N'Sr0x
<3 O
(ji x °
r a yS®
(32)
-xjffi 3[ :&
S3SSST j|^
(128)
"' -Ml
™"O O
"\-\&o 5<'.u';"^ ' ,LX '»»•
V^ y.ai^o7!' •
" «J
'"' '"• (72)
a mixture
69
70,114
103,117
photo: 71
32
128
72
18
poo.
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
NVP-LAQ824
(dacinostat)
(synthetic using
structures from
psammaplin,
trichostatin, &
trapoxin)
sponge
(Psammaplysilla
[also found in a 2-sponge
association ofPoecillastra sp. &
Jaspis sp.])
Indo-West Pacific
antibiotic, anti-tumor, DNA
methyltransferase inhibitor;
extremely potent histone
deacetylase (HDAC) inhibitor;
Phase I trial for hematologic
malignancies
\.J
(18)
; 8.74
sponges
(Theonella sp. and
; 'r< a':h\•< -l^Jii^ L; visi/in> ///^r <
Okinawa, Australia
3 ho to: Theonella cylindr,
onnamide A
onnamide F
onnamide A
2J&
' I • I
-^kL- ^VJ^^^^XXX^-^Y"" N-^^^^"-^*
^•"V
anticancer, antifungal, &
anthelmintic
(75)
panobinostat
(LBH-589,
Faridak®)
sponge
(Psammaplysilla spp.)
Indo-West Pacific
anticancer; synthetic analog of
psammaplin; with Novartis;
Phase III
onnamide F
X
75,76,
114,128
photo: 77
(76)
.8.
panobinostat
78
(78)
commensal bacterium found
on tunicate (Lissoclinum
patella)
Indo-West Pacific
Patellamide A
patellamides
pectenotoxin-6
anticancer; cytotoxic;
patellamides B, C, & D appear
to reverse multidrug resistance
r^tr1
\—l. N
;N
I
8
t
HN
N
79,80,106
photo: 68
anticancer; induces F-actin
depolymerization
128
Pectenotoxm-6
(128)
peloruside A
sponge
(A lycale hentscheli)
New Zealand
anticancer; appears to bind
tubulin and arrests target cell
development at the G2-M
transition stage of the cell
cycle, triggering apoptosis (cell
suicide) before mitosis can
begin; preclinical studies
18,81
OMe
fungus (Phoma sp.) found
on shell of crab
(Chinoecetes opilio)
Japan
phomactins
anticancer; platelet activating
factor (PAF) antagonists
83
photo: 84
phomactin A
plinabulin
(NPI-2358)
fungus
(Aspergillus sp.)
anticancer; synthetic analog of
marizomib; selective vascular
disrupting agent (VDA);
Phase I/II
Plinabulin NPI-2358 f^\
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
psymberin
(irciniastatin)
sponges
(Psammocinia sp. &
Ircinia ramosd)
Indo-West Pacific
anticancer; showed extremely
potent toxicity toward several
cancer cell lines (>104 more
potent than usually observed))
sponge
(Haliclona sp.)
Indo-Pacific
(photo: Haliclona sarai)
salicylihalimides
anticancer & anti-osteoporosis; first
marine Vo-ATPase inhibitor (Vo-
ATPases are eukaryotic enzymes
whose principal role is to pump
hydrogen ions across cell vacuolar
membranes); may mediate bone
resorption; preclinical studies
SalicylihalamideA
18,86,87
photo: 71
saliniketal A
saliniketal B
actinobacterium
(Salinispora arenicola)
worldwide
chemopreventive
salinosporamide A
(NPI-0052,
marizomib)
actinobacterium
(Salinispora tropica)
tropics
antibiotic & cytotoxin; very potent
proteasome inhibitor; A & B
nhibited colon cancer cells in vitro;
A was extremely potent against
NSCLC, CNS, & breast cancer
lines; in Phase I trials with Nereus
Pharma
Salinosporamide A
(44)
fungus (Fusarium sp.) found
on seagrass (Halodule wrightii)
western tropical Atlantic
& Gulf of Mexico
sansalvamide
anticancer; selective
cytotoxicity towards colon &
melanoma cell lines
88,114
photo: 89
sarcodicytin
(related to
eleutherobins &
eleuthosides)
sculezonone A
&
sculezonone B
alcyonarians (soft corals)
(Sarcodictyon roseum [photo],
Eleutherobia aurea, &
Bellonella albiflora)
Mediterranean
anticancer; tubulin interactive
agent; synthetic combinatorial
research using base structures
of sarcodicytins &
eleutherobins; preclinical
testing of derivatives
Me
bacterium
anticancer; inhibits DNA
polymerase
18,86,102
photo: 90
(102)
Scui«zonone A R«M
SculMonom B B-OH (128)
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
soblidotin
(auristatin PE,
TZT-1027)
(synthetic analog of
dolastatin)
sea hare
(Dolabella auricularid)
(chemical possibly from
commensal cyanobacteria
[Symploca hydnoides &
Lyngbya majuscula])
Indian Ocean
anticancer; exhibits potent
antivascular effects in addition
to antitublin activity; Phase III
trials; Phase II trials showed no
positive results when used
alone, but it appears to be
effective in combination
therapy with vinca alkaloids
and bryostatin
18,44
photo: 31
spisulosine
(ES-285)
squalamine
(opthalamic
formulations are
called Evizon™)
Arctic surf clam
(Spisula polynymd)
North Atlantic,
North Pacific, Arctic Seas,
Japan
anticancer; Rho-GTP inhibitor;
appears to alter cell morphology:
treated cells appear to lose actin
stress fibers, (bundles of actin
filaments that appear & disappear in
response to mechanical stimuli);
Phase I trials
18,91
NH,-CI
spiny dogfish
(Squoiy* acanlhjvs)
Northwest Atlantic
(91)
anticancer, antiangiogenic, &
antibiotic; broad spectrum
antibiotic; also shows anti-
angiogenic activity and may be
useful to treat wet-form age-related
macular degeneration; Phase II
trials for nonresponding solid
tumors as part of combination
therapy and for advanced ovarian
5 cancer as primary treatment; Phase
III trials for wet macular
degeneration show significant
activity
18,44,
92,103
photo: 93
(92;
stolonoxides
tunicate
anticancer; mitochondrial
respiratory chain inhibition
SwkxtoxKteA FUM
StotoooxidB A mathyl «t«f fl*CH3 (128)
swinholide A
sponge
anticancer; facilitates outflow
in eye
128
(128)
synthatodin
(ILX651,
tasidotin)
(3rd gen. derivative
of dolastatin 15)
sea hare
(Dolabella auricularid}
(chemical possibly from
commensal cyanobacteria
[Symploca hydnoides &
Indian Ocean
Taltobulin
(HTI-286)
(hemiasterlin
analog)
anticancer; orally-active 3rd
generation analog; Phase I/II
trials for melanoma, breast,
NSCLC; licensed by Ilex from
BASF Pharma
o o o
(18)
18,44,46
photo: 31
Phase I/II trials
(44)
-------
Anticancer drugs
Chemical
Source
Uses & Status
Structure
Citations
theopederins
sponge
(Theonella ^ixhoe')
Indo-Pacific
anticancer
106
thiocoraline
actinobacterium anticancer; DNA polymerase a
(Micromonospora marina) inhibitor; preclinical studies
18,94
commensal bacterium found
on tunicate (Lissoclinum
patella)
Indo-West Pacific
Trunkamide A =
trunkamide
anticancer; has specific &
unusual activity against the
multidrug-resistant UO3 1 renal
cell line
80,106
photo: 68
variolins
Antarctic sponge
(Kirkpatrickia variolosa)
Antarctic
anticancer; Cdk inhibitors;
preclinical studies
18
vitilevuamide
tunicates
(Didemnum cucliferum &
Polysyncraton lithostn 'turn
[photo])
Australia, New Zealand
anticancer; inhibits tubulin
polymerization & can arrest the
cell cycle in the G2/M phase;
tubulin binding & inhibition
occurs at a different site on the
tubulin molecule than used by
dolastatin 10, colchicine, & the
vinca alkaloids; preclinical
studies
18,95
yessotoxin
anticancer; lymphocyte
homeostasis modulation
128
Zalypsis*
(PM1004)
nudibranch
(Jonnma /^m/:!o^}
Western Indian Ocean,
Red Sea
OCH3
53
photo: 96
(53)
-------
Antidiabetic drugs
Chemical
Source
Uses & Status
Structure
Citations
insulin
trodusquemine
(MSI-1436)
European spotted dogfish
(Scyliorhinus caniculd) &,
hammerhead shark
(Sphyrna lewini [photo])
North Atlantic, Indian
Ocean, Red Sea
spiny dogfish
(Squalus acanthias)
worldwide
antidiabetic; high affinity
binding to human insulin
receptor; although markedly
different than human insulin
Hthe binding sites for the human
I receptor in the same location
A Chain GIVDHCCHNTCSLYDLEGYCNO
B Chain LPSQHLCGSHLVETLYFVCGOKGFYYVPKI
Insulin - AA sequence from insulin of the hammerhead shark
(128)
128
photo: 97
antidiabetic; causes fat-specific
weight loss; Phase I trials had very
promising results; Genaera started
Phase I in 2007 and reported
promising results in 2009. Shortly
thereafter, Genaera was dissolved,
• and trodusquemine was sold to Ohr
| Pharmaceuticals; current status
unknown
92,98
photo: 93
Anti-infective drugs
Chemical Source
hydroxy-
curcuphenol
3,3'-oxybis[5-
methyl-phenol]
abyssomicin C
Acyclovir
(ACV,
acycloguanosine,
Zovirax*)
(derived from
spongouridine &
spongothymidine)
aigialomycin D
sponge
(Didiscus oxeatd)
Jamaica
fungus
(Keissleriella sp.)
actinobacterium
(Verrucosispora marts')
sponge
(Tectitethya crypto)
(synonym: Cryptotethya crypto)
Caribbean
mangrove fungus
(Aigialus parvusBCC 5311)
tropics
Uses & Status
antimalarial;
P. falciparum inhibition
antifungal; C. albicans,
T. rubrum &A. niger
inhibition
antibiotic; inhibits biosynthesis
of p-aminobenzoate (pABA), a
pathway not found in humans;
Phase I trials
antiviral; Acyclovir differs
from previous nucleoside
analogues in that it contains
only a partial nucleoside
structure: the sugar ring is
replaced by an open-chain
structure; clinical use
P. falciparum inhibition
Structure
OH
rti
S,^CH,OH
MjC'^CHj
|tH*)-»»«»»»ircm»»plniul (128)
^v^&»
3,3--oxybi*[5-methyl-phenol]
^cf°
' ^^0,
MO
Abyssomidn C ,•99',
HpN^N^Sl f-°H
^ (102)
OH O i
CT****"!
IJrr-
Citations
128
128
99,100,101
102
128
-------
Anti-infective drugs
Chemical Source
amphilactams
Ara-A
sponge
(Amphimedon sp. )
Australia
(photo '.A. queenslandicd)
%BB^ ' v
fi^^^
gorgonian
(Eunicella cavolini)
Mediterranean
^^™j™^«
(vidarabine, H^B^KT J'- ^A.
Vira-A*)
(derived from
spongouridine &
spongothymidine)
Ara-C
(Cytosar-U*
DepoCyt®,
Tarabine PFS®,
cytarabine, 1-p-D-
arabinosylcytosine)
(derived from
spongouridine &
spongothymidine)
arenosclerins
(similar to
haliclonacyclamine)
ascosali-
pyrrolidinone A
HF*I ':' Ml
'jfr • 3 ' ' j.. " ,S
1 J OY"=~Y'
Amphilactam D (103)
MH2
l*f)—U
--C ^
HO, ^
1 O^,^
\ i/
011 (102)
HH;,
HO *&
\* in/
\__y
1
OH (102)
<7y7^
^—\ H ^-K
S 'l
\ ^
arenosclerin A (128)
C4H, VN-f°
c^&^
H
Ascosahpyrrolidinone A (103)
c,X^^x^%Xx
OtSsC*
O 0*"^^^^
® '/.-^O.
