Mapping seagrass by remote sensing is more difficult than mapping mangroves and tidal

marshes due to water turbidity, color sun glint and the epiphytes that cover the blades of

grass may dilute the spectral reflectance signal of seagrasses and hinder the ability of the

instruments to see through the water (Howard et al., 2014). Then to clarify the usefulness and

limitation of remote sensing for mapping seagrasses, a manual for mapping seagrass and

seaweed beds distribution with satellite images was developed in 2015 .

The literature reviewed by the national expert of China reported that five seagrass species are

distributed in the NOWPAP region of China . Zostera marina, Z. japonica, Z.

caespitosa, Phyllospadix japonicus and P. iwatensis are distributed along the coastal areas of

Liaoning, Hebei and Shandong provinces (Zheng et al., 2013; Guo et al., 2010; Yang, 1979;

Zhang et al., 2013; Yang, 2017). Zostera is the dominant species of most seagrass meadows. In

addition, the distribution of Z. japonica is not limited to the northern coast of Liaoning, Hebei and

Shandong, but also to the coast of Fujian, Taiwan, Guangxi, Hong Kong and even Hainan

provinces (Zheng et al., 2013).

Progressing irreversible degeneration of the spatial structure of the seagrass ecosystem of

Alekseev cove (Popov Island, Peter the Great Bay) under the influence of activity on breeding

aquaculture of sea scallops and mussels is reported (Zharikov and Preobrazhensky, 2010). Large

populations of Zostera marina located within the port area in Sovetskaya Gavan Bay are subject

to significant anthropogenic pollution (Dulenin, 2012). Threats by a typhoon to the seagrasses in

Srednaya Bay (Zharikov et al., 2015), threats by the development of ports, pollution and

aquaculture to the seagrasses in Severnaya Bay (Zharikov et al., 2016) and threats by the

development of ports and pollution to the seagrasses in Tokarevsky Koshka (Eastern Bosphorus

Strait) (Kalita and Scriptsova, 2017) are reported.

The mapping of seagrass and seaweed bed distribution with satellite images has been

conducted by scientists since the launch of a satellite sensor into space (e.g. Komatsu et al.,

2012; Noiraksar et al., 2014).

The region as a

whole has seen a rapid population expansion of nearly sixfold between 1900 and 2000 (Jones 2013). The
region is also a global biodiversity hotspot, with high numbers of endemic species in both the marine
and terrestrial environments (Sodhi et al. 2010, Tittensor et al. 2010).

Seagrass meadows are widely recognised as offering a number of key ecosystem services, such as
habitat formation, nutrient cycling, carbon sequestration, and food provisioning (William and Heck 2001,
McGlathery et al. 2007, Fourqurean et al. 2012, Cullen-Unsworth and Unsworth 2013), with many
coastal populations within Southeast Asia directly dependent on these habitats for a living (Unsworth
and Cullen 2010). Seagrasses also continue to decline at a global rate of approximately 7% (Waycott et
al. 2009), and the losses may be more acute in the Southeast Asian region due to the pressure from
increasing coastal populations and developments, and the lack of data on seagrass resources. Perhaps
just as important are the socio-economic-cultural links between seagrass meadows and coastal
populations, which is only now coming to the attention of researchers (Unsworth and Cullen 2010).

In terms of gene flow, this ecoregion is recognised as the Indo-Pacific Barrier (Bowen et al. 2016), the
equivalent of a marine Wallace line that separates populations of marine fauna on either side of the
strait through shifts in sea level during periods of glaciation. This insight is useful in guiding the selection
of sampling locations for seagrass phylogeographic studies that address questions about biodiversity
distributions in Southeast Asia, and specifically in testing hypotheses about Southeast Asia as a center of
overlap, refuge, accumulation or centre-of-origin for seagrass, such as those suggested by Mukai (1993)
and Nguyen et al. (2013), Nguyen et al. (2014).

Halophila major and Halophila ovata are also limited to single ecoregions in the Northern Bay of Bengal,
and the Andaman and Nicobar Islands, but this may be because these forms are taxonomically difficult
to identify (addressed in Taxonomic Highlights below), although recent progress has been made in
molecular approaches (Nguyen et al. 2013, 2014).

