18 - Seagrass Ecosystems Their Global Status and Prospects Pp. 281-294

Cambridge Books Online
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Aquatic Ecosystems
Trends and Global Prospects
Edited by Nicholas V. C. Polunin
Book DOI: http://dx.doi.org/10.1017/CBO9780511751790
Online ISBN: 9780511751790
Hardback ISBN: 9780521833271
Chapter
18 - Seagrass ecosystems: their global status and prospects pp. 281-294
Chapter DOI: http://dx.doi.org/10.1017/CBO9780511751790.025
Cambridge University Press
18 Seagrass ecosystems: their global status and prospects
c a r l o s m. d ua rt e , j e n s b o r u m, f r e d e r i c k t. s h o rt a n d d i a na i . wa l k e r
INTRODUCTION
Seagrasses form important underwater marine and estu-
arine ecosystems on all continents except Antarctica.
Seagrass ecosystems are highly productive, forming
extensive habitats which support highly diverse commu-
nities. The seagrasses themselves assimilate and cycle
nutrients and other chemicals. Their extensive above- and
below-ground biomass traps sediments, reducing coastal
turbidity and erosion, as well as providing habitat for other
organisms both attached and free-living (Hemminga &
Duarte 2000).
Globally, seagrasses are in decline, almost entirely
because of human impact. Causes range from changes in
light attenuation due to sedimentation and/or nutrient
pollution, to direct damage and climate change. Over the
last two decades, the loss of seagrass from direct and
indirect human impacts amounts to 18% of the docu-
mented seagrass area (Green & Short 2003). Seagrass
Aquatic Ecosystems, ed. N. V. C. Polunin. Published by Cambridge University Press. Foundation for Environmental Conservation 2008.
281
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ecosystems are in need of active management to ensure
their persistence and long-term survival, requiring edu-
cation, increased awareness, management and conservation
on a global scale.
Seagrass meadows are coastal benthic ecosystems
found submerged in nearshore waters; the group of plants
collectively known as seagrasses consists of about 60
similarly functioning angiosperm species (den Hartog
1970; Hemminga & Duarte 2000). These are all rhizo-
matous monocotyledons, within two families (Potamoge-
tonaceae and Hydrocharitaceae) encompassing 12 genera,
which are restricted to the marine environment to com-
plete their life cycle. Seagrasses are owering plants that
pollinate on or under the waters surface and produce
fruits and seeds which are important to maintaining the
habitat, although their primary mode of expansion is
clonal growth. Seagrasses are rooted in shallow sandy to
muddy coastal sediments receiving at least 1020 % of
irradiance incident at the water surface (Hemminga &
Duarte 2000; Green & Short 2003). A few species (such
as genera Phyllospadix, Posidonia and Thalassodendron) can
grow on rocks or very thin sediments (den Hartog 1970;
Hemminga & Duarte 2000). Seagrasses thrive in a range
of salinities, from 5 PSU (practical salinity units), or
even lower, to 60 PSU (McMillan & Moseley 1967;
Walker 1989).
Where they occur, seagrasses represent the dominant
component of the shallow marine landscape. Most sea-
grass meadows are monospecic, although tropical and
subtropical meadows may contain up to a dozen species,
particularly in the Indo-Pacic region, which contains the
most diverse seagrass ora (Fig. 18.1). Seagrass meadows
form lush ecosystems and, on an area basis, are some of
the most productive communities on the planet with an
average total seagrass biomass (dry) of 460 g per m
2
, half
of which develops below the sediment as rhizome and
root material, and an average net primary production
(dry) of 5 g per m
2
per day (Duarte & Chiscano 1999).
The primary production of seagrass meadows is aug-
mented by the contribution of micro- and macroepiphytic
and benthic autotrophs, which contribute about as much
to ecosystem production as the seagrass itself (Hemminga
& Duarte 2000). Seagrass habitat interacts with other
critical coastal habitats, namely saltmarshes (Chapter 11)
and bivalve reefs in temperate regions, and mangrove
forests (Chapter 12) and coral reefs (Chapter 16) in the
tropics.
Mapping the extent of the global seagrass ecosystem
is far from complete, the most recent estimate of the global
seagrass area being 177 000 km
2
(Green & Short 2003), but
this is without doubt much too low. In most countries, no
generalized spatial mapping of seagrass distribution has
been conducted and seagrass locations are only known
from localized observations. In many areas of the globe,
seagrass observations have not been made. Over the past
decade, increased investigation has led to the discovery of
seagrass distribution in areas where it was previously
unknown (for example the deep-water seagrass beds in the
Gulf of Mexico: Hammerstrom et al. 2006), and also to
documentation of absence of seagrass where it might be
expected. For instance, seagrasses are often absent from
coastal waters in the vicinity of cities and towns both in
16
14
12
10
8
6
4
2
0
80 60 40 20 0 20 40 60 80
N
Latitude S
N
u
m
b
e
r

o
f

s
p
e
c
i
e
s
Equator
Fig. 18.1. Global species richness of seagrasses by latitude. (Based on Green & Short 2003.)
282 C. M. DUARTE ET AL.
developed and developing countries. More seagrass map-
ping and monitoring are needed to document the status
and trends of seagrass populations. Current knowledge
suggests that seagrasses are being lost at a greater rate than
is being documented.
Seagrasses are important as habitats supporting high
biodiversity, yet they occur at the landsea interface and
are particularly vulnerable to anthropogenic impact. Their
lush canopies provide habitat to a rich faunal assemblage,
which uses seagrass meadows as nursery and recruitment
grounds (Thayer et al. 1975; Green & Short 2003). The
seagrass fauna encompasses various trophic guilds,
including large herbivores in both tropical (for example
dugongs, manatees and sea turtles) and temperate (for
example swans and geese) waters, and invertebrates. Sea-
grasses are also food for sea urchins and sh (Pollard 1984;
Heck & Valentine 1995). Worldwide, seagrasses provide
crucial links in the food web for animals and people,
including subsistence gleaning for protein on tropical reef
ats by villagers, nursery resources for commercially
important nsh species and habitat for commercial and
recreational bivalve sheries.
