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Mangrove Biogeography of the Indo-Pacific

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 Mangrove Biogeography
of the Indo-Pacific 23
P. Saenger, P. Ragavan, C.-R. Sheue, J. López-­
Portillo, J. W. H. Yong, and T. Mageswaran

Abstract using molecular studies warrant a further

Studies on biogeography are useful to under- comprehensive account of the biogeography
stand the present and past distribution pat- of mangroves. Considering these facts, the
terns of biological diversity and their biogeography of the Indo-West Pacific (IWP)
underlying environmental and historical mangroves has been examined in detail, and
causes. The mangrove realm, largely confined the Atlantic East Pacific mangrove region
to sheltered tropical and subtropical coast- (AEP) has only been briefly discussed. The
lines within latitudes of around 32°N and difference in species richness between the
38°S, is divided longitudinally into an Atlantic IWP and the AEP is evidence of the effective-
realm and an Indo-Pacific realm. Because of ness of the African land mass and the East
the relatively recent closure of the Central Pacific Ocean barriers. Within the IWP, two
American Isthmus, a small incursion of the trends in relation to mangrove species rich-
Atlantic realm species has occurred into the ness can be identified: first, maximal species
Indo-­Pacific realm along the tropical and sub- richness which occurs along the shorelines of
tropical American Pacific coast and spread Makassar Strait, between Borneo and
north and south since the last glaciation. Sulawesi in Indonesia, and second, the
Although the biogeography of mangroves has marked attenuation of species numbers with
been widely discussed, recent investigations increasing latitude generally related to limit-
of mangrove floristics and genetic diversity ing temperatures. The contemporary and his-

P. Saenger (*)
School of Environment, Science and Engineering,
Southern Cross University, Lismore, NSW, Australia J. López-Portillo
e-mail: [email protected] Instituto de Ecologia, A.C. (INECOL),
Xalapa, Veracruz, Mexico
P. Ragavan
SERB-National Post Doctoral Fellow, J. W. H. Yong
CSIR-National Botanical Research Fellow, Lucknow, ARC Centre for Mine Site Restoration, School of
Uttar Pradesh, India Biological Sciences, University of Western Australia,
Perth, Australia
C.-R. Sheue
Department of Life Sciences and Center of Global T. Mageswaran
Change Biology, National Chung Hsing University, National Centre for Sustainable Coastal Management,
Taichung, Taiwan MoEFCC, Chennai, Tamil Nadu, India

© Springer Nature Switzerland AG 2019 379

B. Gul et al. (eds.), Sabkha Ecosystems, Tasks for Vegetation Science VI,
https://doi.org/10.1007/978-3-030-04417-6_23
380 P. Saenger et al.

