Chapter 2
Chapter 2
M. F. M. Fairoz
2.1 Introduction
Coral reefs cover about 1 million square kilometers (km2 ) of Earth’s surface. They
are built by coral organisms producing their own mineral substrates, with thousands
of different species living in coral ecosystems. Coral reefs are comparable with
tropical rain forests in terms of their high levels of biodiversity and complexity of
their ecosystems. They are also among the most productive ecosystems, in terms of
M. F. M. Fairoz (B)
Faculty of Fisheries and Ocean Sciences, Ocean
University of Sri Lanka, Colombo 15 Crow Island,, Sri Lanka
e-mail: fairoz@[Link]
© The Centre for Science and Technology of the Non-aligned and Other Developing 21
Countries (NAM S&T Centre) 2022
E. R. Urban Jr. and V. Ittekkot (eds.), Blue Economy,
[Link]
22 M. F. M. Fairoz
gross organic production (Odum and Odum 1955; Lewis 1977; Birkeland 1983) and
are often an oasis of life in otherwise rather barren tropical marine environments
(Gove et al. 2016). Beside their ecological value, coral reefs are of major economic,
social and cultural importance to the countries they border (Cesar 2000). Almost
half a billion people, 8% of the total global population, live within 100 km of a coral
reef and about 100 developing countries are highly dependent on coral reefs for their
livelihoods.
A large number of studies have estimated the economic value of healthy coral
reefs (e.g., Brander et al. 2012), which come from a variety of renewable resources
that can be harvested (subsistence, commercial and recreational fishing) from reef
areas, and services provided by reefs, including recreational opportunities (e.g., skin
diving and SCUBA, enjoying aesthetic values), tourist development (hotels, vacation
rentals), coastal protection provided by intact reef structures (Sheppard et al. 2005),
and cultural value of reefs to local populations. Brander et al. (2012) compiled 160
separate reef valuation studies and used 45 of these to conduct a meta-analysis of
reef value. The studies included in the analysis used a variety of common valuation
methods. Brander et al. (2012) predicted (through modeling) a loss of 16–27% of
global coral reef area under different IPCC scenarios available at that time. They
estimated an annual loss of $870 billion in 2100 under the IPCC scenario A1 (“a
future world of very rapid economic growth, global population that peaks in mid-
century and declines thereafter, and rapid introduction of new and more efficient
technologies”). However, these figures could be an over-estimate, as they do not
consider the capacity of human populations to adapt, particularly if the ecosystem
changes happen gradually (Hoegh-Guldberg et al. 2019).
This chapter gives a brief overview of reef ecosystems in relation to their role in
blue economies and outlines the latest developments in coral reef monitoring and
research technologies against the background of current scientific research, which is
integral to the successful conservation and management of reef areas.
Fig. 2.1 General structure of the polyp and the underlying skeleton of reef building organism
(Source Lentz 2012)
There have been many attempts to classify different types of reefs. They can be classi-
fied according to their geological history, shape, position relative to land masses, and
by the nature of the material of which they are made. In spite of a great morphological
variety of reef forms, common elements pertaining to the geomorphology such as
buttress system, reef flat, lagoon, patch reef zone, etc. indicate a certain regularity in
their origin and growth on different structural foundations of shelf and island coasts,
across different parts of the coastal zone. This regularity, first recognized by Darwin
(1842), can be distinguished into three basic types of coral reef: (1) fringing reefs, (2)
barrier reefs, and (3) atolls (Fig. 2.2). Fringing reefs are directly attached to volcanic
islands, barrier reefs are separated from volcanic islands or land masses by lagoons,
and atolls are formed by ring reefs which enclose only a lagoon.
Fringing reefs grow in shallow waters and border the coast closely or are sepa-
rated from it by a narrow stretch of water (Fig. 2.2a). They consist of several zones
24 M. F. M. Fairoz
Fig. 2.2 Classification of coral reefs. a Fringing reef. b Barrier Reef. c Atoll (Source Pearson 2016)
that are characterized by their depth, structure of the reef and by plant and animal
communities associated with them. These zones include the reef crest (part of the
reef over which waves break), fore reef (the region of medium energy) and buttress
zone or spur and groove (region of coral growth, which includes rows of corals with
sandy canyons or passages between each row).
Barrier reefs are separated from land by a lagoon; examples are the Great Barrier
Reef off Australia and the Belize Barrier Reef System. These reefs grow parallel to
the coast and are large and continuous. Barrier reefs also include regions of coral
formation that include the zones found in fringing reefs along with patch reefs (small
reefs), back reefs (the shoreward side of the reef), as well as bank reefs (reefs that
occur on deep bottom irregularities) (Fig. 2.2b). Barrier reefs also include reef flats
(the reef not exposed); the reef crest runs parallel to the coast and is protected from
waves and coral terrace (slope of sand with isolated coral peaks). Another coral
terrace and a vertical drop follow these features into deeper waters (Veron 2000).