Seagrasses develop dense canopies that impede water
movement, damping current and wave energy (Grizzle
et al. 1996) and turbulence, reducing sediment resuspen-
sion (Short & Short 1984). Seagrass meadows thus increase
sedimentation rates (Gacia & Duarte 2001; Agawin &
Duarte 2002), acting as a lter of coastal waters. The high
total organic inputs from both the water and the plants fuel
sediment microbial activity, which is typically enhanced in
seagrasses relative to adjacent sediments that are bare of
vegetation (Hemminga & Duarte 2000; Holmer et al.
2001). Calcifying organisms such as coralline algae, mol-
luscs and Foraminifera are important components of sea-
grass meadows, contributing substantially to the formation
of carbonate sediments (Walker & Woelkerling 1988).
Seagrass meadows provide important services such as
nutrient and gas cycling, through the intense biogeo-
chemical processes within the meadows; habitat provision
for a diverse assemblage of plants and animals, often of
commercial value; and erosion control, through the dissi-
pation of wave energy, the stabilization of underlying
sediments and the delivery of important amounts of
organic and inorganic material to other parts of the marine
environment (see Hemminga & Duarte 2000). Seagrass
meadows rank amongst the ecosystems giving greatest
added value in terms of the services they provide to marine
ecosystem processes and ultimately to society (Costanza
et al. 1997), which should inspire conservation efforts to
preserve them.
Seagrass meadows are and have been highly dynamic
ecosystems. Because of the narrow fringe that seagrass
meadows occupy in the coastal zone, these ecosystems
have experienced major, undocumented regression and
progression with sea-level change through geological time.
Although recent rates of sea-level change have been in the
order of, at most, a few millimetres per year, these changes
may translate to signicant rates of horizontal inundation
or desiccation with increasing or decreasing sea level,
respectively. However, seagrasses cannot keep up with
human-imposed change as it is now happening globally.
While seagrasses are recognized as priority subjects
for conservation efforts in international (such as the Con-
vention on Biological Diversity and the EU Habitats
Directive) and national frameworks, there is evidence of
signicant widespread decline (Short & Wyllie-Echeverr a
1996; Hemminga & Duarte 2000; Green & Short 2003;
Spalding et al. 2003). These declines have multiple and
sometimes synergistic sources, many of which are related
to direct and indirect anthropogenic pressures including
climate change (Short & Neckles 1999; Duarte 2002). As
many of these pressures are expected to continue and even
increase over the coming decades, there is a need to
examine the present status and likely prospects for seagrass
meadows globally, thereby helping to inspire effective
conservation efforts. In this chapter, the current status of
seagrass worldwide is examined, including what is known
to date of seagrass ecosystem distribution. Threats to
seagrass are discussed that are largely human-induced but
sometimes natural, as well as the measures that can be
adopted to control, or ideally reverse, seagrass decline.
Finally, the prospects for seagrasses to the 2025 time
horizon are considered.
SEAGRASS STATUS
The extent of the seagrass habitat is documented for only a
fraction of the worlds coastline (Green & Short 2003).
Major losses of seagrass have been documented in Europe
(for example De Jonge & De Jong 1992; Marba` et al. 1996;
Hily et al. 2003), in the developed parts of the USA (for
example Orth & Moore 1983; Koch & Orth 2003; Short &
Short 2003), as well as in Australia (Walker & McComb
1992) and in localized areas of the developing world
(Spalding et al. 2003). The loss of seagrass has in some
locations led to loss of sheries, changes in bottom
Seagrass ecosystems 283
substrata and breakdown of ecosystem structure. World-
wide, vast healthy seagrass meadows still exist in areas not
impacted by human activity. The magnitude of seagrass
loss has been greatest in developed countries, but the
current rate of loss is highest in the developing world.
Regional seagrass status and trends
AFRI CA
Very little is known about seagrasses in West Africa, with
most of the reports pertaining to Cymodocea nodosa and
Zostera noltii in the Canary Islands and on the Mauritanian
coast (Van Lent et al. 1991; Vermaat et al. 1993; Reyes
et al. 1995). In East Africa, there is high species diversity in
mixed species stands along the central coast (Bandiera &
Gell 2003; Ochieng & Erftemeijer 2003), which are heavily
used by humans for gleaning and artisanal sheries, both of
which result in signicant loss of seagrasses. In many areas,
seagrasses grow adjacent to mangrove forests (Chapter 12).
The main human impacts on seagrass result from sedi-
mentation following watershed clearing and development
as well as direct human activities in the coastal zone, but
losses have not been measured.
MI DDLE EAST
In the Red Sea, there are 11 seagrass species, often growing
in mixed beds (Lipkin et al. 2003). One species (Halophila
stipulacea) can penetrate to great depths (70 m). Nutrient
pollution and coastal development are impacting seagrass
beds in the Red Sea. In the Arabian Gulf, there are
extensive seagrass beds supporting large dugong popula-
tions, but also more direct industrial impacts, because of oil
extraction, than in other parts of the region (Phillips 2003).
I NDI A
Many seagrass species are found extensively around the
subcontinent of India and its islands, although little infor-
mation exists on their spatial distribution ( Jagtap et al.
2003). Upland deforestation, mangrove destruction, shore-
line construction and storms have all had important impacts
on seagrasses. Dugong populations were recently eliminated
in India after a 50-year period of decline. Large human
population pressures along the coast continue to threaten
seagrasses through vessel trafc and sewage disposal.
AUSTRALI A
Australia hosts the highest number of seagrass species of
any land mass in the world (about 30 species: Walker 2003;
Coles et al. 2003a), with large multispecies meadows
across vast shallow areas of the coastal fringe. The major
threats to seagrass are industrial development, nutrient
loading near population centres, port development, coastal
agriculture and sheries. However, seagrasses are better
protected by legislation and enforcement in Australia
than anywhere else, and because of this and small
human populations, seagrass losses are lower than on other
continents.