" «H (103)
HO^-^^A
~L /--OH
/^^
ct^CH>
L CH. averol
(107)
0
w
f^T^t^
ST^
•-',.. (103)
103
photo: 104
1 O 1 Q
i o, i y,
105,102
photo: 105,
18,19,
102,103
128
103
103
106,107
103
-------
Anti-infective drugs
Chemical
axisonitrile-3
azidothymidine
(AZT,
zidovudine,
Retrovir*)
(derived from
spongouridine &
spongothymidine)
basiliskamide A
&
basiliskamide B
bengamide
derivative
bengazole A
bogorol A
bromosphaerone
calyceramides
A-C
Source
sponge
(Acanthella klethra)
Australia
:V*\
(3p"
sponge
(Tectttethya crypto)
(synonym: Cr\-ptotethya crypto)
Caribbean
bacterium
(Bacillus laterosporus)
sponge
(Jaspis sp.)
n~
sponge
(Jaspis sp.)
Fiji
•K
^E_
bacterium
(Bacillus sp.)
red algse
(Sphcerococcus coronopifolius)
Atlantic coast of Morocco
sponge
(Discodermia calyx)
Japan
Uses & Status
antituberculosis
antiviral; reverse-transcriptase
inhibitor; clinical use
antifungal; C. albicans &
A. fumigatus inhibition
anticancer & antihelminthic;
methionine aminopeptidase
(Met- API) inhibitor; licensed
to Novartis; withdrawn from
Phase I trials in 2002
antifungal
antibacterial; inhibits
antibiotic-resistant
S. aureus & enterococci
antibiotic
antiviral; neuraminidase
inhibition
Structure
i-""\/ S.
Axisonitrile-3 (103)
^NH
HTX
P1
N=N=N (1Q9)
bamlllskamld. A § (12g)
.
Y W
—
"•'
(22)
HO HO HO
N^i xxxx
Bengazole A (103)
W O \^j- )*~f*
\^/ OM
— V_^° bogorol A
NH
(128)
fXftfL
S?
n .; ... i( . (103)
Calyceeamiae A R, • C,^ Hj . djH^
Calvceiafrtde C R,.CnH;4 R2»C)tH24 n28't
Citations
103
photo: 108
18,19,102
106,109
128
18,22
photo: 23
103
photo: 23
128
103
128
-------
Anti-infective drugs
Chemical Source
caminoside A
chalcomycin B
clathsterol
corticatic acid A
&
corticatic acid B
cribrostatin 3
cyanthiwigin C
diazepinomicin
(ECO-4601)
dicynthaurin
didanosine
(2',3'-dideoxy-
inosine, ddl, DDI)
(derived from
spongouridine &
spongothymidine)
didemnin B
sponge
(Caminus sphceroconid)
Caribbean, Brazil
actinobacterium
(Streptomyces sp.)
sponge
(Clathria sp.)
IledSea
sponge
(Petrosia corticatd)
Western Pacific
sponge
(Cribrochalina sp.)
worldwide
sponge
(Myrmekioderma red)
(synonym: Myrmekioderma styx)
Jamaica
actinobacterium
(Micromonospora sp.)
tunicate
(Halocynthia aurantium)
North Pacific
sponge
(Tectitethya crypto)
(synonym: Cryptotethya crypto)
Caribbean
tunicate
(Trididemium solidum)
Caribbean
Uses & Status
antibacterial; inhibits
antibiotic-resistant
S. aureus & enterococci
antibacterial; inhibits
antibiotic-resistant S. aureus
antiviral; HIV reverse
transcriptase inhibition
antifungal; C. albicans &
A. fumigatus inhibition;
selective GGTase 1 inhibition
(enzyme is involved in fungal
cell wall biosynthesis)
antibiotic & anticancer
antituberculosis
antibiotic & anticancer;
Phase I trials (antibiotic &
anticancer)
antibacterial; gram negative &
gram positive inhibition
antiviral; clinical use
anticancer & antiviral interrupts
protein synthesis in target cells by
binding noncompetitively to
palmitoyl protein thioesterase;
cytotoxic for lymphomas, some
leukemias & melanomas;
antiviral for herpes simplex and
several others; Phase II trials
showed significant toxicity at
efficacious doses; dropped in
middle 1990s
Structure Citations
O OH
c»mlno*ld« A s**Q f*°«y^-v*OH
•^-^-Oy^Y^a^-o-^-
a"-" Y°°"
un I O-./^.O"
-r^o \ 1
J^ o-V^°"
OH chalcomycirt B -i 28')
r" o
QJ 0>-
, \ \ HOLJL — \. ONa
^ °^sP
/ clathsterol // "ONa
(128)
OH
corticatic acid E (128)
0
yv*i
HINYXN
oX^
Cnbiostatin 3 ' (103)
r~*\js^) "OH
eyanthlwlglnC (128)
'' CH, CM} CHj
H° ux" \ \^ diazepinomicin
H >-v^-0" ECO-4601
HO (35)
ILQKAVLDCLKAAGSSLSKAAITAIYNKIT
dicynthaurin (128)
N~^X
>w
\ / (102)
-(hi
f" 1 1""
A.^ »y"^_o
£j^X ^ ^
•* ^__ (41)
128
128
128
photo: 110
128
103
128
36
128
102
18,24,
39,40,41,41
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
...
dmydroxytetra-
hydrofuran
brown algas
(Notheia anomaki)
Australia
antiparasitic; in vivo tests did
not duplicate in vitro success,
probably because drug is
strongly hydrophobic
Trans-dihydroxytotrahydoluran
(103)
103
sponge
di-isocyano- (Pipestela hooperi)
adociane (synonym: Cyrnhi^>e]a hooperi)
Australia
antimalarial
H/'-NC
Diisocyanoadoctane HQ3)
discorhabdins
sponges
(Latrunculia sp.)
New Zealand
antibacterial, antifungal, &
antitumor
Discorhabdin A
(106)
106,111
ent-8-hydroxy-
manzamine A
sponge
Indo-Pacific
antimalarial;
P. berghei inhibition
(128)
128
enterocin
actinobacterium
(Streptomyces maritimus)
antibiotic & antiviral
gorgonian (sea whip)
(Pseudopterogorgia elisabethce)
Caribbean
ergorgiaene
&
7-hydroxy-
ergorgiaene
antituberculosis
H3C°
Ergorgiaene (103)
antiviral; Four types of
eudistomins: unsubstituted,
pyrrolyl-substituted,
pyrrolinyl-substituted, &
tetrahydro-p-carbolines
H H1 1
12,103
photo : 11
103
photo : 113
Pyrrolyl-substituted Eudistomins with
eudistomins (103) oxathiazepine ring (103)
anticancer & antimalarial
128
fascaplysin
sponge
(Fascaplysinopsis sp.)
Indo-West Pacific
antifungal
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
epiphytic fungus
(Acremonium sp.) on tunicate
(Ecteinascidi-,i turbinatd)
Caribbean
fumiquinazoline
antifungal
gambieric acids
dinoflagellate
(Gambierdiscus toxicus)
Central Pacific, Gulf of
Mexico
antifungal
114
photo: 50
103
sponge
(Geodia sp.)
Australia
geodin A Mg salt
antiparasitic
stalked crinoid (sea lily)
(Neogymnacrinus richeri')
synonym: Gymnocrinus richeri)
New Caledonia
halichondramide
sponge
(Halichondria sp.)
Indo-West Pacific
antiviral
antifungal & antimalarial
(103)
OH 0 OH 00
OH 0 OH HO
Gymnochrome D /103N
103
photo: 115
103
photo: 116
103
Hahchondtumide
O OCH3
(103)
haliclona-
cyclamines
(similar to
arenosclerins)
sponge
(Arenosclera brasiliensis')
Brazil
antibacterial
haltclonacyclamlne E
(128)
-------
Anti-infective drugs
Chemical
haliclonadiamine
halishigamide A
halocidin
halorosellinic
acid
hennoxazole A
leptyl prodigiosin
icadamides
iyengaroside A
Source
sponge
(Haliclona sp.)
Indian Ocean
(photo : Haliclona sarai)
VT* ""-HOf
CT
kJ
sponge
(Halichondria sp.)
Indo-West Pacific
tunicate
(Halocynthia aurannuni)
Northern Pacific
fungus
(Halorosellinia oceanica BCC
5149)
Thailand
sponge
(Polyfibrospongia sp.)
Indo-Pacific
commensal bacterium
found on tunicate
Phillipines
sponge
(Leiosella sp.)
Phillipines
algse
(Codium iyengarii)
Arabian Sea
Uses & Status
antifungal
antifungal
antibacterial; inhibits
antibiotic-resistant S. aureus &
MDR-resistant P. ceruginosa
antimalarial;
P. falcipamm inhibition
antiviral
antimalarial;
P. falcipamm & P. berghei
inhibition
anticancer, antiviral, &
immunostimulant
antibacterial; gram negative &
gram positive inhibition
Structure
i H
(5^^o
Haliclonadiamine (103)
CH3 CH3O 0 CH30 O-^^O ^*"v
— \\$
O OCHj
(103)
WLNALLHHGLNCAKGVLA
1
ALLHHGLNCAKGVLA
Two amino acid sequences (one 18-imit
& one 15-unit) joined by a disull'ide bond
between a cysteine residue in each unit
halocidin (128)
" ^-f K5 'OH
haloro««llinic acid (128)
f
1 9°"'
"^HH^-^UxO,
-^ > (103)
™v.
w£r
(CH,),CH3
h.plyl prodlgloiln (128)
OH 0 COOH
o"0^ S \^f" H.H d
^~Y ' *^-° IcaitonikhB NM
" (106)
i*
Q ly«ngaro*ide A f-i~)o\
Citations
117
photo: 71
103
128
128
103
128
59,106
128
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
sponge
(Jaspis sp.)
Fiji
jasplakinolide
(jaspamide)
antifungal & antiparasitic;
P. falciparum inhibition
103,128
photo: 23
nudibranch
(j'orunna funebris)
Pacific
Jorumycin
jorumycm
antibiotic
103
photo: 96
(103)
sponge
(Acanthella sp.)
(photo:^ caven/c
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
lepadin E
&
lepadin F
tumcates
(Clavelina lepadiformis [photo]
Aplidium tabascum,
Didemnum sp.)
Mediteranean,
Indo-West Pacific
antimalarial;
P. falciparum inhibition
128
photo: 120
(128)
litosterol
alcyonarian (soft coral)
(Litophyton viridis)
Okinawa
antituberculosis
manzamine A
&
ircinol A
(its likely biogenic
precursor that lacks
the p-carboline
moiety)
sponge
(Haliclona sp., Pachypellina sp.)
Indo-Pacific
(photo: Haliclona sarai)
manzamine F
sponge
Indo-Pacific
antitumor, antimalarial, anti-
infective, antituberculosis,
antitoxoplasmosis, &
antineurogenic inflammation;
has shown activity against
malaria, tuberculosis, HIV, and
other inflammatory diseases;
preclinical studies
H CH2OH
103,117
photo: 71
Ircinol A
(117)
antimalarial;
P. berghei inhibition
F (128)
marinomycins
A-D
actinobacterium
(Marinispora sp.)
antibiotic; potent activity
against drug-resistant bacterial
pathogens & some melanomas
44
(44)
bacterium
antibiotic
meridine
sponge
(Corticium sp.)
Bahamas
antifungal
(103)
microspinosamide
sponge
(Sidonops microspinosa)
Philippines
antiviral; HIV-growth
inhibition
103,121,128
mimosamycin
commensal actinobacterium
(Streptomyces lavendulce) found=
on sponge (Petrosia sp.)
Western Pacific
antibiotic; a neutral antibiotic
mainly active against
mycobacteria
106
muqubilin
sponge
(Prianos sp.)
Red Sea
antimalarial
102
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
mycalamide A
&
mycalamide B
sponge
(Mycale sp.)