Atolls are annular reefs that develop at or near the surface of the sea when islands
(usually volcanic) that are surrounded by reefs subside (Fig. 2.2c), and as sea level
changes through glacial cycles (Droxler and Jorry 2021). Darwin (1842) proposed
that atolls as corals continue to colonize sinking volcanic islands that eventually sink
below the sea surface. More recently, Droxler and Jorry (2021) demonstrated that
most modern atolls from the Maldives Archipelago and from the tropical Pacific and
southwest Indian oceans occur on top of late Pliocene flat-topped banks. The volcanic
basement, therefore, has had no influence on the late Quaternary period development
of these flat-topped banks into modern atolls. During the multiple glacial sea-level
low stands that intensified throughout the Quaternary, the tops of these banks were
eroded by dissolution; then, during each of the five mid-to-late Brunhes deglaciations,
coral reoccupied their raised margins and grew vertically, keeping up with sea-level
rise and creating the modern atolls.
Corals belong to the Phylum Cnidaria (Fig. 2.3) and are characterized by the presence
of tentacles, a central digestive cavity and radial symmetry. Although coral reefs are
commonly found in nutrient-poor waters of the tropics, paradoxically, they are among
2 Coral Reefs and Blue Economy 25
the most productive of marine environments (Barnes 1987). Coral reef formation and
maintenance are driven by the symbiosis between the coral animal and single-celled
photosynthetic algae (dinoflagellates) belonging to the genus Symbiodinium that
live within the tissues of stony coral polyps (Levinton 1995). The endosymbiotic
dinoflagellates supply carbon from photosynthesis to the coral host, which in return
provides the algae with nitrogen (Szmant et al. 1990). In addition, coral-associated
bacteria play a key role in the health of the coral (Fairoz et al. 2008; Ziegler et al.
2016). Together this association is referred to as a holobiont (Rohwer et al. 2002)
and the complex interactions among animal, algae and the microbiome intricately
shape coral health and resilience (Bourne et al. 2016; Epstein et al. 2019).
Since the success of this symbiotic relationship relies largely on the photosynthetic
output of the Symbiodinium algae, the distribution and growth of corals is strongly
light-dependent (Levinton 1995), and limits reef growth to the top 100 m of the ocean.
Therefore, coral reefs commonly thrive in clear, nutrient-poor water characteristic
for tropical areas.
It has been estimated that shallow coral reefs occupy less than 0.1% of the world’s
ocean area (Costanza et al. 1997), an area about half the size of Madagascar (Fig. 2.4).
This is less than 1.2% of the world’s continental shelf area (Spalding et al. 2001).
A mass coral bleaching event in 1998 (caused by a major El Niño), resulted in a
decrease of hard coral cover by 8%, but reefs mostly recovered by 2009 (GCRMN
2021). From 2009 to 2018, however, there was a 14% decline in the coral coverage
of shallow-water reef areas globally (Fig. 2.5a), accompanied by an increase in
algal cover of reefs (Fig. 2.5b), due to various natural and anthropogenic processes
26 M. F. M. Fairoz
Fig. 2.4 Distribution of the coral reefs of the world (Source Teh et al. [2013]. Reef areas are
indicated by shading. Attribution 4.0 International [CC BY 4.0])
(GCRMN 2021). The worldwide decline in coral cover is believed to be primarily the
result of recurring coral bleaching events due to increasing sea surface temperatures
(Fig. 2.6), augmented by local outbreaks of coral diseases and crown-of-thorn starfish
(Acanthaster planci), as well as local effects of pollution and destructive fishing
practices. Projected increases in sea surface temperatures with global warming will
undoubtedly lead to continued coral bleaching events and further losses of coral
cover (Schoepf et al. 2015).
The shift in reef areas from coral cover to algal cover (Figs. 2.5a and b), referred
to as a regime shift (Bozec and Mumby 2015), was already noticed more than a
decade ago (Hughes et al. 2007). These will be discussed in Sect. 2.6. Algal over-
growth appears to be most severe in South Asia and the Middle East, where regional-
scale impacts of human activities have prevented coral recovery, even during short
periods of favorable temperatures, particularly when reefs face greater challenges
when exposed to multiple stresses simultaneously.
Fig. 2.5 a Estimated global average cover of hard coral (solid line) and associated 80% (darker
shade) and 95% (lighter shade) uncertainty intervals. b Estimated global average cover of algae
(solid line) and associated 80% (darker shade) and 95% (lighter shade) uncertainty intervals (Source
GCRMN [2021])
2.4.1 Biodiversity
Millions of people living in countries with coral reef ecosystems gain benefits due
to reef’s unique capacity to host diverse groups of marine organisms (Fig. 2.7).