SOUTH- EAST ASI A
South-East Asia is the global centre of biodiversity for
seagrasses, as it is for coral reefs and other biota (Bujang &
Zakeria 2003; Kuriandewa et al. 2003; Supanwanid &
Lewmanomont 2003). There are extensive seagrass beds
throughout the area, but very few have been mapped.
Mining of metals and sand, as well as watershed clearing
for agriculture, aquaculture and industry, all negatively
impact seagrasses (see Fortes 1988; Terrados et al. 1998).
EAST ASI A
Seagrass beds in the countries of temperate Asia ( Japan,
Korea, China and northern Vietnam) are dominated by
Zostera spp. which occupy much of the coastal soft sub-
strate (Aioi & Nakaoka 2003; Lee & Lee 2003). No actual
distribution maps are available. Seagrasses have suffered
severe losses in industrialized parts of the coast, in many
cases from hardening of the shoreline and land reclamation
through lling of tidal areas for industrial development.
Direct sheries impacts also contribute to seagrass loss.
PACI FI C I SLANDS
The many islands of Micronesia, Melanesia and Polynesia
are typically surrounded by coral reef ats with extensive
seagrass habitat and high seagrass biodiversity (Coles et al.
2003b). However, the extent of seagrasses across the region
is poorly documented. The greatest impacts come from
coastal development, artisanal shing and gleaning on the
reef ats, and tourism infrastructure development.
NORTH AMERI CA
Seagrass is found extensively along the coast except in
high-population areas where the habitat has been lost
owing to development and pollution discharge (Wyllie-
Echeverria & Ackerman 2003; Short & Short 2003; Koch &
Orth 2003). The dominant North American seagrasses are
eelgrass (Zostera marina) in temperate waters and turtle-
grass (Thalassia testudinum), accompanied by additional
species, in subtropical areas (Onuf et al. 2003). Eelgrass
experienced a severe die-off during the 1930s on the east
coast of North America and in Europe; although it
recovered in many places, human impacts have prevented
it from re-establishment in parts of its range. In the USA,
seagrass distribution is well documented; major losses have
been identied on both coasts and in the Gulf of Mexico.
Canada and Central America have experienced fewer
declines but are faced with threats owing to increased
coastal development.
CENTRAL AND SOUTH AMERI CA
Seagrasses in the Caribbean are well documented but
only sporadically mapped and are facing threats particu-
larly from agricultural runoff, tourism and urban develop-
ment (Creed et al. 2003). Very little is known about
seagrasses in the rest of South America (Creed 2003). No
one has yet investigated whether seagrass occurs for long
stretches of the South American coast. Brazil has several
tropical species along its coastline, and seagrasses there
experience threats from urban development and watershed
deforestation.
EUROPE
The Mediterranean Sea is dominated by Posidonia ocea-
nica, a reef-forming seagrass that persists over centuries
(Procaccini et al. 2003). Heavy urbanization has impacted
seagrasses, such as through eutrophication, organic inputs
from aquaculture and coastal engineering, and 3040% of
P. oceanica in the western Mediterranean has been lost in
the last few decades. In northern Europe (Hily et al. 2003;
Bostrom et al. 2003), where Zostera species dominate,
losses of seagrass have also been extensive and include
indirect impacts from nutrient pollution and increased
turbidity as well as direct impacts from coastal modica-
tion, shing, boating and aquaculture. For example, in
the Baltic Sea, shifts in eelgrass distribution from deeper
to shallower water reect loss of water clarity over time
(Fig. 18.2).
Global impacts and losses
Globally, the estimated loss of seagrass from direct and
indirect human impacts amounts to 33 000 km
2
, or 18% of
the documented seagrass area, over the last two decades
1900 1900
1997
1997
30
20
10
0
40
30
20
10
0
30
20
10
0
30
20
10
0
2 4 6 8 10 12 14 16 2 4 6 8 10 12 14 16
Colonization depth (m)
F
r
e
q
u
e
n
c
y

o
f

m
e
a
d
o
w
s

(
%
)
Open coasts Estuaries
Fig. 18.2. Colonization depth of Danish eelgrass meadows along open coasts and in estuaries, 190097. (Modied from Bostrom
et al. 2003.)
(Green & Short 2003), based on an extrapolation of known
losses (Short & Wyllie-Echeverria 2000). Reported losses
probably represent a small fraction of those that have
occurred and many losses may remain unreported; indeed
actual losses may never be known because most seagrasses
leave no long-term record of their existence. Seagrasses
exist at the landsea margin and are highly vulnerable to
pressures from human populations, which live dis-
proportionately along the coasts (Nicholls & Small 2002).
Human population growth, with concomitant increased
pollution, hardening and alteration of coastlines and
watershed clearing, threatens seagrass ecosystems and has
resulted in substantial and accelerating seagrass loss over
the last 20 years (Hemminga & Duarte 2000; Short &
Wyllie-Echeverria 2000; Spalding et al. 2003).
The likely primary cause of seagrass loss globally is
reduction in coastal water clarity, both from increased
nutrient loading and increased turbidity. The primary
cause of nutrient enrichment in coastal waters is anthro-
pogenic loading from coastal watersheds (Valiela et al.
1992; DAvanzo & Kremer 1994; Duarte 1995; Short &
Burdick 1996; Short & Wyllie-Echeverria 1996; Tomasko
et al. 1996; Borum 1996; McMahon & Walker 1998). In
general, pristine coastal seas are nitrogen limited, and
nitrogen inputs from point and non-point sources cause
eutrophication (Ryther & Dunstan 1971; Nixon & Pilson
1983) and alter ecosystem structure and function (Valiela
et al. 1992; Borum 1996). Increased nutrient inputs are also
occurring adjacent to industrialized regions of the world
through direct atmospheric deposition of nitrogen (Paerl
1985). Equally, losses of water clarity come from increasing
inputs of nitrogen and phosphorus from waste discharge,
atmospheric deposition and land runoff along temperate
more-industrialized coasts.