torical processes leading to these trends are Recent Taxonomic

summarized and discussed. Finally, the and Distributional Changes
threats to the mangroves of the various in the Mangrove Flora
regions within the IWP are subsequently
briefly reviewed, and given the ecological and Recent taxonomical studies led to an understand-
economic values of mangroves, the potential ing of extended distribution of few extant man-
of mangrove restoration/rehabilitation to off- grove species and the discovery of new entities.
set mangrove losses is also evaluated. Some of the significant findings are as follows.
The taxonomic relationships of the Australian
Keywords Excoecaria species were examined by Maguire
Biogeography · Diversity · Mangrove · and Saenger (2000) using both leaf morphologi-
Rehabilitation · Restoration · Species richness cal data and DNA sequence data from the internal
transcribed spacer (ITS) region of ribosomal
genes. The nucleotide differences in the exam-
ined ITS1 region showed that E. agallocha con-
Introduction sisted of two species – the widespread E.
agallocha and the species confined to northern
Studies on biogeography are useful to understand Australian mangroves, previously described as E.
the present and past distribution patterns of bio- ovalis. The leaf morphological data also support
logical diversity and their underlying environ- the re-erection of E. ovalis: single factor analysis
mental and historical causes. The mangrove of variance consistently separated E. ovalis from
realm, largely confined to sheltered tropical and E. agallocha on the basis of leaf width, leaf
subtropical coastlines within latitudes of around length, length of petiole and leaf margin.
32°N and 38°S, is divided longitudinally into an A recent re-examination of the genus
Atlantic realm and an Indo-Pacific realm Kandelia, long considered monotypic, showed
(Tomlinson 1986; Ricklefs and Latham 1993). that two well-differentiated populations existed,
Because of the relatively recent closure of the separated by the South China Sea. These were
Central American Isthmus (about 3.5 million established as two distinct species Kandelia can-
years ago), a small incursion of the Atlantic realm del and K. obovata (Sheue et al. 2003).
species has occurred into the Indo-Pacific realm The genus Ceriops has also been revised
along the tropical and subtropical American recently (Sheue et al. 2009a, b, 2010) and has
Pacific coast and spread north and south since the been found to consist of three species on the basis
last glaciation (Sandoval-Castro et al. 2014). that the species formerly known as C. decandra
Although the biogeography of mangroves has consisted of a species complex: Ceriops decan-
been widely discussed, recent studies of man- dra occurs in India, Bangladesh, Myanmar and
grove floristics and genetic diversity using Thailand; Ceriops pseudodecandra occurs in
molecular studies warrant a further comprehen- Australia, Papua New Guinea and in the
sive account of the biogeography of mangroves. Indonesian provinces of Irian Jaya and Seram;
Considering these facts, the biogeography of the and Ceriops zippeliana occurs in Indonesia,
Indo-West Pacific (IWP) mangroves has been Malaysia, Vietnam and the Philippines.
examined in detail, and the Atlantic East Pacific Sheue et al. (2009a) have re-examined the dis-
mangrove region (AEP) has only been briefly tinctiveness of Ceriops tagal and Ceriops austra-
discussed. The threats to the mangroves of the lis. In the process, Ceriops australis was found to
various regions within the IWP are subsequently occur widely in Indonesia, constituting a major
briefly reviewed, and given the ecological and increase in its known distribution from Australia
economic values of mangroves, the potential of to the southern coast of New Guinea.
mangrove restoration/rehabilitation to offset Sheue et al. (2005) recorded the following
mangrove losses is also evaluated. species of Bruguiera from Singapore:
23 Mangrove Biogeography of the Indo-Pacific 381