Globally, it is estimated that there are between 600,000 and more than 9 million reef
species (Knowlton 2001). There is evidence that ecosystems with higher biodiversity
are more stable. Marine biodiversity can yield both ecosystem services and products
that benefit blue economies. For example, biodiversity is an important attraction for
marine ecotourism (see Chap. 6 on Tourism) and makes it possible for fishers to catch
a variety of species (see Chap. 4 on Fisheries). The multitude of different species
28 M. F. M. Fairoz
Fig. 2.6 The estimated global average hard coral cover (lighter sold line starting around 33% in
1977) is overlaid with the sea surface temperature (SST) anomaly from 1977 to 2020 (darker line
starting at about 0.25 in 1977). The 80% (darker area) and 95% (lighter area) confidence intervals of
coral cover estimates are shown. Periods of rapid increase in SST anomaly are indicated by asterisk
[*] marks (Source GCRMN [2021])
Table 2.1 Values of coral reef ecosystem services quantified in the literature, from a global
perspective
Statement References
More than 275 million people worldwide live within 30 km of reefs Burke et al. (2011a, b)
(and <10 km from the coast), and ∼850 million people live within
100 km of coral reefs
At least 500 million people rely on coral reefs for food, coastal Wilkinson (2004)
protection, and livelihoods
In developing countries, coral reefs contribute about one-quarter of Moore and Best (2001)
the total fish catch, providing food to an estimated one billion people
in Asia alone
More than 150,000 km of shoreline in 100 countries and territories Burke et al. (2011a, b)
receive some protection from reefs
Globally, coral reefs provide 130 billion USD of flood protection Beck et al. (2018)
from 100-year storm events
Some 30% of the world’s reefs are of value in the tourism sector, with Spalding et al. (2017)
a total value estimated at nearly US$ 36 billion, or over 9% of all
coastal tourism value in the world’s coral reef countries
There are at least 6 million reef fishers in 99 countries worldwide, Teh et al. (2013)
accounting for about 25% of small-scale fishers
2 Coral Reefs and Blue Economy 29
Fig. 2.7 Patterns of biodiversity of the coral reefs (Source Spalding et al. [2001])
produce many different classes of marine natural products that could serve as models
for compounds useful for medicine, agriculture, and other purposes (see Sect. 2.5.3).
Blue carbon ecosystems play an important role in CO2 sequestration and storage
in the form of organic matter, and coral reefs can help protect these systems.
Chapter 3 on Mangroves and Seagrasses provides a more detailed description of
the benefits provided by these ecosystems. Blue carbon has gained much attention in
efforts related to reducing atmospheric CO2 . For example, recent studies have shown
that intact reefs can increase the capacity of seagrass meadows to store carbon by
promoting carbon accumulation and avoiding erosion and export of carbon (Guerra-
Vargas et al. 2020). These economic valuations are considered as “indirect use value”
30 M. F. M. Fairoz
(De Groot et al. 2010) or “hidden value” (Van Beukering et al. 2007; Sarkis et al.
2010) because these services are not visible in daily life.
2.5.1 Tourism
2.5.2 Fisheries
Coral reefs provide habitats for enormous biomass, including finfish and shellfish
resources that are the basis for reef fisheries. A well-managed reef can provide 0.2–
40 tons of seafood per km2 annually (UNEP 2004), with an average of about 5 tons
of seafood per km2 per year. This translates to an annual yield of 1.42 million tons of
seafood from coral reefs worldwide. Coral reef-associated fisheries are a source of
income for many coastal communities, mainly in the developing world (Burke et al.
2011a, b). In terms of nutritional value, reefs supply half of the protein requirement
of the people in some areas, and reef fish are important sources of vitamins A, B, and
D, calcium, iron, and iodine. Across reef nations and territories, people consume an
average of 29 kg of fish and seafood each year (Newton et al. 2007), with consumption
of reef fish being the highest in the Maldives (180 kg/person/year). Reef fish and
shellfish are also harvested for aquaria and curio exports. The economic importance
of coral reef products varies by region. Southeast Asian countries generate $2.4
billion per year in products, and in the Caribbean $395 million per year is derived
from reef-related fisheries (Teh et al. 2013). Reef-associated exports are valued at
more than 1% of total exports in 21 countries and territories, and more than 15%
of total exports in six other countries (Burke et al. 2011a, b). See Chap. 4 for more
information about fishery contributions to blue economies.
2 Coral Reefs and Blue Economy 31
Coral reefs are adapted to thrive in a stable environment and are therefore easily
affected by sudden changes of abiotic factors, such as decreased salinity and pH,
increased temperature, hurricanes and wave action, nutrient pollution, and siltation.
Human threats include overfishing, destructive fishing practices, damage by tourists,
and destruction as part of coastal development. Over the last four decades, catas-
trophic events have affected coral reef systems globally, resulting in their destruction
over vast areas.
A combination of increased seawater temperatures and food availability promote
devastating outbreaks of the predatory crown-of-thorn sea stars that feeds exclusively
on scleractinian corals and has caused extensive damage on the Great Barrier Reef
and in other areas of the tropics (Uthicke et al. 2015).