New Zealand
antiviral; protein synthesis
inhibitors; mycalamide B has
shown greater antiviral activity
OCH3
Mycalamide A H
Mycalamide B CH (103)
nafuredin
fungus
(Aspergillus niger)
anthelmintic; inhibition of
helminth NADH-fumarate
reductase; competes for the
quinone-binding site
HOC
namenamicin
ommensal bacterium found
on tunicate (Polysyncraton
lithostrotuni)
Australia, New Zealand
antitumor & antibiotic; the
enediyne moiety is very
reactive with DNA, making
these chemicals extremely
cytotoxic for all cells & among
the most potent antitumor
agents known
.
Y-N
CM,
Narocnimicjn
(72)
72
nco-Kauluamine
«eo-kauluamine
sponge
Indo-Pacific
antimalarial;
P. berghei inhibition
(103)
103,128
sponges
(Theonella sp. &
Trachycladus Icevispimlifer)
Okinawa, Australia
(photo: Theonella cylindrical
onnamide A
onnamide A
&
onnamide F
anticancer, antifungal, &
anthelmintic
pannosanol
&
pannosane
red algse
(Laurencia pannosa)
Malaysia
(75)
antibacterial
75,76,
114,128
photo: 77
papuamide A
sponges
(Theonella mirabilis &
T. swinhoei [photo])
Indo-West Pacific
antiviral
NH „ o°VNH
VvVv
' OH
o ' OH (103)
103
patagonicoside A
echinoderm (sea cucumber)
(P solus patagonicus)
Southern Ocean
antifungal; Cladosporium
cucumerinum inhibition
128
pestalone
marine fungus (Pestalotia sp.)
found on brown algse
(Rosenvingea sp.)
Bahamas
Pestalone
P- OH
antibacterial
103,128
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
sponge
(Phorbas sp.)
Australia
ioto: P. tenacior)
phorboxazole A
antifungal
i
plakortide F
sponge
(Plakinastrella onkodes)
Jamaica
antimalarial;
P. falcipamm inhibition
Plakortido F
103
photo: 122
plakortolide
&
plakortolide G
(a 2nd cyclic
peroxide)
sponges
(Plakortis sp. [photo] &
Plakinastrella onkodes)
Pacific (Plakortis sp.)
Jamaica (P. onkode:
antiparasitic & antiprotozoal;
active agamsiLeishmania spp.
parasites
2ntl cyclic peroxide (103)
polyacetylenetriol
sponge
(Petrosia sp.)
Mediteranean
antiviral; RNA- & DNA-
directed DNA polymerase
inhibition
polyac«tyl«netriol
Q28)
103,128
photo: 123
polyester
15G256P
fungus
(Halorosellinia oceanicd)
(synonym: Hypoxylon
oceanicum)
Indian Ocean
antifungal; cell wall
biosynthesis inhibition
poly»»t»r 1SG236A
(128)
gorgonian (sea whip)
(F'seudopterogorgia elisabethce)
Caribbean
pseudopter-
oxazole
antituberculosis
X
Pseudopteroxazole (103)
ptilomycalin A
sponge
(Ptilocaulis spiculifer)
Australia
antifungal
Ptilomycalin A
12,103
photo: 11
103
photo: 124
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
puupehenone
sponges
(from Order Verongida &
Order Dictyoceratida)
Hawaii
antituberculosis
Puupehenone (103)
103
renieramycins
sponge
(Reniera sp.)
Pacific
antibiotic;
(102)
102
safracins
bacterium
(Pseudomonasfluorescens)
antibiotic & antitumor
R Sriracln: R = OH
Cytmosafracin: R = CN
(106)
106
saframycins
actinobacterium
(Streptomyces lavendulce)
antibiotic & anticancer
102
salinosporamide A
actinobacterium
{Salinispora tropicd)
tropics
antibiotic & cytotoxin; very
potent proteasome inhibitor; A
& B inhibited colon cancer
cells in vitro; A was extremely
potent against NSCLC, CNS, &
breast cancer lines;
in Phase I trials
Salinospotamide A
(44)
32,44
sigmosceptrellin sPonges
(Sigmosceptrella sp. &
Diacarnus erythrceanus)
Red Sea
&
sigmosceptrellin B
antimalarial & antiparasitic
gorgoman
(Briareum asbestinum)
Gulf of
Mexico
solenolide A
antiviral
103
photo: 125
Solenolide A
(103)
-------
Anti-infective drugs
Chemical
Source
Uses & Status
Structure
Citations
deep-water sponge
(Spongia sp.)
(photo: S. officinalis)
Caribbean
spongiadiol
antiviral
spiny dogfish
;Sqi<>.:l;fS 'jcantkias)
worldwide
squalamine
anticancer, antiangiogenic, &
antibiotic; broad spectrum
antibiotic; also shows anti-
angiogenic activity and may be
useful to treat age-related macular
degeneration; Phase II trials for
nonresponding solid tumors as part
of combination therapy and for
advanced ovarian cancer as primary
treatment; Phase III trials for wet
macular degeneration show
significant activity
103
photo: 126
18,44,
92,103
photo: 93
Sumiki's acid
(acetyl derivative)
antibacterial; B. subtilis &
S. aureus inhibition
Yl
o
COOH
(128)
Sumiki's acid
sponge
swinhoeiamide A (TheonelL: ^*i>ihoc^
Indo-West Pacific
antifungal; C. albicans &
A. fumigatus inhibition
swinhoeiamide A
(128)
sea grass
(Thola^ia icwdinun
Gulf of Mexico
O OH
thalassiolins
A-C
:O3SO OH
antiviral; HIV-1 integrase
inhibition
'O
CH;>OH
128
photo: 127
Thalassiolin A: R=OH
Thalassiolin B: R=OMe
Thalassiolin C: R=H
(128)
thyrsiferol
red algse
(Laurencia vem^ii
Indian Ocean
HO ,
antiviral
iroir-'
Thyrsiferol 18S. 19R
wailupemycins
actinobacterium
(Streptomyces maritimus)
antibiotic & antiviral
Waifupemycin D
106
(106)
xestodecalactone B
fungus (Penicillium cf
monanense) found on
sponge (Neopetrosia exigua)
(synonym: Xestospongia exigua)
Indian Ocean
antifungal; C. albicans
inhibition
xektodecalactone B
(128)
P ° antibacterial; Rhodospmllum
zamamistatm (Pseudoceratina vurpurea) . .f.
„, . salexieens inhibition
Okinawa
128
-------
Anti-infective drugs
Chemical
zopfiellamide A
&
zopfiellamide B
Source
fungus
(Zopfiella latipes}
Uses & Status
antibacterial; gram negative &
gram positive inhibition
Structure Citations
"^V-X zopflflllamide A R'CH,
'.' y — lopfloll.mld. B H3CH.CH,
(128)
128
Cardiac & circulatory system drugs; Anti-angiogenesis drugs
Chemical
Photo of Source
Uses & Status
Structure
Citations
2,5,6-tribromo-l-
methylgramine
(TBG)
bryozoan
(Zoobotryon verticillatum)
(synonym: Z. pellucidum)
worldwide
cardiovascular; vasorelaxation;
Ca2+ inhibition & increase
cyclic AMP
-CK,
128
aeroplysinin-1
sponge
antiangiogenic
Aeroplysinin-1 (128)
bryoanthra-
thiophene
bryozoan
inhibits angiogenesis
128
cortistatms
sponge
(Corticium simplex)
Australia
possesses a highly selective &
perhaps mechanistically unique
antiangiogenic activity
129
tunicates
(Clavelina moluccensis &
C. lepadiformis [photo])
Mediteranean,
Indo-West Pacific
lepadiformine
cardiovascular; inhibition of
cardiocirculatory system;
reduction in inward K+ current
128
photo: 130
Lepaditormine
(128)
polymeric 1,3-
alkylpyridinium
salts
sulfated
a-L-fucan
sponge
(Haliclona (Rhizoniera) sarai)
(synonym: Reniera sarai)
Mediteranean
sea urchins
(Echinometra lucunter [photo] &
Strongylocentrotus franciscanus)
Brazil & Gulf of Mexico
blood coagulation & platelet
aggregation; induces blood
coagulation, platelet
aggregation, & cytotoxicity in
rats; previously found to be
cholinesterase inhibitors
128
photo: 71
polymeric 1,3-alkylpyridinium salts
anticoagulant
128
photo: 131
sulfated a-L-fucan
S. franciscanus
(128)
-------
Cardiac & circulatory system drugs; Anti-angiogenesis drugs
Chemical
Photo of Source
Uses & Status
Structure
Citations
sea urchins
(Echinometra lucunter &
Strongylocentrotus franciscanus
[photo])
Brazil & Gulf of Mexico
sulfated
a-L-galactan
wondonin A
&
wondonin B
sponge
II
anticoagulant
HO
tjJ/CHjOH
-03SO/
0--
128
photo: 132
sulfated oc-L-galactan
£. lucunter
WondonviA repimer
WondoTin B 2 «
modulation of angiogenesis
sponge
(Xestospongia sp.)
(photo: Xestospongia
testudinarid)
Okinawa
xestospongin C
vasodilation; a potent, cell-
permeable inhibitor of Ca;
inhibits voltage-dependent Ca
& K currents
133
photo: 134
Immunological drugs
Chemical
Source
Uses & Status
Structure
Citations
sponge
(Stylissa massd)
(synonym: Stylotella aurantium)
Indo-West Pacific
(-)-palau'amine
immunosuppressant; strongly
cytotoxic; antibiotic
contignasterol
sponge
(Neopetrosia contignatd)
(synonym: Petrosia contignatd)
Indonesia
antihistamine & antiasthmatic
(-)-palau'amine
135
photo: 136
16,18
discodermolide
deep-water sponge
(Discodermia dissolutd)
Caribbean
anticancer & immunosuppres;;:ivc:
tubulin polymer stabilizer (like
taxol); essentially arresting cells at <
specific stage in the cell cycle and
halting cell division; Phase I trials;
licensed to Novartis by Woods
Hole; may be used in combination
with Taxol
^
24,25,42,43
(43)
-------
Immunological drugs
Chemical
Source
Uses & Status
Structure
Citations
Domoic acid
(causes amnesic
shellfish poisoning
in humans)
diatom
(Pseudo-nitzschia sp.)
worldwide
COOH
immune system; limits TNF-a
& matrix metalloproteinase-9
release from brain microglia
halipeptin A
&
halipeptin B
sponge
(Haliclona sp.)