B. ­sexangula, B. gymnorhiza, B. parviflora and 1986; Duke and Kudo 2018). More recently it
B. cylindrica, as well as the new record of seemed that buoyant propagules are better
Bruguiera hainesii, which has also recently been adapted to longshore and interisland dispersal
recorded from northeastern Queensland (Cooper rather than intercontinental transmission by oce-
et al. 2016). anic currents (Clarke 1993; McGuiness 1997).
The distributional limits of Rhizophora spe- Mangrove dispersal is thus limited in different
cies have been extended in the southwestern
Pacific by Duke (2010), and new records of
Table 23.1 Indo-Pacific mangrove hybrids with putative
Sonneratia ovata and S. lanceolata, together with parental species
the hybrids S. × urama, S. × gulngai and
Combretaceae
Rhizophora × annamalayana, have been made
Lumnitzera × rosea (L. littorea × L. racemosa)
from the Andaman and Nicobar Islands, India Lamiaceae
(Ragavan et al. 2016). Unnamed hybrid (Avicennia marina × A. rumphiana)
Considering the above taxonomical results, Lythraceae
the mangrove flora of the world is updated and Sonneratia × gulngai (S. alba × S. caseolaris)
listed in Appendix 1. The mangrove niche is Sonneratia × hainanensis (S. alba × S. ovata)
inhabited by around 81 tree and shrub species of Sonneratia × urama (S. alba × S. lanceolata)
30 genera from 17 families worldwide. Eleven Unnamed hybrid (Sonneratia alba × S. griffithii
Pteridaceae
named and unnamed hybrids (Table 23.1) and
Unnamed hybrid (Acrostichum aureum × A. speciosum
several subspecies/varieties of mangroves have
Rhizophoraceae
been described. Hybrids are not discussed further Bruguiera × rhynchopetala (B. gymnorhiza × B.
because they are usually of limited distribution sexangula)
and, as they occur within the range of the putative Unnamed hybrid (Ceriops tagal × C. australis)
parental species, they do not offer a great deal of Rhizophora × annamalayana (R. apiculata × R.
biogeographic information. The same goes for mucronata)
varieties and subspecies. For further information *Rhizophora × harrisonii (R. mangle × R. racemosa)
Rhizophora × lamarckii (R. apiculata × R. stylosa)
on mangrove hybrids, see the recent review by
Rhizophora × mohanii (R. mucronata × R. stylosa)
Ragavan et al. (2017). Recently Ono et al. (2016)
Rhizophora × selala (R. stylosa × R. mangle)
reported on the hybrid origin of B. hainesii (B. Rhizophora × tomlinsonii (R. apiculata × R. mangle)
gymnorhiza × B. cylindrica), although here it is Notes: The hybrid Rhizophora × brevistyla was reduced
considered as distinct species until further confir- into synonymy with R. mangle by Hou (1960). Similarly,
mation is available. Hou (1960) reduced Rhizophora samoensis into synon-
ymy with R. mangle. This has been followed by later
authors, e.g. Breteler (1977), Tomlinson (1986) and Tyagi
(2003), although Duke (2010) maintains its species status.
 iogeography of Indo-West Pacific
B There have been suggestions that Rhizophora harrisonii is
Mangroves a hybrid between R. mangle and R. racemosa (Keay 1953;
Breteler 1969; Wilcox 1985), but data from foliar wax
chemistry did not support any hybrid status in that R. har-
The species diversity and distribution of man- risonii showed no intermediate characteristics from those
groves are variable at different spatial scales, i.e. of its putative parental species (Dodd et al. 1995). More
global, regional, estuarine and intertidal. On a recent molecular data, however, supported the hybrid
global scale, mangrove distribution appears to be nature of this taxon (Cerón-Souza et al. 2010). Recently
Ono et al. (2016) reported on the hybrid origin of
limited by temperature, while at a regional scale, Bruguiera × hainesii (B. gymnorhiza × B. cylindrica):
various biotic and abiotic factors control their here, because of its high fruit set, it is considered as a dis-
distribution. In the past, it was assumed that tinct species until more data are forthcoming
because of long distance dispersal by buoyant Hybrids confined to the Atlantic East Pacific only are indi-
cated with an *
propagules, mangrove species were widely and Finally, since this paper was prepared another hybrid
continuously distributed in suitable habitats mangrove has been described from north-east Australia
within the region of their occurrence (Tomlinson (Duke and Kudo (2018)
382 P. Saenger et al.