Seagrass species diversity is highest in the Philippines (19 species), and lowest in Brunei (7 species),
which is the country with the most recent additions to its species list (Lamit et al. 2017).

tiny in this region, which we briefly summarise below. In Southeast Asia, Halophila major has been
studied in greater detail than other species in its genus, through a combination of morphological and
molecular traits. As a result, it has become a recent entry into the species

records of Indonesia (Tuntiprapas et al. 2015), Thailand (Tuntiprapas et al. 2015, Myanmar (Nguyen et
al. 2014), Viet  Nam (Nguyen et al. 2013, 2014), and the Philippines (Kim et al. 2017).
Halodule pinifolia is considered to be the narrow-leaved form of Halodule uninervis (Waycott et
al.2004), but these have been shown through genetic analysis on samples in the Philippines to be
separate species that have morphological variations in leaf width because of site-specific differences,
density and exposure (Wagey and Calumpong 2013).

It has only been in recent years that areal estimates for seagrass meadows have begun to emerge more
rapidly as a result of advancements in remote sensing technology and well-funded regional projects
such as the UNEP/GEF South China Sea Project (UNEP 2008), the Bay of Bengal Large Marine Ecosystem
Project (BOBLME 2015), and the JSPS-Asian CORE Project.

We consider meadow size data to be particularly critical for moving seagrass conservation and
management forward in the region because these provide Halophila major from Tanjung Adang Laut,
Sungai Pulai estuary, Johor (refer also to Nguyen et al. 2014 for morphological and genetic identification
of the species).

Because they are physically isolated, these sites are natural laboratories for testing ideas about
allopatric differentiation in seagrass and seagrassassociated species, and to elucidate seagrass dispersal
routes in the region. To take the example of fish larvae in the Spratly Islands, larval drift time and vector
current charts indicate that the western Philippines, Taiwan, south-eastern China, Brunei, and Malaysia
are direct sink habitats for coral reef fish from the Spratlys (McManu 2017). Long-distance dispersal in
seagrasses has also been shown to be possible through seeds, fruits, viviparous seedlings and vegetative
fragments, all of which have the capacity to move over hundreds of kilometers in distance (Kendrick et
al. 2012).

Island-building often involves sanddredging and land reclamation on shallow coral reefs and in lagoons.
To date, two-thirds of currently occupied atolls in the Spratlys have been shown to have proportionally
less reef extent than unoccupied atolls, implying the detrimental effects of island-building on reef
systems (Asner et al. 2017).

Seagrass research in Southeast Asia is increasing based on the number of research papers that have
been produced in the last decade (Ooi et al. 2011a), and an update of the number of research articles,
reports and studies that have been reported in the region has shown a steady increase in research
output decade by decade for most countries.

Taxonomic research continues to be a mainstay of the research efforts in this region, with new species –
particularly in the Halophila ovalis species complex (Waycott et al. 2004) – being described (see Kuo
2007, Japar Sidik and Muta Harah 2011, Fortes 2013).

However, there is an obvious need for traditional morphological seagrass

taxonomy to embrace new methods in the field. There is currently already a Global Initiative to Barcode
Seagrass (GIBS) based at the State Herbarium of South Australia and the University of Adelaide (GIBS
2018), and more participation from researchers in the Southeast Asian region may be beneficial for the
advancement of seagrass taxonomy.There is an increasing trend of research output for the region as a
whole (Ooi et al. 2011a), and there are some distinct trends in the intensity of work within each
thematic research area across the decades .

For example, there is virtually no social science research examining the social, cultural and economic
aspects of human-seagrass interactions, and there is research emerging that points to important
relationships between coastal populations and seagrass beds (Cullen-Unsworth et al. 2014).
Furthermore, while the significance of seagrass fishery activity has been acknowledged globally
(Nordlund et al. 2017), the lack of information on seagrass fishery resources and the flow-on benefits
from ecosystem services it provides in the Southeast Asian region is a data gap that needs to be
addressed (Unsworth et al. 2009).

These impacts need to be addressed in order to stem further deterioriation of seagrass meadows across
the region. Although attempts are occasionally made to undertake seagrass relocation and restoration,
these are often not the most cost-effective solution as success rates are quite low despite a large input
of effort and funds (van Katwijk et al. 2015). These ineffective solutions are also often compounded by
the lack of effective linkages between science, government, and private sectors, leading to poor
management and conservation actions taken. There is currently a big gap or disconnect between
seagrass science, policy, and practice (Fortes 2018).

Designation of marine parks is also often inadequate, with fishing, aquaculture, or other anthropogenic
activities that impact seagrass meadows continuing to take place (e.g. Guimarães et al. 2012). One of the
major difficulties surrounding implentation and enforcement of enviromental laws is often resistance
from the local seafaring communities (Bennett and Dearden 2014).