In addition to such natural threats, various human activates are responsible for reef
degradation. Tourism and recreational activities, and use of explosives (dynamite
fishing), poison and destructive gear for reef fisheries are examples for destructive
human activities. Impact from these activities, coupled with land-based activities such
as shore development, farming, mining, deforestation, reclamation, sewage disposal
exert tremendous stress on the ecological balance of reefs. Segel and Ducklow (1982)
found that most corals are weakened by stress, particularly due to diseases caused
by bacteria as a result of anthropogenic activities (see next paragraph). Changes
in the extent of biotic and abiotic stressors are likely to be further compounded
32 M. F. M. Fairoz
Coral reefs support numerous blue economy activities and global warming is a
challenging and emerging threat to the survival of coral animals. Tropical coral
species generally live close to their upper temperature limits (Berkelmans and van
Oppen 2006) and an increase in sea surface temperature of 0.5 °C can exceed a
species’ heat tolerance, which causes corals to expel their endosymbionts, leading to
a phenomenon known as “coral bleaching”. Because corals depend on algae for the
majority of their energy requirements, prolonged coral bleaching frequently results
in disease, reduced growth and reproduction and often large-scale coral mortality
(LaJeunesse et al. 2018; Hughes et al. 2019; Sully et al. 2019). The mechanisms
related to bleaching and subsequent mortality of different taxa are still not well
understood (Hughes et al. 2018) and thermal tolerance of the symbiotic relationship
can vary greatly between and within species as well as across different geographical
areas (e.g., Cunning et al. 2021).
In order to mitigate the effect of coral bleaching, it is important to understand the
underlying physiological mechanisms responsible for the dissociation between both
symbiotic partners. Although coral reefs may be largely eliminated worldwide by
the end of this century as a result of coral bleaching (Hoegh-Guldberg 1999; Hoegh-
Guldberg et al. 2007; Veron et al. 2009; Bay et al. 2017; Heron et al. 2017), recent
studies also suggest that many species are able to adapt and increase their resilience
to reoccurring heat exposure (Matsuda et al. 2020).
Over the past few decades, there has been a marked rise in coral diseases (Rohwer
et al. 2002) and epizootics of coral disease have resulted in significant losses of
coral cover (Aronson and Precht 2001; Walton et al. 2018), The most notable disease
outbreak was reported from Florida reef tracts from 2014 and it is ongoing to date
(Precht et al. 2016; Walton et al. 2018). Although there is still a lot of uncertainty
surrounding specific pathogens and/or their vectors, disease prevalence has been
linked to an array of environmental disturbances pertaining to water quality such
as nutrient load and sewage (Redding et al. 2013), as well as increases in seawater
temperature (Maynard et al. 2015). Determining the drivers of disease outbreaks is
an ongoing area of vigorous research aiming to disentangle the relative effects of host
resilience and destabilized homeostasis within the holobiont on disease outbreaks
(Burge et al. 2014; Thompson et al. 2014; Peixoto et al. 2017).
2 Coral Reefs and Blue Economy 33
Ocean acidification threatens many ocean ecosystems including coral reefs (see
Chap. 11 for details on ocean acidification and its economic impacts). Economic
impacts of ocean acidification are difficult to predict because the biological effects
of acidification are not yet fully known or predictable. Pelagic organisms that live in
the open ocean and benthic animals that live in shallow habitats or in deep waters
are threatened by ocean acidification, but corals and pteropods appear to be most at
risk (Caldeira and Wickett 2003; Fabry et al. 2008).
In the late 1980s, it was inferred from the geologic record that the chemical
condition of the ocean controls the rate of production of carbonate shells and skeletons
in the marine environment over geologic time scales (Veron 2000). Experiments
performed on tropical corals showed coral growth rates slowing dramatically as
seawater CO2 concentrations increase (Gattuso et al. 1998). Soon after, a ground-
breaking study combined model predictions of future ocean chemistry with results
from laboratory experiments on corals. This study concluded that coral growth would
be severely impeded later this century and that tropical reefs are likely to be most
vulnerable (Kleypas et al. 1999).
The scientific and policy needs for coordinated, worldwide information gathering
on ocean acidification and its ecological impacts, recognized by the United Nations
General Assembly (UNGA 2013), and by many governmental and non-governmental
bodies, resulted in the creation of the Global Ocean Acidification Observing Network
(GOA-ON) (Newton et al. 2015) and inclusion of ocean acidification monitoring as a
goal of Sustainable Development Goal 14. The methods of monitoring and research
for ocean acidification are interdisciplinary, including carbon chemistry, meteo-
rology, oceanography, biogeochemistry, ecology, and biology. Basic data collection
includes temperature, salinity, water depth, oxygen concentration and carbon-system
parameters, plus fluorescence, and irradiance. Benthic status (e.g., coral and coralline
algae cover) is also included in monitoring coral reef habitats to correlate with the
physical and chemical variations in the water column that correspond to those spec-
ified for the Global Coral Reef Monitoring Network (GCRMN), a program of the
International Coral Reef Initiative. These methods will detect ocean acidification as
more acidic conditions erode dead carbonate structures in coral reefs (Campbell et al.
2014; Hoegh-Guldberg et al. 2017).
Several international projects and activities contribute data on ocean and marine
biodiversity parameters, including data related to coral reefs. The following are
among the notable high-level activities:
34 M. F. M. Fairoz
• The Global Ocean Observing System (GOOS) is a platform of the United Nations
Education, Scientific and Cultural Organization’s Intergovernmental Oceano-
graphic Commission, UNESCO-IOC, along with other co-sponsors (Miloslavich
et al. 2018b). GOOS established a new panel in 2015 to extend Essential Ocean
Variables (EOVs) to the biological and ecosystem components of the ocean. The
Biology and Ecosystems Panel identified a set of biological EOVs and is working
on strengthening and developing coordinated observing networks around each of
these (Miloslavich et al. 2018b) in response to identified global needs. See also
Chap. 14.