ways by both land barriers and wide expanses of Excoecaria ovalis are restricted to Australasia,
water, which in turn cause the disjunct pattern of and Camptostemon philippinense, Ceriops zippe-
contemporary mangrove distribution. As dis- liana and Aegiceras floridum are restricted to
cussed below, recent phylogeographic studies, South East Asia. Both Kandelia obovata and
using molecular methods, also have shown the Acanthus xiamenensis are found only in East
existence of disjunction in populations of the Asia.
mangrove species throughout the Indo-West
Pacific.
The well-known disjunction pattern of man-  egional Distribution of Mangrove
R
groves is the species-poor Atlantic East Pacific Species Around the Indo-Pacific
(AEP) and species-rich Indo-West Pacific (IWP).
Despite the equivalent area of mangrove habitat Based on recent taxonomic and distributional
in each region, mangroves of the IWP are three records discussed above and the personal knowl-
times more diverse than in the AEP. Mangroves edge of the authors, the distribution pattern and
of IWP consist of 69 mangrove species, whereas gradients in five geographical regions of the IWP
the AEP region has 13 species (Appendix 1). (East Africa, Middle East, South Asia, South East
Only one species (Acrostichum aureum) and Asia, East Asia and Australasia) and of the East
three genera (Acrostichum, Avicennia and Pacific are summarized in Tables 23.2, 23.3, 23.4,
Rhizophora) have a common distribution in both 23.5, 23.6 and 23.7.
the IWP and the AEP. Except for the family The mangroves of the East Pacific coast are
Tetrameristaceae (formerly Pelliceriaceae), all listed in Table 23.2, which has been compiled
mangrove families of the AEP are also repre- from data given in Breteler (1969), Gentry
sented in the IWP. Difference in species richness (1982), Jiménez (1987), Cerón-Souza et al.
between the IWP and the AEP is evidence of the (2005, 2010), Spalding et al. (2010), Barreto and
effectiveness of the African land mass and the Barreto-Pittol (2012), Castillo-Cárdenas et al.
East Pacific Ocean barriers. (2015) and Santamaria-Damián et al. (2016).
On the whole, 20 out of 25 genera are com- Detailed data on the distribution of mangroves
mon in the IWP. The exceptions are Aglaia, and associated species are given by Ellison
Brownlowia, Camptostemon, Diospyros and (2004) for Central America, Sandoval-Castro
Osbornia. Diospyros is restricted to Australasia, et al. (2014) for Mexico and Gentry (1982) and
and Aglaia, Brownlowia and Phoenix are Cerón-Souza et al. (2005) for Colombia.
restricted to South and South East Asia. No natural mangroves occur in the Hawaiian
Camptostemon and Osbornia are restricted to archipelago, but various species have been intro-
South East Asia and Australasia. The mangrove duced including Rhizophora mangle, R. mucro-
flora of South and South East Asia shares 42 spe- nata, Bruguiera sexangula, B. parviflora, Ceriops
cies, of which 11 species (Acanthus volubilis, tagal and Conocarpus erectus. Of these species,
Aglaia cucullata, Brownlowia tersa, Ceriops only R. mangle, B. sexangula and C. erectus have
decandra, Excoecaria indica, Heritiera fomes, established self-maintaining populations (Allen
Kandelia candel, Phoenix paludosa, Sonneratia 1998). Other introduced species in Eastern
apetala, Sonneratia griffithii and Aegialitis rotun- Polynesia include Rhizophora stylosa on
difolia) are not found in Australasia. Australasia Mo’orea, Tahiti and Bora Bora, Rhizophora man-
shares 40 species with South East Asia, of which gle on Enewetak Atoll and Bruguiera gymno-
9 species (Aegialitis annulata, Avicennia rhiza on Bikini Atoll (Saenger 2002).
rumphiana, Bruguiera hainesii, Brownlowia The mangroves of the East African region and
argentata, Camptostemon schultzii, Ceriops aus- the Middle East consist of 10 and 4 species,
tralis, Ceriops pseudodecandra, Osbornia respectively. Species occurrences are listed in
octodonta and Bruguiera exaristata) are restricted Tables 23.3 and 23.4, respectively. They have
to South East Asia and Australasia. Three species been compiled from data given in Béguinot
Avicennia integra, Diospyros littorea and (1918), Dale (1938), Koechlin et al. (1974),
23 Mangrove Biogeography of the Indo-Pacific 383

Table 23.2 Mangrove species distribution in the East Pacific region

Species Mex Gua ElS Hon Nic CRi Pan Col Ecu Per
Acrostichum aureum * * * * * * * *
Acrostichum danaeifolium * * * * * * * *
Avicennia bicolor * * * * * * * *
Avicennia germinans * * * * * * * * * *
Conocarpus erectus * * * * * * * * * *
Laguncularia racemosa * * * * * * * * * *
Mora oleifera * *
Pelliciera rhizophorae * * * * *
Rhizophora mangle * * * * * * * * * *
Rhizophora racemosa * * * * * * * *
Tabebuia palustris * * *
Species totals 7 7 7 8 9 9 11 11 8 5
Mex Mexico, Gua Guatemala, EIS El Salvador, Hon Honduras, Nic Nicaragua, CRi Costa Rica, Pan Panama, Col
Colombia, Ecu Ecuador, Per Peru

Table 23.3 Mangrove species distribution of the East African region

Species SAf Moz Mad Tan IOI Ken Som Dji
Acrostichum aureum * * * *
Avicennia marina * * * * * * * *
Bruguiera gymnorhiza * * * * * * *
Ceriops tagal * * * * * * * *
Heritiera littoralis * * * * *
Lumnitzera racemosa * * * * * * *
Pemphis acidula * * * *
Rhizophora mucronata * * * * * * *
Sonneratia alba * * * * * *
Xylocarpus granatuma * * * * *
Species totals 6 10 9 10 10 8 6 2
SAf South Africa, Moz Mozambique, Mad Madagascar and Mauritius, Tan Tanzania, IOI Indian Ocean islands of
Comoros, Mayotte and the Seychelles, Ken Kenya, Som Somalia including the island of Socotra, Dji Djibouti
a
Only X. granatum is listed in Spalding et al. (2010), but Dale (1938) mentions ‘the two species of Xylocarpus’ in the
Kenyan mangroves, listing both X. granatum (as X. benadirensis) and X. moluccensis. Other records of X. moluccensis
are from Somalia (Ciferri 1939), Tanzania (Mangora 2011) and Madagascar (Koechlin et al. 1974; Hughes and Hughes
1992)