Indo-Pacific
(photo: Hall lorn -...v ,
*%
MMpenn A
i B n«H
anti-inflammatory; inhibition of
carrageenan-induced edema
hymenamide C
sponge
(Stylissa carteri)
(synonym: Axinella carteri)
Indo-West Pacific
103,117
photo: 71
anti-inflammatory; neutrophil
& macrophage mediator
modulation
HOCX/
manoalide
(AGN-190093)
(related to the
cacospongiolides
from the same
sponge)
sponge
(Luffariella variabilis}
Indo-Pacific
anti-inflammation; antipsoriatic;
nervous system; first substance ever
observed to selectively inhibit
phospholipase A2 (PLA2); inhibits
seizures & epileptogenic properties
of crotoxin; pharmacological
probe; commercially available;
licensed to Allergan; Phase II trials
stopped due to formulation
problems (insufficient dermal
absorption); research continuing
18,24,25,128
137,138
(138)
fungus (Acremonium sp.)
found on tunicate
(Ecteinascidia turbinata)
Caribbean
oxepinamide
anti- inflammatory
114
photo: 50
petrosaspongiolide
sponge
(Petrosaspongia nigrd)
New Caledonia
gorgonian (sea whip)
(Pseudopterogorgia elisabethce')
Caribbean
pseudopterosins
(including
methopterosin, a
derivative)
anti- inflammatory;
phospholipase A2 inhibition
arthritis, anti-inflammation, &
analgesic; modify the arachidonic
acid cascade; inhibit synthesis of
eicosanoids, (hormone-like
substances) in specific white blood
cells (polymorphonuclear
leukocytes); extremely selective;
they appear to be pharmacologically
distinct from other NSAIDs; novel
MOA; Phase II trials; sold as a
cosmetic "anti-wrinkle" cream by
Estee Lauder under the name
Resilience
9,10,12,
24,25
photo: 11
pseudopterosin A
-------
Immunological drugs
Chemical
salinamide A
&
salinamide B
scytonemin
Source Uses & Status
bacterium
cyanobacterium
thalassospiramide A
„ bacterium
OL
i( , . • i T-. (Thalassospira sp.)
thalassospiramide B
topsentin Bl
totepsin D
deep-water sponges
(Halichondria genitrix
[synomyn: Topsentia genitrix],
Hexadella sp., Spongosorites
ruetzleri)
Mediteranean, North
Atlantic, Caribbean
sponge
(Spongosorites sp.)
anti-inflammatory
anti-inflammatory &
anticancer; inhibition of PMA-
induced mouse ear edema;
inhibits active cell proliferation
immunosuppressant
anti-inflammatory; suppresses
immunogenic & neurogenic
inflammation; early research
suggests potential use for colon
cancer, Alzheimer's disease, &
inflammatory bowel disease;
preclinical studies
anti-inflammatory
CH.SH, „
Structure
H,C j"'
i..JtH.B. ..CT c
H.c T~¥H"] 3 T 1
OH o J-o!ioo0^N.CHs
-TgS
H Cl— 'on
Citations
>
V (139)
un— $ \—3 n /=\ ScytOftemm
Cr/
XX^
^v\
;^V
(140)
jV\p-
tot«p.m 0
O
JH
(146)
25,139
128
44
140
146
Molecular probes
Chemical Source Uses & Status
jaspaquinol
(AGN-190093)
(related to the
cacospongiolides
from the same
sponge)
okadaic acid
saxitoxin
sponge
sponge
(Luffariella variabilis)
Indo-Pacific
red tide dinoflagellate
(Prorocentrum lima)
Northeast Atlantic,
molecular probe; human 15-
lipoxygenase inhibition
anti-inflammation; antipsoriatic;
first substance observed to
selectively inhibit phospholipase A2
(PLA2); inhibits seizures &
epileptogenic properties of crotoxin;
pharmacological probe;
commercially available; licensed to
Allergan; Phase II trials stopped due
to formulation problems
(insufficient dermal absorption);
research continuing
molecular probe; cellular
phosphorylation processes;
causes Diahrretic Shellfish
Poisoning (DSP); selective
inhibitor of the enzyme protein
phosphatase; pharmacological
probe, commercially available
molecular probe; neurotoxin;
ion channel nerve transmission;
pharmacological probe
Structure Citations
.^. -OH Jaspaquino) \^->v
iTT Jl
HO^^^-^V^-^V^-^X
V (128)
OH
5
H,C / y^\
CH| / T (
M,C 1 ^ (/
/ ^^ H: ^><
(138)
-M^T-«%Cb
(141)
TXtn
Hinx
""" MOXH
°" (142)
128
18,24,25,
128,137,138
12,24,25,
141
12,142
-------
Molecular probes
Chemical
Source
Uses & Status
Structure
Citations
tetrodotoxin
horseshoe crab (Limulus
polyphemus [NW Atlantic &
Gulf of Mexico],
Carcinoscorpius rotundicauda
[Asia], Tachypleus sp. [Asia]),
blue-ringed octopus
(Hapalochlaena lunulata
[Western Pacific] [photo],
Hapalochlaena maculosa
[Australia])
molecular probe; neurotoxin;
on channel nerve transmission;
pharmacological probe
HO
NH
(143)
12,142,143
photo: 144
Neurological drugs
Chemical Source Uses & Status Structure Citations
mollusk (sea snail)
(Hexaplex trunculus*)
1 Mediterranean
6-bromoindirubin
j^^fe
^^
anabaseine
(hoplonemertine
toxin)
antillatoxin B
aplysiallene
bipyridinyl
analog of
anabaseine
(synthetic analog of
anabaseine)
chlorogentisyl-
quinone
nemertine worm
anti-Alzheimer's; selective
inhibitor of
GSK-3P
anti-Alzheimer's & memory
enhancement; stimulates
vertebrate neuromuscular
(Paranemertes peregrind) nicotinic receptors & increasing
North Pacific cholinergenic transmission; has
cyanobacterium
(Lyngbya majuscula)
Indian Ocean, Gulf of
Mexico, Southeast Atlantic
sea hare
nemertine worm
(Paranemertes peregrind)
North Pacific
fungus
potential as a treatment of
cognitive function loss
nervous system; activates
voltage sensitive Na channel
nervous system; Na+, K+—
ATPase inhibition
nervous system; treatment of
neurodegenerative diseases;
patented by Memory
Pharmaceuticals Corp.
nervous system; neutral
sphingomyelinase inhibition
o (^~~~~h
H \ /
ff^f'-'x /^Y
^fl ~VNH
o
6-bromoindirubin ,14g.
N— v
/ \ / \
\i ff~~~\ /
\ / M f
anabaaeinv (147)
II ? 1
'"- f^^^u-^ ,O
Artilaloxin D fj ^^
? H f>~
^TX5V^*rVo''^^NYi
° Vs!
^ (128)
lT**l T^
TrV^N
(146)
146
photo: 145
18,146,147
128
146,147
128
-------
Neurological drugs
Chemical Source Uses & Status Structure Citations
conantokin G
conantokin L
Debromo
hymenialdisine
(DBH)
DMXB
(DMXB-A,
GTS-21)
mollusk (cone snail)
(Conus geographies')
Indo-Pacific
^^^P
P .
;^^^^Spf*
mollusk (cone snail)
(Conus lynceus)
Indo-Pacific
sponge
(Stylotella aurantium)
Palau
analgesic & anti-seizure; 17-
amino-acid competitive
antagonist of N-methyl-D-
aspartate (NMDA) receptors;
Phase I trials discontinued
anti-seizure & neuroprotective;
N-methyl-D-aspartate (NMDA)
receptor antagonist
anti-Alzheimer's &
osteoarthritis; acts as a highly
selective inhibitor of a specific
target cell DNA damage
checkpoint enzyme during the
G2 phase of the cell cycle;
Phase I trials
anti- Alzheimer' s & memory
enhancement; stimulates vertebrate
neuromuscular nicotinic receptors &
increasing cholinergenic trans-
nemertine worm mission; has potential as a treatment
(Paranemertes peregrina) of cognitive function loss; Phase II
(synthetic analog of North Pacific trial for treatment of Alzheimer' s
anabaseine)
dysiherbaine
halenaquinol
sponge
(Lamellof lysidea herbacea)
(synonym: Dysidea herbacea)
Indian Ocean
sponge
echinoderm (sea cucumber)
(Holothuria leucospilotd)
and schizophrenia; licensed to
Taiho by U. Florida; only marine
drug in trials for pallative treatment
of Alzheimer's (2005)
nervous system; induces
convulsant action in mice;
inhibits kainic acid & mGluRS
glutamate receptors
nervous system; Na+, K+-
ATPase inhibition
Indian Ocean
i ^fc ' ' ^B ' ' f '
HLG-1, 3l
HLG-2
nervous system
HLG-3 iBMtt^^k.
w ^^^B
1 SiB
precursor amino acid sequence
shown at (128, Figure 2, p. 275)
precursor amino acid sequence
shown at (128, Figure 2, p. 275
..
HN. y \\ H
\ — / J —
°-=._r \c^L L
"l f^Tl^>***NHj
HN J __
^^ (149)
1 N
[| J
^O ^^\
\
f^^^^f^ N
1
O'^^^^ ^^^
1
OMXBA (GTS-21) ,14g.
'
•a»>e i'19S'i
I, 1 Z.O }
O
J?
iOcoH
Halenaquinol Q 2 8 ^
MUM sfV^
U|y"^K^. 1...19
OCHjCOHH &*:)COM7JA_rt3iCOOM ' O*1 ™ "" * 19'3C'!I
A °\l I/V-OM coon 1^ V-wO i QH
Jv
(128)
1,2,18
photo: 3
128
photo: 148
149
photo: 149
146,147
128
128
128
photo: 150
-------
Neurological drugs
Chemical
Source
Uses & Status
Structure
Citations
sponges
(Axinella damicornis [photo],
Axinella verrucosa, Stylissa
carteri [synonym: Acar^Al;
aurantiacd\)
Mediteranean, Atlantic
Ocean, Indian Ocean
hymenialdisine
anti-Alzheimer's & anti-
Parkinson's; ATP-competitive
inase inhibitor; inhibits cyclin-
dependent kinases & blocks in
vivo phosphorylation; Potent
inhibitor of mitogen-activated
protein kinase kinase-1
(MEK-1)
hymenidine
sponge
(Hymeniacidon sp.)
Okinawa
anti-Alzheimer's & anti-
Parkinson's; a potent antagonist
of serotonergic receptors
146,151,152
photo: 153
hymenialdisine
(146)
146,154
(146)
iantheran A
&
iantheran B
sponge
nervous system; Na+, K+—
ATPase and plasmin inhibition
128
la-tltwran A R.H
n A duicetat* R-Ac
(128)
linckoside A
&
linckoside B
starfish
nervous system; induces
neuritogenesis
L —^
maitotoxm
algiE
nervous system; modulates
calcium & sodium influx
12S
(128)
manoalide
(AGN-190093)
(related to the
cacospongiolides
from the same
sponge)
sponge
{ftlrn?Ua variab>i
Indo-Pacific
anti-inflammation; antipsoriatic;
nervous system; first substance ever
observed to selectively inhibit
phospholipase A2 (PLA2); inhibits
seizures & epileptogenic properties
of crotoxin; pharmacological
probe; commercially available;
licensed to Allergan; Phase II trials
stopped due to formulation
problems (with dermal application,
insufficient amount absorbed
through skin); research continuing
18,24,25,
128,137,13!
-------
tunicate
(Aplidium meridianum)
Southern Ocean
sponge
(Smenospongia aured]
Jamaica
N-3'-
ethylaplysinopsin
neodvsiherbaine A
mollusk (sea slug)
Gulf of California
(photo: Onchidella sp.)
sponge
(Agelas oroides')
Mediteranean
oroidm
(related to
hymenialdisine)
gorgoman (sea whip)
(F'seudopterogorgia elisabethce]
Caribbean
pseudopterosins
(including
methopterosin, a
derivative)
sponge
(Haliclona sp.)
Indo-Pacific
(photo: Haliclona sarai]
salicylihalimides
Neurological drugs
Uses & Status
Structure
Citations
anti-Alzheimer's; protein
kinase inhibitors
Mtridiiinin A
Mil iili:nriii It
Ml'l illi.1llill (
Mu jlljimli! F.
Mi'i iiliiinin I
Mi-liiliililin I.
4 5
OH H
Oil II
II Kr
II II
OH H
II
H
II
Hi
H
II,
•I
7
H
II
II
il
Br
H
II
nervous system; binds to
human serotonin 5-HT2c
receptor
155
photo: 156
nervous system; induces
convulsant action in mice;
inhibits kainic acid glutamate
receptors
128
anti-Alzheimer's; an active
site-directed irreversible
inhibitor of AChE
onch.d.il O
anti-Alzheimer's & anti-
'arkinson's; a potent antagonist
of serotonergic receptors
(146)
arthritis, anti-inflammation, &
analgesic; appear to modify the
arachidonic acid cascade; inhibit
synthesis of eicosanoids, (locally
functioning hormone-like
substances) in specific white blood
cells called polymorp ho nuclear
leukocytes; their extreme selectivity
intrigues to researchers; they appear
to be pharmacologically distinct
from other NSAIDs & their MOA
seems novel; preclinical studies;
sold as a cosmetic anti-wrinkle
cream by Estee Lauder under the
name Resilience
pseudopterosin A
anticancer & anti-osteoporosis;
first marine Vo-ATPase
inhibitor (Vo-ATPases are
eukaryotic enzymes whose
principal role is to pump
hydrogen ions across cell
vacuolar membranes); may
mediate bone resorption;
preclinical studies
OH 0
SalicylihalamideA
146
photo: 157
146
photo: 158
9,10,12,
24,25
photo: 11
18,86
photo: 71
-------
Neurological drugs
Chemical | Source
trans-3-
cinnamylidene
analogs of
anabaseine
tridentatol
nemertine worm
(Paranemertes peregrind)
North Pacific
hydroid
(Tridentata marginatd)
Gulf of Mexico
Uses & Status Structure
nervous system; developed by
Univ. of Florida; potentially
useful for treating brain
nicotinic subtype receptors
antioxidant & UV protection
p ,jo 6
N IT ^N*
tran«-3-clnnamylld«n0 analog* of anal
Citations
fc
•~k» (146)
146,147
114
Templates
Chemical
Source | Uses & Status
spongouridine sPonge
v 6 (Tectitethya crypto)
spongothymidine
(synonym: Cryptotethya crypto)
Caribbean
template for antiviral &
antitumor agents; template for
nucleosides with sugars other
than ribose or deoxyribose; led
to synthesis of zidovudine
Structure Citations
ff s
H0 0*^rT HQ 0<^N^
YT yy
(AZT) Ol ' *
spongouridine spongothymidine (102)
19 102
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kinase inhibitors isolated from the AscidianAplidium meridianum. Bioorganic & Medicinal Chemistry Letters, 14:1703-1707.