• The Marine Biodiversity Observation Network (MBON) of the Group on Earth
Observations Biodiversity Observation Network (GEO BON) (Mueller-Karger
et al. 2018) was established to develop a global community of practice for the
collection, curation, analysis, and communication of marine biodiversity data.
This requires coordination and collaboration among countries, organizations and
individuals involved in the Group on Earth Observations (GEO) and many other
organizations.
These groups provided leadership in developing EOVs and Essential Biodiversity
Variables (EBVs) for biological and ecological parameters (Table 2.2) and to promote
the integration of regional datasets shared among scientific communities through
global systems such as the Ocean Biogeographic Information System (OBIS).
Hard coral cover and composition was identified as one of the leading biological
EOVs (see specification sheet at [Link]/eov), partly due to the two
decades of coordinated coral reef monitoring already undertaken by the GCRMN
(e.g., Wilkinson 2000, 2008). The GCRMN has been the unofficial global observing
system for coral reefs since 1997 with the publication of the first global status of reefs
report (Wilkinson 2000) that was motivated by the first global coral bleaching event
of 1997–1998. After that, several reports on coral reefs on global and regional levels,
including research methods, were published. These reports were incorporated into
the Convention on Biological Diversity (CBD) (e.g., [Link] Tittensor
et al. 2014) and global assessments (GRID-Arendal and UNEP 2016).
The GCRMN was established by the International Coral Reef Initiative (ICRI) in
1995, initially with the primary task of reporting on the condition of the world’s
coral reefs in the context of the development of the ICRI “Call to Action” (Dight
and Scherl 1997). Since then, the GCRMN has produced a range of global, regional
and thematic reports on coral reef status and trends, with the support of regional
coordinators and managers of marine parks.
The GCRMN has recognized that coral reefs in more than 100 countries are under
significant pressure from human activities, and are uniquely vulnerable to climate
change, as well as ocean acidification. This makes coral reefs a sensitive indicator
2 Coral Reefs and Blue Economy 35
Table 2.2 Essential Ocean Variables (EOVs) important for monitoring and reporting coral reef
health, and levels proposed to assist in assessing data quality by the Global Coral Reef Monitoring
Network (GCRMN)
EOV name Description and notes Levels 1, 2, and 3*
Hard coral cover and Hard corals are the architects of 1-total hard coral cover (%)
composition coral reefs, justifying this as the 2-cover by functional
most important indicator of coral group/growth form
reef presence and health. 3-cover by genus or species
Three-dimensional reef structure
is strongly determined by the
growth form of coral colonies.
For the GCRMN, the growth
(functional) form and genus-level
identification provide sufficient
detail for monitoring
Fleshy algae cover Different algal groups serve 1-total fleshy algal cover (%)
unique functional roles in reef 2-cover by functional group (turf,
communities. In general, fleshy, fleshy/macro, CCA, and
macro and turf algae are primary calcareous)
competitors to corals for 3-cover by genus or functional
occupying reef substrates; some group, with canopy height
also release dissolved organic
carbon into the water which fuels
microbial activity that inhibits
corals. Crustose coralline algae
(CCA) are key contributors to
reef building. Fleshy algal
cover is the aggregation of fleshy,
macro and turf algal forms
Fish abundance and Fish are highly diverse, occupy a Assigning levels for fish data is
diversity range of functional roles, are premature. The most basic is
mobile, and their size matters. abundance of key taxa, the most
The selection of target fish for complex is biomass of all species
monitoring is challenging and in key families’ status
highly variable. Monitoring
programs are therefore
encouraged to focus on a subset
of fish families and to record all
species within them, though in
many cases target species/taxa are
more manageable
*Data levels rank from Level 1 (minimum requirements) to Level 3 (maximum requirements) See
additional details below from 2000 to 2008
system for coastal ocean health, climate change and ocean acidification impacts, and
their implications for society.
The GCRMN Implementation and Governance Plan (Obura et al. 2019) identifies
four goals for the GCRMN, with subsidiary objectives:
36 M. F. M. Fairoz
• Goal 1: Improve understanding of coral reef status and trends, globally and
regionally.
• Goal 2: Analyze and communicate coral reef biophysical, social and economic
trends, providing science-based recommendations in support of raising awareness,
management and policy development.
• Goal 3: Enable and facilitate greater utilization of coral reef data, including in
research.
• Goal 4: Build human and technical capacity to collect, analyze and report
biophysical and socio-economic data on coral reefs.
Tracking and reporting on coral reef status and trends is needed to understand the
extent and rate of change, and to design appropriate responses. As coral degradation is
taking place at the global level, driven by global as well as local processes, systematic
coral reef observation is required. These observations directly support planning and
tracking coral reef health in relation to sustainable development, climate change
and biodiversity conservation, and have broad application in awareness raising and
outreach. High-quality coral reef data will also support research, including in relation
to ecology and ecosystem service provision, and observational data are needed for
modelling to better predict future reef responses to climate stress.