In contrast, in tropical areas, the major impact on water
clarity is the discharge of vast quantities of sediment into
coastal waters as a result of poor land-use practices, watershed
deforestation and coastal clearing (leading to erosion), and
lack of erosion controls (Fortes 1988; Duarte et al. 1997; Bach
et al. 1998; Terrados et al. 1998). The relatively high light
requirements of seagrasses make them vulnerable to reduced
light penetration of turbid coastal waters and thus erosionand
sediment transport lead to seagrass elimination. This is one of
the major threats to seagrass ecosystems in South-East Asia
(Fortes 1988; Bach et al. 1998; Terrados et al. 1998), where
deforestation leads to sediment yields to the coastal ocean ten
times higher than in any other region of the world (Miliman
& Meade 1993). The deposition of 681 g dry per m
2
per
day experienced in a Thalassia hemprichii bed in Bay Tien
(Vietnam) (Gacia et al. 2003) causeda level of burial that cannot
be sustained by most seagrass species (Duarte et al. 1997).
Direct human impacts on seagrasses threaten seagrass
habitat, particularly in areas with dense human popula-
tions. The direct impacts include: (1) shing and aqua-
culture, (2) introduced exotic species, (3) boating and
anchoring and (4) habitat alteration (dredging, reclamation
and coastal construction). Fishing methods such as
dredging and trawling (Riemann & Hoffmann 1991) may
signicantly affect seagrasses by direct removal and by
modifying the benthos (Fig. 18.3). Damage to Z. marina by
scallop dredging reduces shoot density and plant biomass
(Fonseca et al. 1984), and digging for clams can also exert
extensive damage (Orth et al. 2002). Many of these impacts
remain unquantied as yet, and their long-term effects are
unknown. In the Mediterranean, the use of certain types of
shing gear like bottom trawls has detrimental effects on
seagrass beds; in some areas, trawling marks cover 18% of
the meadow surface (Pasqualini et al. 2000) (Fig. 18.3a).
Mussel harvest in the Dutch Wadden Sea is believed to be
a major factor in the loss of Z. marina and Z. noltii there
(De Jonge & De Jong 1992).
Worldwide, coastal areas are being targeted for aqua-
culture developments. Aquaculture of sh and algal biomass
has been shown to produce major environmental impacts,
particularly because of shading, eutrophication and sedi-
ment deterioration through excess organic inputs (Seymour
& Bergheim 1991; Ackefors & Enell 1994; Shireman &
Cichra 1994; Dosdat et al. 1995; Holmer et al. 2001). The
effects of sh farms (Fig. 18.4) and other aquaculture
developments are of concern as areas of productive seagrass
habitats are often targeted for such developments, as in
the Philippines (Holmer et al. 2001), the Gulf of Thailand
(T. Ruangchoy, personal communication 1996) and along
the Mediterranean coast (Delgado et al. 1999; Pergent et al.
1999). Fish pens can cause seagrass loss (Delgado et al. 1999;
Pergent et al. 1999), and mussel culture adversely affects Z.
marina and Z. noltii beds in France (De Casabianca et al.
1997). Extensive and intensive aquaculture developments
are expanding worldwide, increasing the risk of more sea-
grass loss.
The introduction of exotic marine organisms, from
accidental release, vessel ballast water, hull fouling and
aquaculture, remains an area of concern, particularly where
the introduced species are competitors for soft-bottom
substratumsuch as the alga Caulerpa taxifolia (Meinesz et al.
1993) and the fan wormSabella spallanzanii (Lemmens et al.
1996). Large-scale engineering projects have also resulted in
species invasion, such as that by Caulerpa racemosa, intro-
duced to the Mediterranean through the Suez Canal (Lipkin
1975; Piazzi et al. 1994), which also allowed the introduction
of the seagrass Halophila stipulacea (Lipkin 1975).
Boat-propeller damage to seagrass communities is
prevalent in shallow areas with heavy boat trafc, particu-
larly in the Florida Keys (Zieman 1976). Boat anchoring
leaves scars in P. oceanica landscapes (Francour et al. 1999)
(Fig. 18.3), as do boat moorings (Walker et al. 1989).
Return of large temperate meadow-forming seagrasses to
mooring scars may take decades, and docks and piers shade
shoreline seagrass, an effect that may fragment the habitat
(Burdick & Short 1999). Boating may also be associated
with organic inputs in areas where boats do not have
holding tanks (Marba` et al. 2002).
Development of the coastline, particularly related to
increased human population pressure, leads to alteration
and fragmentation of coastal seagrass habitats. Coastal
development (construction of ports, marinas and groynes)
is usually localized around human population centres.
Housing developments impact coastal water quality, the
number of houses in a watershed being directly correlated
with the rate of seagrass loss (Short & Burdick 1996; Short
et al. 1996). Construction of the causeway at the southern
end of Cockburn Sound (Western Australia), in combin-
ation with industrial pollution, destroyed existing seagrass
(Cambridge et al. 1986). Construction of roads through
shallow waters which modies water ow may affect cir-
culation and lead to seagrass loss, such as observed in Cuba
where coastal waters were rendered hypersaline by the
effects of road construction over shallow lagoon areas.
Dredging and reclamation of marine environments,
either for extraction of sediments or as part of coastal
engineering or construction, can remove seagrasses. Land
reclamation directly eliminates seagrass habitat and results
in hardening of the shoreline, further eliminating seagrass
habitat, as seen throughout Tokyo Bay ( Japan). Groynes
alter sediment transport in the nearshore zone. Dredging
removes seagrass habitat as well as the underlying sediment,
leaving bare sand at greater depth, resulting in changes to
the biological, chemical and physical habitat values that
seagrasses support (Walker et al. 2001). Beach nourishment
(see Chapter 17) may impact adjacent seagrasses by
(a)
(b)
Fig. 18.3. Side-scan sonar echograms of two Posidonia oceanica beds at c.7 m depth in the Spanish Mediterranean littoral showing
(a) impacts of trawling as parallel scars running vertically along the image, and (b) scars left by anchor displacement along the meadow.