156. http://www.boldsystems.org/views/taxbrowser.php?taxid=299527
157. http://en.wikipedia.Org/wiki/File:Onchidella.jpg
158. http://en.wikipedia.org/wiki/File: Agelas_oroides_Capo_Gallo_025.JPG
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Chapter 6
Coral Reefs: Summary
The ESRP coral reefs project is intended to advance the understanding of coral reef ecosystem services, how they are affected
by human activities, and how management and policy decisions influence their delivery. Clear definitions of the ecosystem
services provided, including the reef attributes that support these services are needed.
In the previous chapters, we have provided a review of past studies focused on four primary ecosystem services provided by
coral reefs (shoreline protection, fishing, tourism and recreation, and natural pharmaceutical and biochemical products). The
chapters have summarized the economic benefits provided by each of the services, the methods used to quantify the services,
and how those ecosystem services were linked to characteristics (attributes) of coral reefs. Table 6-1 shows the combined final
ecosystem services table.
6.1 Lessons learned
The management of ecosystem services and the establishment of policies that will result in the preservation of coral reefs will
require extensive collaboration between natural and social scientists. We need to better understand:
• The physical and biological processes that provide the ecosystem services;
• The nature of the ecosystem services and how they can be quantified;
• Sustainable exploitation of reef services; that is,
>• How do we define sustainable levels of exploitation?
>• How do we estimate a reef's carrying capacity or potential to supply a service? and,
• The social benefits derived from ecosystem services and how they can be measured.
Our decision framework needs to change. Decisions must move towards sustainability (that is, consideration of the "triple
bottom line", where economic, social, and ecological aspects receive equal consideration in decision-making). We are currently
hindered by our incomplete understanding of all the benefits provided by coral reefs. We need to look at this from two
directions:
• On one hand, what does society gain by having the reef.
• On the other hand, what do we lose if the reef goes away, (e.g., "How much of the protein consumption in the USVI or
Samoa comes directly or indirectly from their coral reefs?" "How would that be replaced and at what cost.")
Measuring an ecosystem service is often confused with valuing that service. For the past twenty years, economists have been
conducting studies to estimate the economic value of coral reefs. These studies have often focused on a single ecosystem
service, and data limitations have often precluded comprehensive measurements and valuations. In addition, some services
provided by coral reefs have been ignored in these studies. For example:
• Conotoxins provide significant improvement in quality of life as compared to opiate painkillers in some therapeutic
situations, because conotoxins reduce pain more effectively than opiates without inducing sleep or impairing cognitive
functions. How do we most appropriately capture the value of this improved quality-of-life.
• There are over 10 million surfers. Many of the best surfing locations are provided by reef breaks. This service has not
yet been quantified.
• Coral reefs provide protection from flooding during storms. Better quantification of this service could potentially
support reduced flood insurance rates for those areas protected by reefs.
Valuation tends to be more accurate at the micro level, where economists are able to apply methods for eliciting individuals'
values (both market and nonmarket). However, aggregation and estimation of nonmarket benefits at the macro scale remain a
challenge. As a result, coral reef ecosystem services have been undervalued, perhaps significantly. Most studies have not
directly measured the ecosystem services.
There is a need to consider all aspects of the ecosystem service that can be provided in the valuation process. The present value
of the actual use of the ecosystem service, the service that could potentially be available, and the service that would be available
under management for sustainable use should all be determined. For example, current practice does not include services being
provided but not exploited because supporting services (e.g., access) are not available. However, the development of supporting
infrastructure brings environmental impacts to the reef that must be considered in the light of sustainable use. We recommend
that all three aspects of the ecosystem service be quantified, when possible.
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6.2 Next steps
The ultimate goal of the ESRP coral reefs project is to identify coral reef indicators that can be used to estimate the quantity of
ecosystem services being delivered by coral reef ecosystems and to predict the extent to which the current quantities are likely
to change in the future. In 2011, we will fully document the linkages between coral reef condition, function, and ecosystem
services and the coral reef attributes that support those services. We will also create the coral reef system model that links the
coral reef ecological production functions with policy/management-relevant final ecosystem services. We will identify a
manageable suite of candidate indicators of final ecosystem services and directly tie the valuation activities to particular
management decision(s).
Our literature review has shown that many of the ecosystem indicators are already being collected. However, the aggregation of
the indicators and of the underlying data and relating them to ecosystem services is not being done. Missing items include
landscape attributes (both onshore and offshore), many of which have been collected but not analyzed for the purpose of
relating them to ecosystem services. We plan to begin research in this area using existing and new ORD expertise and
resources.
We intend to design the next generation of survey questionnaires, which will address the value that stakeholders assign to coral
reef attributes. In addition, modern technology would support web-based surveys, which could greatly facilitate the ease of
collecting and analyzing the survey data. One of our collaborators, NOAA's Bob Leeworthy, successfully used a web-based
survey questionnaire in a recent valuation of Hawaii's coral reefs. We are collaborating with Bob to design a web-based study
for the Guanica Bay Watershed. This is a longer-term research activity (2012-2013).
Table 6-1. Combined final ecosystem services and supporting features for coral reefs
Ecosystem Service(s)
Final (FES)
Ecosystem-
Natural Features
Social Values w>mpiememary
Intermediate eooas & services
Benefits
Potential
Indicators of Final
Ecosystem
Service(s)
Tourism & recreation
Recreational
Fishing
Opportunity
Recreational
Diving/Snorkeling
Opportunity
Production of benthic
and aquatic prey for
consumption by
recreational fish
Fish diversity and
abundance
Desirability offish
species and size
for rod-and-reel
catches
Adequate
infrastructure (boats,
marinas, etc.)
Revenues from
tourism and
recreation
activities
_ . . Coral diversity, Desirability of Access to reef, safe
' abundance and coral reef for swimming conditions, Revenues from
maintenance ot water ,,.,,., ^-11 , + >. • i
, . _ . . ,, health; fish recreation based adequate tourism and
clarity; Production ot , . . , , 1-1 • ^ + + „ + ,
, . , . diversity and on physical infrastructure (hotels, recreation
benthic and aquatic , , . ,. , ...
„ . . abundance; water appearance (color, dive boat operators, activities
prey for consumption . . . ...... . \ . ,
f . , .5 , clantv visibility, etc. ) etc. )
by recreational tish
^.».»..wwx x»..uuu»u Coral diversity, Desirability of 1 Access to reef, safe
Recreational . ' abundance and coral reef for swimming conditions, Revenues from
TT , , maintenance ot water ,,.,,-, ^-111^ >. • i
Underwater , . _ . ,, health; fish recreation based adequate tourism and
„, . clarity; Production ot , . . , , 1-1 • ^ + + „ + ,
Photography . ,. . . diversity and on physical infrastructure (hotels, recreation
°. .. benthic and aquatic , , . ,. , ...
Opportunity „ , . abundance; water appearance (color, dive boat operators, activities
prey for consumption . . . ...... . \ . ,
: . .-. clantv visibility, etc.) etc.)
Recreational
Surfing
Opportunity
Opportunity to
View Nature and
Wildlife
by recreational tish
Reef breaks
Biological integrity
Opportunity to Water quality,
Sunbath and Swim shoreline protection,
at the Beach sand production
Opportunity to
Collect Objects
(beachcombing)
Water quality
3-D reef structure
Biodiversity (birds,
marine mammals,
turtles)
White coralline
sands; calm waters
Desirability based
on wave size and
speed
Desirability of
species (rarity,
size)
Desirability of
coralline sand
beach for
sunbathing (size,
cleanliness,
appearance)
.... . . Desirability ot
Wide sandy „ . , ,
. . walking on beach
beaches, , ° ,.
.... . , and ot finding
biodiversity, , ,.,-,„
/' beautiful &
occasional storms . . .
Access to reef,
adequate
infrastructure (hotels,
board shops, etc.)
Access to reef and
adequate
infrastructure (boats,
tour guides)
Access to beach
Access to beach
unusual objects
Revenues from
tourism and
recreation
activities
Revenues from
tourism and
recreation
activities
Revenues from
tourism and
recreation
activities
Revenues from
tourism and
recreation
activities
Abundance of
catchable snappers
and groupers
Taxa richness, size
and density of reef
organisms
Taxa richness, size
and density of reef
organisms
3-D structure and
proximity to deep
ocean
Taxa richness,
presence of
specific species
Areal extent of
beach, color of
beach, water
temperature, days
of sunshine, beach
trash
Areal extent of
beach, frequency
of storms,
proximity of reef,
taxa richness of
invertebrates
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Table 6-1 (continued)
Ecosystem Service(s)
Final (FES) Intermediate
Ecosystem-
Natural Features Social Values Complementary Derjved
Goods* services Benefjts
Potential
Indicators of Final
Ecosystem
Service(s)
Fishing
Seafood Products:
Fish, shellfish,
algae
harvested
Aquarium
Products (live fish
& coral
taken)
Material for
Curios and
Jewelry removed
Biological
integrity
Biological
integrity
Biological
integrity
Shoreline protection
Protection from Reduction in
shoreline erosion;
Decreased erosion
in kg/ha/y
wave energy,
velocity, or
height
coastal inundation
during extreme
events area in
hectares protected
Reduction in
wave set-up, or
storm surge
Fish diversity and
abundance; coral health; Desirability of Adequate
seascape connectivity; species based on infrastructure (boats,
and structural taste marinas, etc.)
complexity
Coral diversity,
• 1.-1-
abundance and health; ^*-°"-""""j ^
,_ , ,. . . species for aquaria
tish diversity and , , , • ,
based on physical
abundance; seascape . .
.. .. , appearance (color,
connectivity; and 1 . .
structural complexity
'
Coral diversity, Aesthetic values
abundance and health; and artistic
water clarity inspiration
Diving and boating
infrastructure
Revenue from
commercial
seafood fisheries
Revenue from
sales of
aquarium fish
and coral
Revenue from
sales of curios
and jewelry
Presence of reef; reef
height, width, slope, &
roughness; reef ... ,,1
. . Attractiveness of
continuity. . . . Absence of
„, . , ... sandy beaches; .
Physical variables: , .,.,.. f constructed
/-,«• u desirability of
Offshore wave energy, , . , breakwaters
... ... housing near water
wave height, tidal
depth; beach elevation,
sediment grain size
Presence of reef; reef
height, width, slope,
roughness; reef
continuity.
Physical variables:
Wave energy; distance
from hazard event;
slope of coastline;
frequency & intensity of
extreme events
Past history of
extreme events
Absence of
constructed
breakwaters;
location, intensity,
and value of coastal
development
Higher property
values;
opportunity to
use beaches (see
Ch. 2)
higher property
values;
lower property
damage and loss
of life
Abundance of
commercially
desirable fish
species
Species abundance
and diversity of
target populations
Species abundance
and diversity of
target populations
% reduction in
rates of shoreline
erosion due to
presence of reef
% reduction in
coastal inundation
due to presence of
reef
Pharmaceuticals from natural products
Marketable natural T , .
i >. >. Unique
product or a tern- .... ,.