Such initiatives provide long-term monitoring data of seagrass meadows at a much lower cost, allowing
research funding to be directed at other research avenues (Theil et al. 2014). A successful example of a
long-term citizen science monitoring program in Southeast Asia is TeamSeaGrass (Yaakub et al. 2014b).
TeamSeaGrass is part of the Seagrass-Watch Network and the data collected by volunteers have
contributed towards scientific publications (Yaakub et al. 2014c, McKenzie et al. 2016), and been shared
with managers from the National Parks Board (NParks) in Singapore.

Twelve ISBWs have been held and each followed individual themes reflecting both a geographical or
institutional interest, and current trends in seagrass research as new knowledge and techniques became
available (Coles et al. 2014).

Seagrasses are angiosperms thriving best in slightly

reducing sediments of shallow tropical and subtropical

coasts. Here, they form an ecosystem, dominating it as a

discrete functional, not as a taxonomic group (McKenzie

et al. 2010)
This is largely due to the fact that

the interests of marine scientists focus mainly on coastal

resources with immediate economic value or impacts (e.g.

coral reefs, seaweeds, animals, or fish, tourism, coastal

development) (Fortes 1989, Fortes 2012).

This effort, nevertheless, should be

science-based (Fortes O010a, O010b). Ogawa et al. (O011)

gives a more recent account of the status and trends in

seagrass resources in 5 countries in fhe region (including

Japan). Coles & Fortes (O001) reported the approaches and

methods to protect seagrass. In Southeast Asia where the

second highest seagrass diversity in the world is found, seagrass ecosystem has been a focus of scientific
inquiry

only in the last 30 years and, as an object of natural

resource conservation, only in the last 15 years! It took

more than five decades after the morphological account

of Pascasio & Santos (1930) to follow up the works on

taxonomy and distribution and initiate an ecological

research on seagrasses in the Philippines (Fortes 2012).

In the last 50 years, a mix of research works appeared,

presenting various methods of conservation, rehabilitation

and management of the seagrass habitats in the country,

together with the organisms associated with them. This

impetus resulted into a rapidly increasing importance

of integrated studies, including genetic markers as a

means to understand their phylogeography in the region (Matsuki et al. 2012, Nakajima et al. 2012).

The

shifting needs of the times, aggravated principally by an

alarming reduction and loss in resources resulting from

a decline in coastal water quality and degradation of the

environment, dictated a corresponding shift in seagrass

research focus from basic to its applications, from purely

scientific initiatives to those that now require support

and collaboration from social and behavioral sciences

(Fortes 2012).

Hence, for the first time in decades, a seagrass (Halophila

becarrii) has been included in the IUCN Red Book as

a locally threatened species (Short et al. 2011).

The region as a whole has seen a rapid population expansion of nearly six-fold between 1900 and 2000
(Jones 2013).

The region is also a global biodiversity hotspot, with high numbers of endemic species in both the
marine and terrestrial environments (Sodhi et al. 2010, Tittensor et al. 2010).

Perhaps just as important are the socio-economic cultural links between seagrass meadows and coastal
populations , which is only now coming to the attention of researchers (Unsworth and Cullen 2010).

In terms of gene flow , the ecoregion is recognized as the Indo-Pacific Barrier (Bowen et al. 2016) , the
equivalent of a marine Wallace line that separates populations pf marine fauna on either side of the
strait through shifts in sea level during periods of glaciation.

Seagrass species diversity is highest in the Philippines (19 species) ,and lowest in Brunei (7 species),
which is the country with the most recent addtions to its species list (Lamit et al. 2017).

In Southeast Asia, Halophila major has been studied in greater detail than other species in its genus,
through a combination of morphological and molecular traits. As a result, it has become a recent entry
into the species records of Indonesia ( Tuntiprapas et al. 2015), Thailand ( Tuntiprapas et al. 2015,
Myanmar(Nguyen et al. 2014), Viet nam (Nguyen et al. 2013,2014), and the Philippines (Kim et al. 2017).
Halophila minor and Halophila ovata were treated as synonyms for the seagrass floral of Singapore
(Yaakub et al.2013).

However , H. Ovata was subsequently considered an illegitimate name proposed as Halophila

gaudichaudii by Kau et al. (2006).