Each image corresponds to an area c.35 m 15 m. (Images courtesy of Mediterraneo Servicios Marinos.)
delivering sediment that may shade or bury the seagrasses.
Beach nourishment can also impact seagrasses growing in
areas where sediments collect, often at depths <30 m (N.
Marba` & C. M. Duarte, unpublished data 2001).
CLIMATE CHANGE
By 2025, climate change will not yet have had a major
impact on seagrasses globally. In some areas, there will
be warming with accompanying species shifts and sea-
level rise. For the near future, the impacts of climate
change are dwarfed by direct and indirect human impacts
(above), all of which will be negative. Climate change
looms as a possible major impact to the 2100 time horizon
and some impacts of climate change on seagrasses may be
positive.
Global climate changes derived, at least partly, from
anthropogenic combustion of fossil fuels, emission of
greenhouse gases and changes in land use with increasing
concentrations of carbon dioxide will most likely have
substantial long-term impacts on seagrass ecosystems
(Brouns 1994; Short & Neckles 1999; Duarte 2002).
Climate-related changes of potential importance for sea-
grass growth and distribution include global warming,
rising sea-level, increase in atmospheric and oceanic carbon
dioxide, and the increasing frequency and strength of
storms (IPCC [Intergovernmental Panel on Climate
Change] 2001). While the increase in carbon dioxide can be
predicted with relatively high precision, global warming
and especially its meteorological implications are more
difcult to foresee, climate changes interacting with other
human-caused changes in the marine environment making
such prediction even more complex.
The expected increase in global temperature may in
due course have numerous effects on seagrass performance
(Short & Neckles 1999). Temperature affects almost every
aspect of seagrass metabolism, growth and reproduction
(see Biebl & McRoy 1971; Evans et al. 1986; Durako &
Mofer 1987), and also has important implications for
geographical patterns of seagrass species abundance
and distribution (Walker 1991). Progressively increasing
temperature may be a major threat to local populations of
seagrasses, especially where they live close to low-latitude
borders of distribution (Spalding et al. 2003). Seagrass
Fig. 18.4. Heavily epiphytized seagrass (Posidonia oceanica) under sh cages in Sicily. (Photograph by S. A. Srensen.)
distribution shifts could be even greater if oceanic circu-
lation were to change in response to global warming,
leading to abrupt changes in water temperature beyond
those directly resulting from warming, as water masses
shift at the edge of present biogeographical boundaries
between seagrass oras (Hemminga & Duarte 2000; Green
& Short 2003).
While rising temperature may have major negative
effects on local seagrass beds, there seems to be less reason
for concern for seagrasses on the global scale. In the lit-
erature on effects of temperature on seagrasses, there
seems to be a bias towards the detrimental effects of high
temperature and less focus on the negative impact of low
temperatures. Seagrasses probably evolved in warm waters,
suggested by the high species diversity of seagrass beds in
the tropical Indo-Pacic region (Duarte 2001), and
although a few genera such as Zostera have had great
success colonizing cold temperate waters, most species
grow in warm waters (Fig. 18.1). There may be no
apparent physiological limitations latitudinally constrain-
ing seagrass distribution (Duarte et al. 2002). Nevertheless,
it is reasonable to expect that increasing global temperature
will enhance species diversity in subtropical regions and
allow cold-water species to expand their geographical
distribution further towards higher latitudes, thereby
increasing the importance of seagrass ecosystems at the
global scale (Fig. 18.1).
Warming to the year 2025 is projected also to raise sea
level by 1015 cm through thermal expansion of the ocean
and, to a lesser extent, melting of glaciers and ice sheets
(IPCC 2001). The rise in sea level has implications for
circulation, tidal amplitude, current and salinity regimes,
coastal erosion and water turbidity, each of which could
have major negative impacts on local seagrass performance
(Short & Neckles 1999).
The present atmospheric level of carbon dioxide is
relatively low considered over a geological timescale, and
the photosynthetic systems of most types of autotrophic
organisms evolved during times of much higher levels of
carbon dioxide than at present (Raven et al. 1995). With
climate change, increasing levels of dissolved carbon
dioxide in seawater may increase the competitive advantage
of seagrasses over algae because seagrasses are currently
more carbon-dioxide-limited than algae; increases in
carbon-dioxide will stimulate seagrass productivity (Beer &
Koch 1996; Zimmerman et al. 1997; Invers et al. 2001), as
it will that of terrestrial higher plants (see Bowes 1993;
Amthor 1995), rooted aquatic macrophytes (Madsen &
Sand-Jensen 1991) and oceanic phytoplankton (Hein &
Sand-Jensen 1997).
Increase in the frequency and strength of storm
events will result in increased turbidity of coastal water
and poorer light conditions for benthic plant commu-
nities (Duarte 2001). Many species of seagrasses can
survive periods of low light and partial burial (Duarte
et al. 1997), but storm events often reduce growth
and survival and require new colonization by seeds to
re-establish seagrass beds (e.g. Cabello-Pasini et al.
2002). Conversely, physical disturbance represents an
energetic subsidy and may be of advantage to species
diversity and improve growth conditions for climax plant
species (Odum et al. 1979). Uprooting of slow-growing
seagrass species forming dense canopies opens space for
colonization of more opportunistic species such as
Halophila spp. (Duarte 2002). Overall, the net effect of
increasing frequency and strength of storm events on
seagrasses is unclear.