, biologically
plate that results in , . ,
. . . active secondary
a marketable + , ,•+
. metabolite
product
Pharmaceutical
Shallow, marine Desirability of research programs for
biodiverse, species- good health and both field collection
dense ecosystem well-being and laboratory
analysis
Increased
revenues from
Pharmaceuticals;
increased health
and well-being
Species density,
biological integrity,
sponge diversity,
rare species
Definitions (proposed by the Ecosystem Services Research Program and currently under discussion by the Program)
• Final Ecosystem Service - Output of ecological functions or processes that directly contributes to social welfare or has the potential to do so
in the future (broadly based on Boyd & Banzhaff [2007]).
• Intermediate Ecosystem Service - Output of ecological functions or processes that indirectly contributes to social welfare or has the potential
to do so in the future.
• Natural Features - The biological, chemical, and physical attributes of an ecosystem or environment.
• Social Values - The social attributes that influence economic demand for an ecosystem service.
• Complementary Goods & Services - Inputs (usually built infrastructure or location characteristics) that allow a good or service to be used by
complementing the ecological condition. For example, complementary goods and services that allow the presence of fishable fish to become
an opportunity for recreational fishing will include aspects of site accessibility, such as road access, available parking and the presence of a
fishing pier, all of which make fishing at the site possible and enhance enjoyment of the activity.
• Ecosystem-Derived Benefits - The contribution to social welfare of ecosystem goods and services. In the ESRP, the term applies specifically
to net improvements in social welfare that result from changes in the quantity or quality of ecosystem goods and services attributable to
policy or environmental decisions.
• Indicator of Final Ecosystem Service - Biophysical feature, quantity, or quality that requires little further translation to make clear its
relevance to human well-being (i.e., "public-friendly" measurement)
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Coral Reefs: Glossary
Words in bold are defined in this glossary.
Acropora A genus of stony corals that contain the elkhorn and staghorn corals (NOAA 2010).
actinomycetes Gram-positive bacteria of the order Actinomycetales in the phylum Actinobacteria that are mostly aerobic but can
be anaerobic. Some resemble fungi, because they produce a characteristic, branched mycelium. Actinobacteria
are well-known sources of secondary metabolites having pharmaceutical uses. Often found in symbiont
relationships with megafauna, notably sponges.
alcyonarian An octocoral.
algae A large and diverse group of simple unicellular or multicellular organisms that use chloroplasts for
photosynthesis, although they are not plants. Algas, which are chiefly aquatic, form the basis of the marine food
chain. Common algas include dinoflagellates, diatoms, seaweeds, and kelp.
B
antifouling Agent that inhibits the growth of barnacles and other marine organisms on a ship's bottom (an antifouling paint or
agents other coating). Organotin compounds have been the most often used agents in this application since they are
effective against both soft and hard fouling organisms. However, in spite of their performance, they have a
negative impact on the marine environment, and their long half-life in the environment has prompted marine
paint manufacturers to look for a nonpersistent alternative.
artificial A bank or levee of stones or a timber structure, used to break the force of the sea in its entrance into a harbor or
breakwater roadstead.
ascidian Sac-like filter feeders that belong to the class Ascidiacea within the subphylum Tunicata. Often called sea squirts.
atoll reef A type of coral reef that encircles a lagoon partially or completely.
attribute Any measurable component of a biological system (Karr & Chu 1999).
Ayurveda An ancient medical treatise of the Hindu art of healing and prolonging life; sometimes regarded as a 5th Veda.
back reef The landward side of a reef between the reef crest and the land.
barrier reef A type of coral reef near the shoreline, but separated from it by a deep lagoon.
Bayesian belief A graphical network for modeling probabilistic interrelationships between events. Events are represented by
network (BBN) nodes in the network and causative relationships are represented by directed arrows between the nodes. A BBN is
especially useful when individual nodes of the network will be updated with evidence. For example, a BBN could
represent the probabilistic relationships between diseases and symptoms. Given symptoms, the network can be
used to compute the probabilities of the presence of various diseases. Decision and utility nodes can be added to a
BBN to represent and solve a decision problem following maximum expected value criterion (this is called an
influence diagram).
beachcombing The recreational activity of searching the beach and the intertidal zone for items that have washed in with the tide
(e.g., corals, seashells, sponges, sea fans, fossils, pottery shards, artifacts, sea beans, sea glass, and driftwood).
benefit transfer Techniques to estimate values of ecosystem goods and services based on previously conducted valuation studies.
Benefit transfer is conducted by either taking average values of existing studies or by using a transfer function to
transfer values from primary studies (study sites) to new locations (policy sites). A transfer function is often
developed through meta-analysis, which is a statistical (usually regression) technique to model differences in
values among primary valuation studies. A transfer function allows values to be transferred from study sites to
policy sites based on a set of independent variables that capture the degree of similarity between the study sites
and policy sites (Wainger & Mazzotta 2009).
biochemicals Chemicals that result from biological and chemical processes in living organisms.
biogeography The study of ecosystem geography to understand why flora and fauna are found in certain places.
biological diversity
(biodiversity)
The variability among living organisms from all sources including terrestrial, marine, and other aquatic
ecosystems and the ecological complexes of which they are part; this includes diversity within and among species
and diversity within and among ecosystems (MEA 2009).
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biophysical Pertaining to the biological, chemical, and physical attributes of an ecosystem or environment.
bombora A shallow area some distance from the shoreline that causes sea waves to break.
bryozoans Aquatic animals comprising the phylum Bryozoa that form mossy colonies of small polyps each having a curved
or circular ridge bearing tentacles; they attach to stones or seaweed and reproduce by budding.
carbon dioxide An odorless colorless gas formed during respiration and by the decomposition of organic substances; absorbed
(CO2) from the air by plants in photosynthesis. It is also a byproduct of burning fossil fuels and biomass, as well as land
use changes and other industrial processes. It is the principal anthropogenic greenhouse gas that affects the
earth's radiative balance.
charismatic Large animal species with widespread popular appeal that environmental activists use to achieve conservation
megafauna goals well beyond just those species.
clinical trials A scientifically designed and executed investigation of the effects of a drug (or vaccine) administered to human
volunteers. The goal is to define the safety, clinical efficacy, and pharmacological effects (including toxicity, side
effects, incompatibilities, or interactions) of the drug.
cnidarian Multicellular animals comprising the phylum Cnidaria (silent c), including the stony corals (scleractinians), soft
(the c is silent) corals (octocorals), anemones, sea fans, sea pens, hydroids, and jellyfish.
commercial fishing Fishing for profit.
complementary
goods and services
connectivity
contingent valuation
method (CVM)
Inputs (usually built infrastructure or location characteristics) that allow a good or service to be used by
complementing the ecological condition. For example, complementary goods and services that allow the presence
of fishable fish to become an opportunity for recreational fishing will include aspects of site accessibility, such as
road access, available parking and the presence of a fishing pier, all of which make fishing at the site possible and
enhance enjoyment of the activity.
A topological property relating to how geographical features are attached to one another functionally, spatially, or
logically.
A valuation method that estimates consumers' preferences by asking them how much they are willing to pay for a
benefit (willingness-to-pay or WTP), or what they are willing to accept by way of compensation to tolerate a loss
(willingness-to-accept or WTA).
coral The regulatory definition is: species of the phylum Cnidaria, including: (a) all species of the orders Antipatharia
(black corals), Scleractinia (stony corals), Gorgonacea (horny corals), Stolonifera (organpipe corals and others),
Alcyanacea (soft corals), and Coenothecalia (blue coral), of the class Anthozoa; and (b) all species of the order
Hydrocorallina (fire corals and hydrocorals) of the class Hydrozoa (16 U.S.C. 6401 et seq 2000). Current
taxonomy has corals and sea anemones grouped into the class Anthozoa within the phylum Cnidaria. Anthozoa is
divided into two subclasses, Octocorallia and Hexacorallia. Soft corals (including gorgonians) (order
Alcyonacea) and blue corals (order Helioporacea) are under Octocorallia, and stony corals (order Scleractinia),
black corals (order Antipatharia), and zoanthids (order Zoantharia) are under Hexacorallia. Neither fire corals nor
hydrocorals are technically corals: they are classified as phylum Cnidaria, class Hydrozoa, order Capitata.
coral bleaching The process in which a coral polyp, under environmental stress, expels its symbiotic zooxanthellse from its body.
The affected coral colony appears whitened (NOAA 2010).
coral cover The covering of the sea floor by coral. It can be measured in square miles, square kilometers, or as a percentage
of area with cover.
coral reef Any reefs or shoals composed primarily of corals.
coral reef ecosystem
Coral and other species of reef organisms (including reef plants) associated with coral reefs, and the nonliving
environmental factors that directly affect coral reefs, that together function as an ecological unit in nature.
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cultural services Cultural services are the nonmaterial benefits people obtain from ecosystems through spiritual enrichment,
cognitive development, reflection, recreation, and assthetic experiences, including: cultural diversity, spiritual and
religious values, knowledge systems (traditional and formal), educational values, assthetic values, social relations,
sense of place, cultural heritage values, and, recreation and ecotourism. In the lexicon of the ESRP, many of the
elements in the MEA category (e.g., spiritual and religious values) could be considered ecosystem services that
are best defined specifically in terms of beneficiaries' ethical or cultural value systems, while the benefits derived
from others (e.g., certain aspects of recreation and ecotourism) might be valued in more generic terms. See also,
provisioning services, regulating services, and supporting services (MEA 2005).
cyanobacteria Photosynthetic aquatic bacteria that compose the phylum Cyanobacteria. They are often called blue-green algas,
but have no relationship to algae. Cyanobacteria get their name from the bluish pigment phycocyanin, which they
use to capture light for photosynthesis. They also contain chlorophyll a, the same photosynthetic pigment found
in the chloroplasts of plants. Not all "blue-green" bacteria are blue; some common forms are red or pink, resulting
from the pigment phycoerythrin (NOAA 2010).
D
decision maker Individual(s) or groups of people responsible for making choices or determining policy that impacts the functions,
processes, and condition of ecological systems. Decisions may be local, regional, or national in scale.
demand Generally, the amount of a particular good or service that a consumer or group of consumers will want to
purchase at a given price. Demand for a good or service is determined by many different factors other than price,
such as the price of substitutability and complementary goods and services. Along with supply, demand is one of
the two key determinants of the market price.
direct use values Economic values derived from direct use or interaction with a biological resource or resource system.
ecological endpoint
A biophysical feature, quantity or quality that requires little further translation to make clear its relevance to human
well-being (i.e., "public-friendly" measurements). Ecological endpoints are the ecological inputs that, along with
complementary goods and services inputs and demands by people, produce ecosystem services. For example, the
abundance of watchable birds at a site is an ecological endpoint that, when combined with complementary inputs
such as transportation infrastructure and demand by birders, produces the ecosystem service of recreational bird
watching. Specified changes in ecological endpoints can be used in economic surveys to gauge people's
willingness-to-pay for (or willingness-to-accept) increases or decreases in potentially valued ecosystem services,
thereby providing quantitative information with which to evaluate decision/management mandates (adapted from
Boyd [2007], Boyd & Banzhaf [2007], Wainger & Boyd [2009] and Wainger & Mazzotta [2009]).
ecological integrity
ecological
production function
(EPF)
The capability of supporting and maintaining a balanced, integrated, adaptive community of organisms having a
species composition, diversity, and functional organization comparable to that of natural habitat of the region
(Frey 1975; Karr and Dudley 1981; Angermeier and Karr 1994; Karr et al. 1986).