Related studies

Foreign
Southeast Asia is a biologically diverse region with a high diversity of marine species globally
(Groombridge and Jenkins 2002, Spalding et al. 2003, Kulbicki et al. 2013, Huang et al. 2015).
Bordered by the Bay of Bengal to the west, the Indian Ocean to the South, and the Pacific Ocean
to the east, this region has approximately 117,763 km of coastline (Fortes et al. 2018), and two of
the largest archipelagic nations (Indonesia and the Philippines) in the world. The region is also
undergoing rapid economic development and coastal change, with many marine ecosystems
being threatened by anthropogenic activities and, by extension, environmental change
(Wilkinson and Salvat 2012).
The coastal and maritime nature of the region translates into a close relationship between the
sea and marine resources, with the cultures, communities, and economies of this region (Hung
et al. 2013). 
Seagrass habitats play an important role in the biological, socio-cultural and economic
landscape in Southeast Asia (Unsworth and Cullen 2010), but suffer disproportionately in terms
of conservation priority. 
Seagrasses are declining at an unprecedented rate globally (Orth et al. 2006), but information
on the loss of seagrass habitats in Southeast Asia is limited, although there is growing evidence
of widespread decline and loss throughout the region (Yaakub et al. 2014, Unsworth et al. 2018).
Much of the early research on seagrass in Southeast Asia was concentrated in Indonesia and the
Philippines, but the publications and study sites in these early papers are clustered around
Northwest Luzon in the Philippines and South Sulawesi in Indonesia (Ooi et al. 2011).
Knowledge on seagrass is increasing across Southeast Asia, with a variety of research themes
emerging resulting from access to technology and resources, but is hampered by lack of
coordination and collaboration within the region (Fortes et al. 2018). The review by Fortes et al.
(2018) highlights locations within the Southeast Asian region where information and research
on seagrass resources remain limited, such as in the Andaman and Nicobar Islands, Cambodia,
Myanmar, Timor Leste, and Viet Nam. Within this special issue, information on aspects of
seagrass community ecology (Phan et al. 2018), fisheries (Jones et al. 2018), and biogeography
(Savurirajan et al. 2018), are emerging from some of these data-deficient areas. 
Remote sensing and the availability of quality license-free satellite imagery has created an
opportunity not only for plugging the knowledge gaps in the distribution, location, and extent of
seagrass habitats in Southeast Asia, but in monitoring meadow-scale changes and dynamics of
seagrass beds (Bramante et al. 2018).
For example, seagrass habitats in Southeast Asia lie mostly in shallow, semi-enclosed coastal
systems and large estuaries, where tidal fluctuations bring about considerable daily and seasonal
variations in key drivers like light availability and salinity. Here we see that fluctuations in
physicochemical parameters such as irradiance (Phandee and Buapet 2018) and salinity
(Kongrueang et al. 2018) can affect seagrasses at the physiological level as well as whole plant
communities at the broad scale (Phan et al. 2018).
Environmental and historical drivers also play an important role in determining how seagrass
populations form, and are maintained. There is evidence that complex geological processes have
contributed to the genetic structure observed in the Sunda Shelf and Banda Sea (Wainwright et
al. 2018). This study contributes to the growing body of knowledge on seagrass population
genetics in Southeast Asia (Nakajima et al. 2014, Arriesgado et al. 2015, Hernawan et al. 2017).
The trophic interactions found in tropical seagrass meadows are poorly understood, despite
being an important aspect of seagrass meadow maintenance, and the work of Fong et al. (2018)
documents the diversity of gastropod grazers in seagrass meadows, as well as important trophic
pathways between seagrasses, their associated epiphytes and the gastropod grazers that feed on
them. The overexploitation of seagrass fauna could have important consequences for seagrass
meadows and the communities that rely on them for food provisioning services. This is
especially important in data deficient areas such as Myanmar, where a depauperate diversity of
motile fauna could be heralding larger complications for marine ecosystems in the region (Jones
et al. 2018). 
Seagrass restoration is necessary to stem seagrass decline and to return seagrass meadows to a
healthy state and is able to provide critical ecosystem services (Orth et al. 2006, van Katwijk et
al. 2016). 
While seagrass restoration is not new in Southeast Asia, there has been varying degrees of
success in the survival of transplants in the tropics (Paling et al. 2009). Asriani et al. (2018) in
this special issue report on a novel approach of transplanting mixed-seagrass species
combinations and found that transplantation increased with species richness. 
Widespread anthropogenic threats to seagrass habitats and the rapid rate of coastal change in
the region call for quick and decisive action on seagrass conservation. This may not always be
feasible when there is a paucity of information on the seagrass meadows. Tan et al. (2018)
propose a simple method for prioritising sites for conservation based on a combination of two
well-defined indexes: ecosystem service assessments and habitat vulnerability analyses. 
Conservation of seagrass meadows will ultimately rely on the ability of scientists and managers
to work together across the region to effect a change in the attitudes of people towards seagrass
habitats, and the review paper in this special issue provides a roadmap for seagrass conservation
in the region (Fortes et al. 2018).