Losses as a result of natural causes
A marine slime mould caused the large-scale wasting dis-
ease seen on both sides of the Atlantic in the 1930s (Renn
1936; Rasmussen 1977) which destroyed 90% of eelgrass
and affected scallop, waterfowl and sh populations. A
smaller-scale recurrence of the wasting disease in the 1980s
was caused by the slime mould Labyrinthula zosterae and
not evidently by human agency (Short et al. 1986, 1988;
Muehlstein et al. 1988, 1991). Wasting disease is endemic
in eelgrass; large-scale die-offs have some association with
increased salinity and water temperature (Burdick et al.
1993), and the 1930s die-off occurred at the time of a
warming period (Stevens 1936).
Seagrass is removed by large storms, cyclones and
hurricanes (see for example Patriquin 1975; Poiner et al.
1989; Preen et al. 1995), but impacts are variable and there
is no relationship between the strength of cyclones and
their impact on seagrass beds (Poiner et al. 1989).
Increasing frequency of storm events will probably
increase the damage inicted on seagrass beds. Other
extremes of climate, such as desiccation (Walker et al.
1988), erosion (Marba` & Duarte 1995) and ice scour
(Robertson & Mann 1984), can also remove seagrass.
Grazing by swans and other water birds can remove
large amounts of seagrass (Supanawanid et al. 2001).
Large-scale herbivory impacts may follow other causes
of decline. Outbreaks of sea-urchin grazing have been
reported from Cockburn Sound (Western Australia)
(Cambridge et al. 1986) and Botany Bay (New South Wales)
(Larkum & West 1990), where seagrasses were already in
decline from eutrophication effects and were unable to
recover from grazing. Dugongs and turtles can also graze
seagrass beds (Bjorndal 1980; Lanyon et al. 1989), reducing
leaves down to lengths of a few centimetres, repeated
cropping often leading to seagrass loss. These animals are
themselves under threat from human activities.
CUMULATI VE I MPACTS
There have been few coherent, broad-based studies in
either time or space of the cumulative impacts of multiple
stressors on the structure of marine communities, includ-
ing pollution, siltation, habitat fragmentation and intro-
ductions of invasive species. Further effort is needed to
investigate the inuence of these human activities on sea-
grass communities, although such multidisciplinary studies
will require a long-term funding commitment.
The most potent danger to the likely future of seagrass
ecosystems is that synergistic effects of human-caused
disturbances may arise. For instance, the increase in pre-
cipitation due to climate change and concomitant increases
in runoff of nutrients and silt from land, partly caused by
changes in land use, are potentially severe combined
threats to the distribution and performance of seagrass
beds. The loss of eelgrass from Waquoit Bay (Massachu-
setts, USA) resulted primarily from nutrient enrichment of
this enclosed coastal pond (Short & Burdick 1996), but the
rapid decline of eelgrass that occurred in shallow beds
along the shoreline was exacerbated by increasing numbers
of residential boat docks which fragmented the already
stressed beds (Burdick & Short 1999). In the Dutch
Wadden Sea, extensive losses of eelgrass occurred in the
1970s from the cumulative effects of turbidity in con-
junction with increased shellsheries, construction activity
and nutrient loading (Hily et al. 2003). Local human
impacts on Posidonia oceanica beds combined with large-
scale erosion, derived from urbanization and sea-level rise,
have caused widespread decline of north-west Mediterra-
nean seagrass beds (Marba` et al. 1996; Marba` & Duarte
1997). Anthropogenic pressures also render seagrass eco-
systems more vulnerable to negative impacts from climate
change. For instance, modication of the coastline by the
construction of harbours and other structures on the shore
leads, in the presence of sea-level rise, to considerable
beach erosion (see Chapter 17), which propagates down-
slope to cause losses to seagrass ecosystems (Marba` &
Duarte 1997).
While examples of the effects of cumulative impacts
exist, a framework within which to forecast the responses
of seagrasses to multiple stresses is still lacking, so that the
preceding discussion of likely impacts of direct and
indirect human pressures and climate change on seagrass
ecosystems must be considered a conservative scenario of
the possible losses.
PROSPECTS
The current rate of seagrass loss illustrates the imperilled
status of the ecosystem and the need for increased public
awareness, expanded protective policies and active man-
agement. In order to achieve such goals it is important
to focus resources to monitor seagrass habitat trends,
conserve existing seagrass resources, act to attenuate the
causes of seagrass loss, and develop knowledge and tech-
nologies to reverse ongoing seagrass decline.
The widespread loss of seagrasses is largely a consequence
of the rapid growth in human activities and transformation of
the coastal zone. Global human population growth is con-
centrated in the coastal zone (Cohen 1995; Nicholls & Small
2002), which also harbours a disproportionate fraction of the
worlds wealth. Some rapidly growing industries are linked to
the marine environment, such as tourism, maritime transport
and aquaculture. Consequently, human activity in the coastal
zone is likely to continue to increase, with a potential for even
greater impacts on seagrasses (Duarte 2002). Future devel-
opment-derived impacts on seagrass ecosystems are likely to
be greatest in developing countries, which contain much of
the extant global seagrass area (Green & Short 2003). There,
human population growth is forecast to be fastest, and the
potential for adopting sustainable approaches to development
is constrained by resources available for management.
Moreover, should resources become available, these are likely
to be diverted to ensure the conservation of ecosystems for
which there is greater public awareness both locally and
globally, such as coral reefs (Chapter 16). Indeed, only
recently have seagrass ecosystems been targeted in designing
marine protected areas (MPAs). The prospect for seagrass
ecosystems is therefore not positive and this prognosis can
only be reversed through the development of a new attitude,
involving proactive policies and efforts to manage, protect
and restore seagrass ecosystems.
MANAGING, PROTECTING AND MONI-
TORING SEAGRASS
ECOSYSTEMS
Increasing awareness of the importance of seagrass has led
many countries to enact legislation to prevent destruction of
these habitats. Their value is recognized in the Convention
on Biological Diversity (Coles & Fortes 2001). Environment
Australia (EA) has federal powers to manage dredging across
Australia to minimize seagrass damage. The Environmental
Protection and Biodiversity Conservation Act legislates
against seagrass destruction. State agencies, such as those
in Western Australia and Queensland, have policies to
prevent removal or destruction of seagrasses and a mandate,
where necessary, to replace them. Strong measures to pro-
tect seagrass have also been implemented through the
Habitats Directive of the European Commission, which
specically recognizes seagrass meadows as preferential
habitats for conservation policies. Many countries in Europe
(such as France and Spain) have implemented legislation to
protect seagrass meadows. However, some of these laws
restrict coastal zone use to such an extent that they are
impractical to apply and are therefore ineffective for
management.