The role of the GCRMN is to provide these coral reef data, aggregating from
national to regional levels, and then to a global level. The EOVs proposed by the
GCRMN are important for monitoring and reporting coral reef health (Table 2.2).
Greater precision and consistency in how data are measured and reported is a priority
focus for the GCRMN in the coming years. The GCRMN is adopting the Essential
Variable approach (Obura et al. 2019) and a data quality model (Table 2.2) that
enables (a) effective submission of minimum data required to monitor and assess
reef health (hard coral cover and composition), and (b) procedures for strengthening
and extending the data to additional key variables (e.g., algae and fish), and improving
the resolution and “quality” of each variable.
The data quality model scores three levels of data, from minimum requirements
(Level 1) to maximum (Level 3), providing specific guidance on how to improve data
quality from Level 1 to 3, and incentivizing monitoring teams to improve quality. The
key method for ensuring data quality is the preparation and submission of appropriate
metadata (GCRMN 2018a, b) that include information on sites and sampling, variable
precision and replication, and any data processing or transformations applied after
data collection. This approach enables two objectives: first, to maximize submission
of data that meet a minimum standard from all parts of the globe, and second, to
provide a pathway for capacity building to improve data quality to the highest levels
possible.
2 Coral Reefs and Blue Economy 37
For developing countries, there are specific needs related to observational monitoring
and ecosystem-based management approaches at local levels (Obura et al. 2019).
Awareness and access to available new technologies and capacity building are among
some of the priorities, including for personnel at the science-policy interface. These
aspects are discussed in detail in Chaps. 14 (Ocean Observations) and 15 (Capacity
Development). This section presents a description of the science and technology
options to better manage reef ecosystems in order to avoid threats to sustainable blue
economies and discusses some of the latest studies on improving our understanding
of coral reefs.
A large body of literature shows that the major threat to coral reefs globally is
the series of unprecedented coral bleaching incidents that have occurred in various
parts of the world. The major cause of this coral bleaching is global warming, caused
by increased sea surface temperature. Research on coral physiology and adaptive
capacities may help in devising approaches to mitigate the impacts of global warming.
Stony corals owe their success as reef builders to their symbiosis with endosymbiotic
dinoflagellate algae of the genus Symbiodinium. These algae live in coral tissues in
extremely high densities (greater than 106 cm−2 ) and provide up to 90% of a coral’s
nutritional requirements (Muscatine and Porter 1977). Researchers have studied the
connections between loss of endosymbionts, the global incidence of coral bleaching,
and the loss of coral cover in recent decades. The effects of thermal stress on corals
over the last century (Lough 2000) yield clues to the future of coral reefs in the
context of global warming in the future (IPCC 2021).
Coral-Symbiodinium symbiosis is temperature sensitive; increased sea surface
temperature can disrupt coral symbiotic complexes to cause coral bleaching resulting
from the expulsion of symbiotic algae from the coral tissues due to the thermal stress.
It is sometimes possible for the algae to recolonize coral tissues after the temperature
returns to normal. This situation is called temporary bleaching and corals can recover
within a couple of days. However, if elevated temperatures last for a longer period,
bleaching can become permanent and there is no chance for the coral to maintain the
symbiotic system (Muscatine 1990).
Recent research to understand coral bleaching and to improve knowledge of the
effects of temperature on the breakdown of coral symbiosis also includes microbial
ecology, to integrate into a larger explanation of status of the reef coral health (Dove
and Hoegh-Guldberg 2006), as explained in the following section.
38 M. F. M. Fairoz
Fig. 2.8 The coral holobiont consists of symbiotic zooxanthellae, the coral animal and associated
viruses, microbes (both bacteria and archaea), and endolithic organisms. Most of the viruses are
bacteriophages, which attack the microbes. The endolithic organisms include algae, fungi, sponges,
and microbes that bore into the coral skeleton. The holobiont likely occupies different coral reef
niches by mixing and matching different components (i.e., the Probiotic Hypothesis). Zooxanthellae
and protist micrographs by Linda Wegley and Ian Hewson, respectively (Source Rosenberg et al.
2007; Attribution 4.0 International [CC BY 4.0])
2 Coral Reefs and Blue Economy 39
holobiont. Microbes included in the coral holobiont are also described as “benefi-
cial microorganisms for corals” (BMCs) (Santoro et al. 2021). These coral-associated
beneficial microbes generally fix atmospheric nitrogen, maintain sulfur cycling, scav-
enge reactive oxygen species (ROS) that otherwise harm coral tissues, and produce
antibiotics to thwart pathogens (Robbins et al. 2019). The functional changes and
host association of BMCs are largely unknown.
Knowledge gained from coral reef microbial ecology studies has been applied to
explain the large-scale ecological status of coral reefs. Dinsdale et al. (2008) charac-
terized microbial communities from pristine to human-populated atolls in the Pacific
Ocean, including four coral atolls in the Northern Line Islands located in the central
Pacific Ocean. The differences in microbial communities across atolls reflected the
variation in the oceanographic conditions and direct and indirect human impacts,
such as fishing and land use (Dinsdale et al. 2008; Sandin et al. 2008). Data from this
study could serve as a baseline for future studies on reef microbes and the associa-
tion of microbes on degradation of coral reef systems worldwide. Techniques used
in this study represent a combination of methods from metagenomics, microscopy,
microbial culture methods, and water chemistry.