Zahran (1977), Ruwa (1993), Banyikwa (1986), atively simple and feature a decline in species
Ormond et al. (1988), Cornes and Cornes (1989), richness with increasing latitude associated with
Hughes and Hughes (1992), Faye (1993), temperature and/or aridity.
Spalding et al. (2010), Kumar et al. (2010a, b, South Asia (also known as Indian subconti-
2011), Moore et al. (2015) and Almahasheer nent) represents approximately 7% of the global
(2018). Limited areas of suitable habitat and a mangrove area and hosts 42 species belonging to
prevalence of harsh environmental conditions 22 genera. The mangroves of South Asia are
limit extensive growth of mangroves in these listed in Table 23.5, which have been compiled
regions. Mangroves of this region are considered from data given in Naskar and Mandal (1999),
as a subset of the highly diverse mangroves of Spalding et al. (2010) and Ragavan et al. (2016).
other regions of the IWP (Duke et al. 1998), and No endemic species of mangroves are known
no endemic species have been reported. from South Asia. Maximum number of species
Distributional gradients in this region appear rel- occurs in mainland India (37 species), with a
384 P. Saenger et al.

Table 23.4 Mangrove species distribution in the Middle East

Species Eri Sud Egy SAr Yem Oma UAE Qat Bah Kuw Ira
Avicennia marina * * * * * * * * * ** *
Bruguiera gymnorhiza * * * * **
Ceriops tagal *
Rhizophora mucronata * * * * ** *
Species totals 3 2 3 2 3 1 1 1 1 0 3
Eri Eritrea, Sud Sudan, Egy Egypt, SAr Saudi Arabia, Yem Yemen, Oma Oman, UAE United Arab Emirates, Qat Qatar,
Bah Bahrain, Kuw Kuwait, Ira Iran
**
Introduced

marked decline in species numbers with the possible dispersal route between these South
increasing distance from the continental land
­ Asian countries and mainland India through the
mass (e.g. Sri Lanka, the Maldives and the Strait of Malacca to the shores of the Bay of
Chagos Archipelago). Different geomorphologi- Bengal. However, the disjunct distribution of
cal settings between west and east coasts of India Lumnitzera littorea in this region is difficult to
and absence or presence of river deltas are explain: it is restricted to Sri Lanka and ANI but
responsible for the greater species richness of the is not found on either coasts of mainland India.
east coast. None of the species on the west coast The mangroves of the East Asia are listed in
is unique whereas several species (Scyphiphora Table 23.6, which has been compiled from data
hydrophylacea, Acrostichum speciosum, given in Li and Lee (1997), Hsueh and Lee (2000)
Aegialitis rotundifolia, Aglaia cucullata, and Spalding et al. (2010). The most notable fea-
Heritiera fomes, Brownlowia tersa, Pemphis ture of this region is the high level of species rich-
acidula, Nypa fruticans, Phoenix paludosa and ness across the region, with a rapid loss of species
Acanthus volubilis) are restricted to east coast of to the north-east (China and Japan) as a result of
India. unfavourable temperature regimes (Li and Lee
Another important mangrove region in South 1997; Chen et al. 2017). These trends are dis-
Asia comprises the Andaman and Nicobar Islands cussed further below.
(ANI) that host 34 mangroves species. The east The mangroves of the Pacific and Australasian
coast of these islands, in particular, has well-­ region are listed in Table 23.7, which have been
developed contiguous mangroves. Rhizophora compiled from data given in Saenger et al. (1977),
stylosa, Lumnitzera littorea, Sonneratia ovata, Maguire and Saenger (2000), Duke (2006),
Sonneratia griffithii and Sonneratia lanceolata Ellison (2009), and Spalding et al. (2010). The
are restricted to the ANI in South Asia. Earlier continental land masses in this region (Australia
reports of the occurrence of Rhizophora stylosa and Papua New Guinea) show high levels of spe-
and Sonneratia griffithii on the east coast of India cies richness and a degree of endemism
are erroneous. It is often noted that floristically (Avicennia integra, Diospyros littorea and
ANI has a close similarity with Southeast Asian Excoecaria ovalis), but the various island groups
countries due to its geographical proximity. to the south and east show a rapid decline in spe-
However, mangrove floristics of ANI show a cies numbers, reflecting both dispersal limita-
closer similarity with mainland India, sharing tions due to distance and limited availability of
only five species with South East Asia suitable mangrove habitat. This is particularly
(Rhizophora stylosa, Lumnitzera littorea, well illustrated by the tropical Christmas Island
Sonneratia ovata, Sonneratia griffithii and (10° 25′S; 105° 40′ E), lying some 360 km south
Sonneratia lanceolata). In contrast, there is high- of Java with its diverse mangroves and with an
est similarity between mainland India and the annual rainfall in excess of 2000 mm, which has
Southeast Asian countries like Thailand, four recorded mangrove species. Clearly, disper-
Myanmar, Malaysia and Singapore. It indicates sal distance and adverse currents, together with
23 Mangrove Biogeography of the Indo-Pacific 385