Even under these best-case scenarios of management
and protection, losses still occur, and the global status of
seagrass management and protection is poor. Most seagrass
habitat is unprotected, and where existing policies are
protective they are rarely enforced. Active management of
seagrass habitat occurs in only a very small proportion of
the worlds seagrasses. In many areas where management
does occur, conicting uses of the coastal zone nonetheless
often result in seagrass decline or loss. As these are
underwater habitats that are not easily seen from shore,
they are often overlooked or damaged before their exis-
tence is noted. Seagrass protection policies, protective
measures and enforcement are very variable globally
(Table 18.1).
Increasing awareness and understanding of the
importance of seagrasses is critical to their survival. Edu-
cating the public to appreciate the importance of aquatic
resources and effective ways to manage them, in combin-
ation with establishment of effective MPAs, is one way of
ensuring seagrass survival. In addition to legislation that
protects seagrass beds across countries or continents as in
Europe and Australia, efforts to protect seagrass beds
through the designation of seagrass meadows as MPAs are
growing worldwide. The implementation of the Natura
2000 network of sites, aimed at preserving a fraction of the
territory of European Union member states, has resulted
in a major increase in the area of Mediterranean seagrass
beds under protection. Some seagrass beds are now central
elements of World Heritage sites, such as the Posidonia
oceanica meadow in Ibiza-Formentera (Spain). The Shark
Bay World Heritage Site (Western Australia) contains
some 5000 km
2
of seagrass beds of high diversity, as well as
populations of turtles, dolphins and more than 10 000
dugong (Walker 1989). The Great Barrier Reef World
Heritage Site (Queensland, Australia), contains lagoonal
and deep-water seagrasses (Poiner et al. 1989). In the USA,
the Florida Keys National Marine Sanctuary has been
declared to help protect and conserve seagrass commu-
nities, as well as the coral reefs, hard-bottom habitats,
mangroves and the marine life (NOAA [National Oceanic
and Atmospheric Administration] 2006). The development
of MPAs must form part of the framework for sufciently
comprehensive and representative marine conservation.
Clearly, management of seagrasses is inadequate on a
global scale. Even in areas with a high degree of manage-
ment, development and commercial use of coastal waters
and upland watersheds create indirect impacts on seagrass
ecosystems. Moreover, evaluation of the effectiveness of
protection of seagrass in MPAs is still insufcient, so that
the effectiveness of current conservation practices remains
poorly evaluated. In the Cabrera Archipelago National
Park (Spain), while P. oceanica beds are recovering, a slow
process that will require centuries to be completed, their
continued recovery is jeopardized in places by organic
inputs from visitors to the Park (Marba` et al. 2002).
Identifying such responses requires efcient monitoring
programmes.
An unresolved issue in most monitoring programmes
is their power to detect change. Most monitoring pro-
grammes use estimates of seagrass cover or density, which
are generally poor at detecting change (Heidelbaugh &
Nelson 1996), and substantial seagrass loss may remain
undetected. Improved monitoring methods are required.
Several seagrass monitoring programmes have been ini-
tiated across the globe at various levels. Monitoring pro-
grammes at local scales are many, including the French
programme started in 1984, which surveys the upper
and lower limits of P. oceanica meadows at 33 sites in
the Mediterranean, and which has shown the beds to be
increasing in size as wastewater treatment plants have
reduced nutrient inputs to coastal waters (Boudouresque
et al. 2000). Citizen monitoring using community volunteers
can be a valuable and effective mechanism for increasing
public awareness of seagrass habitat and tracking large-scale
changes. In Queensland (Australia) and the Western Pacic,
Seagrass-Watch is an outstanding example, with many
participants, a regular newsletter and ongoing tracking of
habitat extent and change (McKenzie et al. 2000; Coles et al.
2003b). Similar volunteer programmes have been estab-
lished in Spain.
Thorough monitoring programmes at the national level
exist in a few countries, such as that run by Danish county
authorities, which since 1989 has provided excellent data
on changes in seagrass communities at the national level
(rtebjerg et al. 2003). In 1992, in the USA, the NOAAs
Coastal Change Analysis Program (C-CAP) was initiated
to monitor seagrass distribution and establish ongoing
assessment using aerial-photography distribution maps
Table 18.1. Status of seagrass protection by region
Area
Information
on seagrass
protection Legislation
Marine
protected areas
that include
seagrasses
Enforcement of
policies References
a
North Pacic Mostly poor
except
USA
Some in USA,
no net loss
policies and
sh habitat
protection
Country
dependent
None in some
areas to
moderate
(USA)
Wyllie-
Echeverria
et al. (1994),
UNEP
(1999a)
Chile, south-
west Atlantic
Poor Not reported Not reported Not reported Not cited
North Atlantic Good in USA
and EU
Yes, varies
between states
and countries
Few, but high
protection not
guaranteed
Ranges from
poor to high
Wolff (1997),
NOAA (1998)
Caribbean Good in some
countries
Threatened
animal species
and sheries
Yes, but high
protection not
guaranteed
None in some
areas to
moderate
(USA)
Clark (1998),
NOAA (1998)
Mediterranean Patchy Inter-country
agreements in
EU
Yes, but high
protection not
guaranteed
None in some
areas to
moderate
(EU)
UNEP (2000b)
South-east
Atlantic
Poor Not reported Not reported Not reported Not cited
South African Poor Not reported Not reported Not reported Not cited
Indo-West
Pacic
Poor except
Australia
and the
Philippines
Yes, in
Australia,
and the
Philippines,
and in some
Pacic island
nations
Yes, in SE Asia
and the Pacic
including
Australia
Poor to
non-existent,
except high in
Australia
Coles (1996),
Leadbitter
et al. (1999)
a
UNEP, United Nations Environment Programme; NOAA, National Oceanic and Atmospheric Administration.