A description of the type, quantity and interactions of natural features required to generate outputs of natural
products and services. For a simple example, the biophysical characteristics of a coastal wetland (flooding
regimes, salinity, nutrient concentrations, plant species abundance, prey and predator abundances, etc.) can
influence the abundance of a population of watchable wading shorebirds (the ecological endpoint). The outputs
of ecological production functions, when combined with complementary goods and services and demand by
humans, produce ecosystem goods and services (adapted from Wainger & Boyd [2009] and Wainger & Mazzotta
[2009]).
ecological resilience The capacity of a system to absorb disturbance and reorganize while undergoing change so as to still retain
essentially the same function, structure, identity, and feedbacks (Walker et al. 2004; Folke 2006).
ecosystem A dynamic complex of plant, animal, and microorganism communities and their nonliving environment
interacting as a functional unit (MEA 2009).
ecosystem functions Physical, chemical, and biological processes that occur in ecosystems.
ecosystem goods and
services
Outputs of ecological functions or processes that directly (final ecosystem service broadly based on Boyd &
Banzhaff [2007]) or indirectly (intermediate ecosystem service) contribute to social welfare or have the
potential to do so in the future. Some outputs may be bought and sold, but most are not marketed. Often
abbreviated as ecosystem services (modified from EPA [2006]).
ecosystem service Shorthand notation for an ecosystem good or service.
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ecosystem structure The individuals and communities of plants and animals of which an ecosystem is composed, their age and spatial
distribution, and the nonliving natural resources present. The elements of ecosystem structure interact to create
ecosystem functions.
effect on production An estimate of the difference in value of productive output before and after the impact of a threat or a
(EoP) management intervention.
emergy The available energy of one kind previously used-up directly and indirectly to make a product or service. Emergy
is expressed in its own unit, the emjoule, which connotes the energy of equivalent quality (e.g., solar emjoules)
used in the past to make a product or service (e.g., a wetland), as compared with the energy (J) content of the
product or service (Odum 1996).
enzymes Proteins that catalyze (i.e., increase the rates of) chemical reactions.
erosion Wearing away of rock or soil by the gradual detachment of soil or rock fragments by water, wind, ice, and other
mechanical, chemical, or biological forces.
ex-vessel price The price received by a commercial fishing captain for the catch (Thayer et al. 2005).
fauna Animal life, especially the animals characteristic of, or endemic to, a region.
final ecosystem Components of nature directly enjoyed, consumed, or used to yield human well-being (Boyd & Banzhaf 2007).
service
financial analysis Uses the observed current financial activities, revenues, costs, and financial flows in the economy from market-
based uses of the reef (such as diving and snorkeling) to analyze the economic activity generated by use of an
ecosystem good or service.
flora Plant life, especially the plants characteristic of, or endemic to, a region, period, or special environment.
fore reef The seaward edge of a reef that is fairly steep and slopes down to deeper water.
fringing reef A type of coral reef that borders the shoreline, separated from shore by only a shallow lagoon or none at all.
functions The physical, chemical, and biological processes that occur in ecosystems.
fungi A kingdom separate from animals, bacteria, and plants consisting of usually multicellular, heterotrophic
eukaryotes that have multinucleated cells enclosed within cell walls. Fungi obtain nutrition by decomposing dead
and dying organisms and absorbing the decomposition products (NOAA 2010).
gorgonian An octocoral having a horny or calcareous branching skeleton (e.g., sea fans and sea whips).
habitat A place where the physical and biological elements of ecosystems provide a suitable environment including the
food, cover, and space needed for plant and animal livelihood (EPA 2009a).
hardbottom Shallow and deep-water habitats with solid floor that can provide an attachment surface for sessile (nonmoving)
organisms such as corals.
health Health is the general condition of a person in all aspects, including physical and mental. The term health is also
sometimes used to represent the condition of other organisms as well as ecosystems and social structures.
Ecosystem health is an element of overall ecosystem integrity (Campbell 2000). Organism and ecosystem health
usually implies normal functioning of the system and absence of disease as a dominant factor in the system.
Ecosystem health can be thought of as functional integrity. Ecosystems also have structural integrity, which is
related to the presence of all the normally expected elements of the system. Overall ecological integrity is a
combination of the two (i.e., wholeness and normal functioning). For example, a person with only one arm might
be healthy but would not be structurally whole.
herbivore An animal that feeds on plants (EPA 2010).
index A usually dimensionless numeric combination of scores derived from biological measures called metrics (EPA 2000).
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indicator Information based on measured data used to represent a particular attribute, characteristic, or property of a system
(MEA 2009).
indigenous A species is defined as native to a given region or ecosystem if its presence in that region is the result of only
natural processes, with no human intervention. Every natural organism (as opposed to a domesticated organism)
has its own natural range of distribution in which it is regarded as native. Outside this native range, a species may
be introduced by human activity, after which it is referred to as an "introduced species" in such locales.
integrity The extent to which all parts or elements of a system (e.g., an aquatic ecosystem) are present and functioning.
intermediate
ecosystem service
landscape
Components of nature that are not directly enjoyed, consumed or used to yield human well-being, but that are
important for the production of final ecosystem services.
An area of land that contains a mosaic of ecosystems, including human-dominated ecosystems. The term cultural
landscape is often used when referring to landscapes containing significant human populations (MEA 2009).
M
macroalgse Macroscopic, multicellular algas commonly referred to as seaweed.
macroinvertebrate
Animals without backbones of a size large enough to be seen by the unaided eye and that can be retained by a
U.S. Standard No. 30 sieve (28 meshes per inch, 0.595 mm openings) (EPA 2009a).
macrophyte Large aquatic plants that may be rooted or non-rooted, vascular or algiform (such as kelp), including submerged
aquatic vegetation, emergent aquatic vegetation, and floating aquatic vegetation (EPA 2000).
mangrove A general name for several species of halophyte (a plant able to grow in saline conditions) belonging to different
families of plants (including trees, shrubs, a palm tree, and a ground fern) occurring in intertidal zones of tropical
and subtropical sheltered coastlines and exceeding one-half meter in height. The term is applied to both the
individual and the ecosystem (which is termed mangal). Mangroves provide protected nursery areas for juvenile
reef fishes, crustaceans, and mollusks. They also provide a feeding ground for a multitude of marine species.
Many organisms find shelter either in the roots or branches of mangroves. Mangrove branches are nesting areas
for several species of coastal birds. The root systems harbor organisms that trap and cycle nutrients, organic
materials and other important chemicals. Mangroves also contribute to higher water quality by stabilizing bottom
sediments, filtering water, and protecting shorelines from erosion. They protect reefs from land runoff and
sedimentation. Conversely, coral reefs protect mangroves and seagrasses from erosion during heavy storms and
strong wave action. The nations with the largest mangrove areas include Indonesia (with 21% of global
mangroves), Brazil (9%), Australia (7%), Mexico (5%), and Nigeria (5%).The global area of mangroves—
150,000 km —is equivalent to the area of the state of Illinois, or half the area of the Philippines. About one-fifth
of all mangroves are thought to have been lost since 1980, and although loss rates are declining, they are still
three to four times higher than average global forest loss estimates (NOAA 2010).
Marine Protected
Area (MPA)
Any area of the marine environment that has been reserved by federal, state, territorial, tribal or local laws or
regulations to provide lasting protection to part or all of the natural or cultural resources within them. Familiar
examples of U.S. MPAs include national parks, national wildlife refuges, national monuments, national marine
sanctuaries, fisheries closures, critical habitat, habitat areas of particular concern, state parks, conservation areas,
estuarine reserves and preserves, and numerous others. MPAs are sometimes called Marine Managed Areas
(MMA). However, "marine protected area" is a broad, inclusive term that includes both multipurpose sites with
some restrictions as well as the more restrictive "no-take marine reserves" (NOAA 2010).
marine reserve An area in the ocean that is protected from uses that remove animals, plants, and other organisms, or alter their
habitats (NOAA 2010).
marine tourism Includes those recreational activities that involve travel away from one's place of residence and that have as their
host or focus the marine environment (where the marine environment is defined as those waters that are saline
and tide-affected) (Orams 1999).
metabolites A substance that takes part in the process of metabolism, which involves the breakdown of complex organic
constituents of the organism's body with the liberation of energy for use in bodily functioning. The various
compounds that take part in, or are formed by, these reactions are called metabolites (NOAA 2010).
model A physical, mathematical, or logical representation of a system of entities, phenomena, or processes; an
abstracted view of a complex reality.
molecular probe Chemicals that are used to explore and elucidate biochemical structures and processes at the cellular and
molecular levels.
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Montastraea A genus of hard (stony) coral (octocoral) that includes the boulder coral and the great star coral (NOAA 2010).
National Account NAS provide a complete and consistent conceptual framework for measuring the economic activity of a nation.
Systems (NAS) NAS are derived from a wide variety of source data including surveys, administrative and census data, and
regulatory data. Most countries have a national statistical office or central bank that compiles, integrates,
harmonizes, and publishes the data. NAS include a number of aggregate measures (e.g., gross domestic product
[GDP], disposable income, savings, and investment) and other information (e.g., input-output tables that show
how industries interact with each other in the production process).
natural feature A readily observable characteristic of natural systems such as type of vegetation and arrangement of land use
(Wainger & Boyd 2009).
nature and wildlife
watching
The practice of observing nature and wildlife (e.g., birds, dolphins, fish, manatees, turtles, whales) in their natural
habitat.
nonmarket value Value recognized by people but not usually expressed in prices because the thing either is not currently, or cannot
be, traded in markets.
nonuse value The value people hold for a service that they do not directly use. (Sometimes referred to as "passive use value".)
Early literature in environmental economics split nonuse value into three components: existence value, option
value, and bequest value. Nonuse values are theoretically distinct from use values, although the boundary
between use and nonuse values is often fuzzy.
North American The standard used by Federal statistical agencies in classifying business establishments for the purpose of
Industry collecting, analyzing, and publishing statistical data related to the U.S. business economy. NAICS was developed
Classification under the auspices of the Office of Management and Budget (OMB), and adopted in 1997 to replace the Standard
System (NAICS) Industrial Classification (SIC) system. It was developed jointly by the U.S. Economic Classification Policy
Committee (ECPC), Statistics Canada, and Mexico's Institute Nacional de Estadistica y Geografia, to allow for a
high level of comparability in business statistics among the North American countries (U.S. Census Bureau
2010).
nudibranch A marine gastropod mollusk that has no protective covering as an adult. Gills or other projections on the dorsal
surface carry on respiration. They comprise the clade Nudibranchia (formerly a suborder).
nutraceuticals A term that combines the words nutrition and pharmaceutical and is a food or food product that provides health
and medical benefits, including the prevention and treatment of disease (Kalra 2003).
octocoral Aquatic organisms formed of colonial polyps with 8-fold symmetry. They comprise the subclass Octocorallia.
Examples include blue coral, soft corals, and gorgonians (sea fans and sea whips). See also coral.
opportunity cost The cost of something in terms of an opportunity forgone (and the benefits that could be received from that
opportunity), or the most valuable forgone alternative (i.e., the second best alternative).
overfishing Occurs when fishing activities reduce fish stocks below a level that is biologically or economically sustainable.
patch reef Small circular or irregular reefs that arise from the floor of lagoons, behind barrier reefs, or within an atoll.
pharmaceutical Biologically active chemicals used to treat diseases, disorders, and illnesses (NOAA 2010).
preclinical stage Research (sometimes using animal models) to assess whether a candidate drug, procedure, or treatment is likely
of testing to be of therapeutic value in humans. Preclinical studies take place before any testing in humans is done.
presence of reef Quantifies whether or not an offshore reef is present near the coastal area of interest.
protein A large complex molecule made up of one or more chains of amino acids. A typical protein contains 200-300
amino acids, but some are much smaller and some much larger (e.g., titin, a protein found in skeletal muscle,
contains approximately 27,000 amino acids in a single chain). Proteins perform a wide variety of essential
activities in cells, including: (1) they largely form the physical structure of cells and cellular matrices; (2) proteins
are enzymes, which are the catalysts for all biochemical reactions; (3) the transport of materials in body fluids
depends on proteins; (4) the receptors for hormones and other signaling molecules are proteins; (5) motion and
locomotion of cells and organisms depends on contractile proteins; (6) the transcription factors that turn genes on
and off are proteins; and, (7) proteins are an essential nutrient for heterotrophs. The activities of cells and
organisms are largely dependent on the activities of their proteins (NOAA 2010).