Another study was conducted of coral reefs at 11 Line Islands in the central Pacific
Ocean that together span a wide range of biogeochemical and anthropogenic influ-
ences to examine coral microbiomes (Kelly et al. 2014). The percent cover of major
benthic functional groups were significantly correlated with particular microbial
taxa. Reefs with higher coral cover had a coral microbiome with higher abundances
of Alphaproteobacteria (such as Rhodobacterales and Sphingomonadales), whereas
microbiomes of algae-dominated reefs had higher abundances of Gammaproteobac-
teria (such as Alteromonadales, Pseudomonadales, and Vibrionales), Betaproteobac-
teria, and Bacteriodetes. In contrast to taxa, geography was the strongest predictor
of microbial community metabolism. Microbial communities on reefs with higher
nutrient availability (e.g., equatorial upwelling zones) were enriched in genes
involved in nutrient-related metabolisms (e.g., nitrate and nitrite ammonification,
total nitrogen transport). On reefs further from the equator with lower nutrient levels,
microbes had more genes encoding chlorophyll biosynthesis and photosystems I/II.
Another approach to coral reef research has involved the application of metage-
nomics in coral reef studies to obtain a comprehensive view of microbial ecology
of coral reefs. Metagenomics is the study of genetic material recovered directly
from environmental samples, such as seawater. This is a broad field also referred
to as environmental genomics, ecogenomics and community genomics. A detailed
40 M. F. M. Fairoz
metagenomics study from Japanese reefs (Meirelles et al. 2018) suggest that syner-
gistic effects among several regional stressors are driving coral decline. In a highly
hydrodynamic reef environment, high algal/turf cover, stimulated by eutrophication
and low fish abundance due to overfishing, promote water column microbial activity,
together with crown-of-thorns sea star outbreaks and climate changes impacts; these
coral reefs are likely to collapse.
Global coral reef status shows that coral-dominated reefs are becoming algal-
dominant reefs worldwide, recognized as a “phase shift” (Hass et al. 2006). This
shift in coral ecosystems was explained as “global microbialization” of coral reefs,
an observed shift in ecosystem trophic structure towards higher microbial biomass
and energy use. On coral reefs, the primary causes of microbialization are overfishing
of herbivorous fish and eutrophication, both of which enhance the growth of fleshy
algae, allowing it to out-compete calcifying corals and coralline algae. Haas et al.
(2016) propose that the “DDAM (Dissolved organic carbon [DOC], Disease, Algae,
Microorganism) positive feedback loop” gives fleshy algae a competitive advantage.
The DDAM feedback loop involves ungrazed fleshy algae releasing DOC, supporting
pathogenic bacteria that harm corals. Observations from three ocean basins were used
to demonstrate the prevalence of DDAM processes. Besides promoting growth of
fleshy algae, microbialization of reefs can also increase hypoxia and decrease pH
from the microbial respiration of DOC. Following the explanation of microbialization
in coral reefs, interest has increased regarding microbial metabolism and the micro-
biome over reefs and its connectivity to local environmental factors. Microbiome
studies explain the maintenance and adaptation of microbiomes to local conditions,
facilitated by the horizontal transfer of genes responsible for specific metabolic capa-
bilities from one microbial species to another. The core microbiomes are determined
by holobiont macroorganisms as explained above in this chapter.
Global-scale coral bleaching risk alerts have been generated for the past 20 years
based on satellite observations of sea surface temperature (SST), through detec-
tion and mapping of high positive SST anomalies relative to long-term averages.
Bleaching alert products were developed by the U.S. National Oceanic and Atmo-
spheric Administration (NOAA) Coral Reef Watch Program ([Link]
[Link]/satellite/; Liu et al. 2014). Other regional forecast models of coral stress
due to heat, extreme cold winter anomalies, and coral disease have been imple-
mented, such as in Australia (Maynard et al. 2011). These models, combined with
volunteer-based ground-truth monitoring networks (e.g., Gudka and Obura 2017),
help generate awareness of the threat of coral bleaching, and build interest and
2 Coral Reefs and Blue Economy 41
capacity in management responses to mitigate the effects of these events and promote
resilience and recovery from the events (Marshall and Schuttenberg 2006). A typical
major bleaching event occurs over 2–4 months, and monitoring an event requires
different approaches at its start, during the event, and afterward to track long-term
impacts. Improved quantitative data helps to target management and policy responses
more effectively.
Fixed buoys for monitoring aerial and subsurface biophysical properties of coral
reef waters are used for monitoring in some locations, notably the NOAA Coral
Reef Early Warning System (CREWS). Starting in 1997, this system combined soft-
ware and sensor platform development to automatically detect conditions thought
conducive to coral bleaching (Hendee et al. 2007). Buoys in this network of stations
around the Caribbean and in Saipan (in the Pacific Ocean) relay data by satellite
to the Atlantic Oceanographic and Meteorological Laboratory (AOML) in Miami,
Florida, USA.