Source: Adapted from Coles and Fortes (2001).
(Dobson et al. 1995). C-CAPs goal is to map seagrass
distribution at 5-year intervals for the coast of the USA,
using standardized protocols, and has demonstrated
signicant seagrass loss (Short & Short 2003).
Regional-level efforts are also in place, such as
the Caribbean Coastal Marine Production Network
(CARICOMP) started in 1985 to monitor seagrass, coral
reefs and mangroves. It consists of marine laboratories and
conservation groups using standardized techniques, and
aims to determine the dominant inuences on coastal
productivity, monitor ecosystem change and detect human
impacts (Creed et al. 2003).
Individual scientists around the world are monitoring
seagrass ecosystems in their areas. These efforts provide a
valuable assessment of the seagrass resource, but there is no
central data repository and methods are often not com-
parable, making it difcult to assess seagrass status at the
global scale. A global monitoring programme to assess the
status and trends of the seagrass ecosystem (SeagrassNet)
has since 2001 collected quarterly data from xed transects
on seagrass species composition, biomass, distribution and
depth and environmental variables in 14 countries of the
Western Pacic, Australia, Brazil, Africa and the USA,
using a standard protocol (Short et al. 2002c). The aim is a
global report card on the status and prospects of seagrasses
worldwide.
Restoring seagrass ecosystems
Reduced anthropogenic nutrient loading and elimination
of sediment discharge into coastal waters will result in
improved water clarity and allow seagrasses to begin to re-
establish their former distribution, achieve historical depth
limits and reverse the downward spiral of habitat decline
(Duarte 1995). The current trends in human population
growth and distribution make achievement of these goals a
very great challenge with major demands on nancial
investment, political change and environmental awareness.
Direct improvement of seagrass distribution may be
achieved through restoration, which has progressed from
early transplant efforts to scientic investigations of site
selection and improved transplant methodologies (Short
et al. 2002a). The scientic tools for seagrass restoration
are: (1) a quantitative site selection model (Short et al.
2002a), (2) a protocol to identify a sustainable source of
planting stock, (3) a reliable planting method (Davis &
Short 1997; Harwell & Orth 1999; Granger et al. 2000;
Short et al. 2002b), and (4) monitoring using scientic
criteria to identify the outcome of the restoration (Short
et al. 2000). The most important tool is site selection,
ensuring adequate water clarity, low bioturbation and
appropriate sediment and physical conditions (Short et al.
2002a). Determining success of seagrass restoration
requires rigour and demonstrated replacement of habitat
function. Research on all these aspects of seagrass restor-
ation is under way in many parts of the world.
Sound science is important as a basis for restoration
implementation and practice, ranging from providing rig-
orous methodology for fullling the statutory requirements
for compensatory mitigation to offset impacts on seagrass
(Davis & Short 1997; Fonseca et al. 1998; Paling et al.
2001) to simplifying the techniques. In addition, costs have
to be reduced and procedures developed that community-
based volunteer groups can use to restore coastal envir-
onments (Short et al. 2002b). Both kinds of restoration,
namely mitigation and community-based, are needed to
slow the loss and reverse the degradation of seagrass
habitat.
Revegetation efforts are expensive, and have proved
effective only at small spatial scales (areas <1 km
2
), and
mainly in sheltered environments. It has only been
attempted with Z. marina, Z. noltii, Halodule wrightii,
T. testudinum, Syringodium liforme, Posidonia australis,
P. coriacea and Amphibolis grifthii (Fonseca et al. 1998;
Lord et al. 1999). Revegetation cannot fully restore lost
seagrass areas, but can stimulate natural revegetation
processes. Seagrass recolonization is slow, requiring from a
few years for the fast-growing species to centuries for the
slow-growing species (Duarte 1995; Marba` & Duarte 1998;
Kendrick et al. 1999; Marba` et al. 2002). The seagrass area
that can be recovered through revegetation in the most
optimistic scenarios is but a small fraction of the area lost
annually. Hence, while a useful management option,
revegetation efforts must be combined with effective
conservation and improved water clarity (monitoring has
no direct effect on seagrass loss) if seagrass loss is to be
reversed.
OUTLOOK FOR SEAGRASSES I N 2025
Worldwide, seagrasses are being lost faster than they can
regrow or recolonize. By 2025, seagrass losses on all con-
tinents and in the oceanic islands will be greater than they
are now and more noticeable, the extent of this loss
depending on the implementation of immediate steps to
eliminate the global threats to seagrass habitat. Despite
some increased awareness and protection, there is no sign
that global losses of seagrasses will be slowed in the next 20
years. Most impacts on seagrass ecosystems to the year
2025 will be from increasing human population densities,
rather than predicted climate change or sea-level rise.
Human impacts, particularly through eutrophication and
increased sedimentation, will continue to cause seagrass
losses worldwide.
Climate change is likely to have long-term, albeit not
large, impacts before 2025. Increased sea level may per se
have neutral effects on seagrasses, but lead to losses in
already eroding coastal areas, particularly those with the
prospect of increased storm frequency. In addition to
promoting submarine erosion, hardening of shorelines
will limit the progression of seagrasses facing sea-level
rise, restricting the ability of seagrasses to migrate
shorewards and thus reducing available habitat. Increased
concentrations of carbon dioxide may favour seagrass
growth.
Only increasing awareness of the need for development
to be sustainable, and the need to protect and conserve
ecosystems, will help mitigate expected seagrass losses. A
precondition is that the values and services of seagrass
ecosystems become more widely acknowledged in order to
foster efforts to promote legislation and enforce protection
measures for seagrasses worldwide.

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