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R
provisioning A category of ecosystem services as described by the Millennium Ecosystem Assessment. Provisioning services
services (ecosystem goods) are the products obtained from ecosystems, including: food; fiber; fuel; genetic resources;
biochemicals, natural medicines, and pharmaceuticals; ornamental resources; and, fresh water. In the lexicon of
the ESRP and when quantified appropriately, elements in the MEA provisioning services category could be
considered ecological endpoints. Also see the other MEA categories of cultural services, regulating services,
and supporting services (MEA 2005).
recreation "Refreshment of strength or spirits after work; also: a means of refreshment or diversion" (Merriam Webster
2010). Recreational activities are enjoyed by both tourists and residents of a given geographic location. However,
the common practice of economists is to differentiate between tourism and recreation based upon the source of
demand.
recreational
(sport) fishing
Fishing for pleasure or competition. It is included as a component of tourism and recreation.
reef break A wave that breaks over a coral reef or a rock seabed.
reef continuity The uninterrupted distribution the reef, namely the absence of large gaps or fragmentation due to degradation or
coral mining.
reef crest The sharp break in slope, or peak in reef height at the seaward edge of the reef flat.
reef depth The distance from the ocean surface to the top of the reef; may be an assumed or fixed value in simulation
models, or an average value from field observations for the reef under consideration.
reef distance The distance between the reef crest and the edge of the shoreline; essentially the width of the lagoon.
reef flat The relatively shallow, flat expanse of coral reef between the reef crest and the shoreline.
reef height The distance from the top of the reef to its base; may be an assumed or fixed value in simulation models, or an
average value from field observations for the reef under consideration.
reef roughness The bottom drag coefficient that quantifies friction and may be approximated in field studies by variability in
colony height, or other measures of topography, along the reef flat.
reef slope The angle, from gradual to steep, of the reef front where offshore waves are first encountered; may be an assumed
or fixed value in simulation models, or an average value from field observations for the reef under consideration.
reef type Describes the general structure of the reef and its relationship to the shoreline, including fringing reefs that
border the shoreline, barrier reefs that are separated from shore by a deep lagoon, atoll reefs that form a circular
barrier around an island, and patch reefs that are small, isolated reef outcrops.
reef width The length of the reef flat, the flat expanse of reef from where offshore waves first crest over the reef to the edge
closest to the shoreline; may be an assumed or fixed value in simulation models, or an average value from field
observations for the reef under consideration.
refugia An area or refuge where biota can live and breed without suffering excess predation from other organisms.
regulating A category of ecosystem services as described by the Millennium Ecosystem Assessment. Regulating services are
services the benefits obtained from the regulation of ecosystem processes, including: air quality regulation, climate
regulation, water regulation, erosion regulation, water purification and waste treatment, disease regulation, pest
regulation, pollination, and natural hazard regulation. In the lexicon of the ESRP, elements in the MEA regulating
services category could be considered ecological processes that can produce ecological endpoints, that when
combined with complementary goods and services and demand by humans, could produce ecosystem goods and
services. See also the other MEA categories of cultural services, provisioning services, and supporting
services (MEA 2005).
replacement costs The amount that an entity would have to pay to replace an asset at the present time.
residents People who live at a particular place for a prolonged period (Leeworthy 2002; Princeton WorldNet Glossary
2010a).
resilience The ability of a system to absorb or recover from disturbance and change, while maintaining its functions and
services (Carpenter et al. 2001). For example, a reefs ability to recover from a coral bleaching event.
revealed preference
The use of the recovery of expenditure to "reveal" the preference of a consumer or group of consumers for the
bundle of goods they purchase.
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risk assessment The determination of quantitative or qualitative value of risk related to a concrete situation and a recognized
threat (also called hazard).
rugosity Describes the amount of "wrinkling" or roughness of the reef profile. It is an index of substrate complexity. Areas
of high complexity are likely to provide more cover for reef fishes and more places of attachment for algas.
corals, and various sessile invertebrates (2010).
salient A type of beach morphology in which sediments are deposited and accumulate in the lee of a breakwater
structure, growing seaward from the shoreline to form a bell-shaped structure.
scleractinians Corals that have a hard limestone skeleton and belong to the order Scleractinia. See stony coral.
scuba An apparatus carried by a diver that includes a tank holding a mixture of oxygen and other gases, used for
breathing underwater. Scuba is an acronym for "self-contained underwater breathing apparatus".
seagrass A flowering plant, complete with leaves, a rhizome (an underground, usually horizontally-oriented stem), and a
root system. They are found in marine or estuarine waters. Most seagrass species are located in soft sediments.
However, some species are attached directly to rocks with root hair adhesion. Seagrasses tend to develop
extensive underwater meadows (NOAA 2010).
seascape A mosaic of interconnected coastal and marine ecosystems (coral reefs, seagrass meadows, and mangrove
forests).
secondary A substance produced by an organism that seemingly has no direct role in the organism's metabolism, though
metabolite they are often produced via pathways that are derived from primary metabolic pathways. It is believed that they
are created because they confer some evolutionary advantage, particularly in sessile (nonmoving) organisms.
Most often, secondary metabolites are used by the organism in intraspecies or interspecies interactions usually
related to defense or signaling (NRC 1999; Croteau et al. 2000; Seigler 2002; Wink 2003).
services The benefits that human populations receive from functions that occur in ecosystems.
shoreline The intersection of the land, including man-made waterfront structures, with the water surface. The shoreline
depicted on NOAA National Ocean Service (NOS) maps and charts represents the line of contact between the
land and a selected water elevation. In areas affected by tidal fluctuations, the shoreline is the interpreted mean
high water line. In confined coastal water of diminished tidal influence, the mean water level line may be used. In
nontidal waters, the line represents the land/water interface at the time of survey. In areas where the land is
obscured by marsh grass, cypress or similar marine vegetation, the actual shoreline can not be accurately
represented. Instead, the outer limit line of the vegetation area is delineated (where it would appear to the mariner
as the shoreline), in this case, it is referred to as the apparent shoreline (NOAA 2010).
shoreline protection
The ability of reefs to attenuate offshore wave energy, to provide sheltered nearshore waters, and to protect
coastlines from erosion, flooding, and storm damage.
snorkeling The practice of swimming while equipped with a diving mask, a shaped tube called a snorkel, and fins.
species A category of taxonomic classification, ranking below a genus or subgenus and consisting of related organisms
capable of interbreeding. Also refers to an organism belonging to such a category.
species diversity The number of different species in an area and their relative abundance (NOAA 2010).
species richness The number of species in an area or biological collection (NOAA 2010).
sponge A sessile (nonmoving), multi-cellular marine animal whose body consists of a jelly-like endoskeleton sandwiched
between two layers of cells. Sponges comprise the phylum Porifera.
stony coral Corals (comprising the order Scleractinia) that form hard, calcium carbonate skeletons. Examples include the
brain corals, fungus or mushroom corals, staghorn corals, elkhorn corals, table corals, flower pot corals, bubble
corals, and lettuce corals. These corals are largely responsible for the physical form of coral reefs.
subsistence fishing
Fishing for survival. Fishing for food (consumed by the local group of people who do the fishing), not for
commercial sale.
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supporting A category of ecosystem service as described by the Millennium Ecosystem Assessment. Supporting services are
service those that are necessary for the production of all other ecosystem services. They differ from provisioning
services, regulating services, and cultural services in that their impacts on people are often indirect or occur
over a very long time, whereas changes in the other categories have relatively direct and short-term impacts on
people. (Some services, like erosion regulation, can be categorized as both a supporting service and a regulating
service, depending on the time scale and immediacy of their impact on people.) Examples of supporting services
include: soil formation, photosynthesis, primary production, nutrient cycling, and water cycling. See also the
other MEA categories of cultural services, provisioning services, and regulating services (MEA 2005)
surf break A permanent obstruction such as a reef, headland, bombora, rock, or sandbar that causes waves to break
(Silmalis 2007).
surfing The sport of riding a surfboard toward the shore on the crest of a wave.
sustainability A characteristic or state whereby the needs of the present and local population can be met without compromising
the ability of future generations or populations in other locations to meet their needs (MEA 2009).
tombolo A type of beach morphology in which sediments are deposited and accumulate in the lee of a breakwater
structure, growing seaward from the shoreline until they are connected to the structure.
topography The physical features of a surface area including relative elevations and the position of natural and man-made
(anthropogenic) features.
total economic value
(TEV)
The sum of the change in all relevant use values and nonuse values for ecosystem goods and services produced
by a given change in the ecosystem (i.e., the full social benefits). This is distinct from the "total value" of an
ecosystem, which is the value of the entire system (e.g., the value of an entire wetland), but instead is the value of
a marginal change to that ecosystem that results form some action.
Tourism Satellite A statistical accounting framework in the field of tourism that measures goods and services according to
Account (TSA) international standards for concepts, classifications, and definitions, that allow valid comparisons from country to
country in a consistent manner. A complete TSA contains detailed production accounts of the tourism industry
and their linkages to other industries, employment, capital formation, and additional non-monetary information
on tourism.
tourists People who "travel to and stay in places outside their usual environment for more than twenty-four (24) hours and
not more than one consecutive year for leisure, business and other purposes not related to the exercise of an
activity remunerated from within the place visited" (UNWTO 1995).
travel cost A valuation method that uses the travel time or travel costs as a proxy "total entry fee", and therefore, a person's
willingness-to-pay for visiting a particular tourist location.
tunicate Members of the subphylum Tunicata (also called Urochordata); a group of underwater sac-like filter feeders with
incurrent and excurrent siphons that is classified within the phylum Chordata. While most tunicates live on the
ocean floor and are commonly known as sea squirts (ascidians) and sea pork, others—such as salps, doliolids and
pyrosomes—live above in the pelagic zone as adults.
valuation The process of expressing a value for a particular good or service in a certain context (e.g., decision-making)
usually in terms of something that can be counted, often money, but also through methods and measures from
other disciplines (sociology, ecology) (MEA 2009).
value Generally, the worth, merit, or desirability of something. It can be expressed quantitatively (for example, in
monetary terms) or qualitatively. Specifically with respect to ecological benefits, a quantitative or qualitative
description of those benefits. Using this definition, the value of an ecosystem might be defined in terms of its
beauty, its uniqueness, its irreplaceability, its contribution to life support functions or commercial or recreational
opportunities, or its role in supporting wildlife or reducing environmental or human health risks, or providing
many other services that benefit humans (Ecosystem Valuation 2009).
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w
wetlands A type of ecosystem, generally occurring between upland and deepwater areas, that provides many important
functions including fish and wildlife habitat, flood protection, erosion control, water quality maintenance, and
recreational opportunities. A wetland is an area that is covered by water or has water-saturated soil during a
portion of the growing season. In general, it is often considered the transitional area between permanently wet
and dry environments. The Ramsar Convention on Wetlands identifies the following marine/coastal wetlands:
permanent shallow marine waters; marine subtidal aquatic beds (kelp beds, sea-grass beds, tropical marine
meadows); coral reefs; rocky marine shores (including rocky offshore islands and sea cliffs); sand, shingle or
pebble shores; estuarine waters; intertidal mud, sand or salt flats; intertidal marshes (includes salt marshes, salt
meadows, saltings, raised salt marshes); intertidal forested wetlands (includes mangrove swamps, nipah swamps,
and freshwater tidal brackish and freshwater marshes); coastal brackish/saline lagoons; coastal freshwater
lagoons; and marine and coastal karst and other subterranean hydrological systems (RAMSAR 2001).
willingness-to-accept (WTA) The amount of money (or other goods) that a person must be paid to accept the loss of something else.
willingness-to-pay (WTP) The amount of money (or other goods) that a person is willing to give up to get something else.
zooplankton Free-floating or drifting animals with movements determined by the motion of the water.
zooxanthellse A group of dinoflagellates living endosymbiotically in association with one of a variety of invertebrate groups
(e.g., corals). In corals, they provide carbohydrates through photosynthesis, which are used as one source of
energy for the coral polyps. They also provide coloration for the corals and receive a sheltered habitat in return.
(NOAA2010).
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