Fig. 2.9 Application of Remotly sensed measurements to coral reef restoration examples by the
Carnegie Airborne Observatory [Asner et al. 2012]. Symbols are courtesy of the Integration and
Application Network, University of Maryland Center for Environmental Science ([Link]/
symbols/) (Source Foo and Asner [2019] Attribution 4.0 International [CC BY 4.0])
their use for mapping relies on input of covariate biophysical data. Feature extrac-
tion provides an alternative application of machine-learning algorithms to coral reef
imagery.
Remotely sensed measurements can be useful for coral reef restoration activities
(Fig. 2.9). Key criteria considered by restoration practitioners are paired with the
current platform and sensing technology that is most mature and developed, for the
criterion at the required resolution to inform fragment collection, coral out-planting,
and monitoring efforts.
Discrete features visible within images can be directly detected as recurring patterns
across multiple pixels. This is commonly achieved through a family of multi-layered
deep-learning algorithms known as convolutional neural networks (CNN) that were
initially developed for facial and handwriting recognition. These algorithms define a
series of mathematical convolutions to generate the output from the input, allowing
all instances of a feature falling within an image to be located based on a predefined
set of features (LeCun et al. 2015). On sand cay islands and reef flats, features that
are amenable to detection include fallen trees, particularly on narrow beaches, which
signify erosion of sand cay shorelines (Lowe et al. 2019) and individual mangrove
2 Coral Reefs and Blue Economy 43
Fig. 2.10 Example of mapping possible by drone, showing a composite picture composed from
1400 UAV images of Nymph Island on Australia’s Great Barrier Reef. Panel A shows the photo-
mosaic, which Panel C that the detail available can show individual corals (Source Hamylton et al.
[2020] Attribution 4.0 International [CC BY 4.0])
trees, from which forest expansion and contraction can be monitored over time
(Hamylton et al. 2020). Individual corals are now clearly visible in drone images
acquired above reefs (Fig. 2.10) and machine-learning algorithms hold good poten-
tial for detecting reef features, opening up a wealth of potential management applica-
tions for monitoring coral restoration and bleaching. For example, on degraded reefs,
restoration activities include planting tens of thousands of juvenile corals (Lirman
and Schopmeyer 2016), whose planting sites can be optimized by analysis of existing
coral distributions.
As pointed out in Chap. 11 on Ocean Acidification, managers of coral reef areas can
do little to slow the global increase in ocean acidification due to increasing CO2 in the
atmosphere. However, there are some potential means of mitigating ocean acidifica-
tion at local levels, or perhaps repair damage to coral reef ecosystems due to ocean
acidification and other stresses. Albright and Cooley (2019) described a variety of
passive and active approaches to help protect coral reefs or to help them recover if
damaged. Passive measures are those that reduce controllable stresses on coral reefs
and increase the resilience of reef areas. Examples of passive approaches include
more conservative management of fisheries and coral reef areas to make them more
resilient, implementation of marine spatial planning to separate conflicting uses from
reef areas, protecting reef areas as part of marine protected areas, factoring reef health
and sustainability into national blue economy plans (Andersson et al. 2019), and
improving wastewater treatment to decrease eutrophication and other coastal pollu-
tion (Guan et al. 2020). Active measures to reef protection and mitigation mentioned
by Albright and Cooley (2019) include using aquatic plants such as seagrasses to
remove CO2 from seawater, chemical balancing of pH in reef areas, reef restoration
from coral nurseries, and “assisted evolution” for both corals and the dinoflagellates
they host. Application of these passive and active approaches will require significant
research, technology transfer, and capacity development.
world. Arks will create an entirely new way to study coral reefs and allow for the
continued discovery of new marine natural products. Arks could serve as midwater
nurseries and reservoirs for restoration projects, providing flora and fauna to restore
reef communities following damage or degradation, and critically preserving coral
reefs of the world.
2.9 Conclusions
Coral reef ecosystems provide critical services and products that are foundations of
several conventional (tourism, fisheries) and emerging blue economic sectors. Intact
reef ecosystems also provide protection against or reduce threats from natural hazards
and extreme events. Climate change and other human activities have led to a decline
in coral reefs globally, limiting their ability to provide products and services. This
decline in live coral cover over the last two decades has been due to coral bleaching,
coral diseases, and mechanical destruction from uncontrolled tourism and destructive
fishing methods. In many regions, there have been regime shifts due to impacts of
complex natural and anthropogenic factors still under investigation, where coral cover
is being replaced by algal cover. There is strong evidence-based research on mech-
anisms explaining why corals die. However, research and monitoring are needed
to continue to better understand and observe coral ecosystem dynamics and reef
behavior in support of ecosystem-based management of resources. New scientific
understanding could lead to new methods of conservation and launch coral restora-
tion initiatives. Shallow-water corals are mostly bordered by developing countries
in the tropics with specific local needs for observational monitoring and manage-
ment approaches. Awareness and access to available new technologies and capacity
building, including for personnel at the science-policy interface, are among some of
the priorities.
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