Table 23.5 Mangrove species distribution in South Asia restricted availability of mangrove habitats, are
Species Pak Ind Mal ANI Sri Ban the major limiting factors.
Acanthus ebracteatus * *
Acanthus ilicifolius * * * *
Acanthus volubilis * * Contemporary Biogeographical
Acrostichum aureum * * * * *
Processes Affecting Mangrove
Acrostichum speciosum * *
Aegialitis rotundifolia * *
Distributions
Aegiceras corniculatum * * * * *
Aglaia cucullata * * A much-debated question concerning mangroves
Avicennia alba * * is where they first arose and how they got where
Avicennia marina * * * * * * they are today (Duke 1995; Ellison et al. 1999).
Avicennia officinalis * * * * Whatever the exact historical origin(s) and dis-
Brownlowia tersa * * persal routes of mangroves, the present distribu-
Bruguiera cylindrica * * * * tions of mangroves show several features
Bruguiera gymnorhiza * * * * * * reflecting some of the more modern processes
Bruguiera parviflora * *
described below (Saenger 1998). These modern
Bruguiera sexangula * * * *
constraints are manifested as a reduction of man-
Ceriops decandra * * *
Ceriops tagal * * * * * * grove species with increasing latitude on the one
Cynometra iripa * * * hand and selective removal or loss of species by
Dolichandrone * * * natural upheavals or human activity on the other.
spathacea The latitudinal limits of mangrove vegetation
Exceocaria agallocha * * * * * on the major land masses of the Indo-Pacific
Excoecaria indica * * * * (Table 23.8) show that these limits are quite vari-
Heritiera fomes * * able. They can, however, be broadly related to
Heritiera littoralis * * * * *
temperature and/or aridity (Saenger and Moverley
Kandelia candel * *
Lumnitzera littorea * *
1985; Quisthoudt et al. 2012; Chen et al. 2017). It
Lumnitzera racemosa * * * * should also be noted that there is a gradual reduc-
Nypa fruticans * * * * tion of mangrove development (e.g. height and
Pemphis acidula * * * * extent of mangrove vegetation) with latitude
Phoenix paludosa * * (Saenger and Snedaker 1993) and that an attenu-
Rhizophora apiculata * * * * * * ation of species with increasing latitude precedes
Rhizophora mucronata * * * * * * these latitudinal limits. By way of example, the
Rhizophora stylosa * * southward attenuation of species on the non-arid
Scyphiphora * * *
east coasts of Africa and Australia is shown in
hydrophylacea
Sonneratia alba * * *
Table 23.9. This comparison indicates that on
Sonneratia apetala * * continental coasts where mangroves are not lim-
Sonneratia caseolaris * * * * * * ited by aridity, temperature is the major factor in
Sonneratia griffithii * * reducing number of species with latitude. Clearly
Sonneratia lanceolata * while there is some intraspecific variability
Sonneratia ovata * between the two continents, the data support the
Xylocarpus granatum * * * * suggestion that, in the presence of adequate rain-
Xylocarpus * * * fall, latitude as a proxy for temperature is related
moluccensis
to the southern limits of mangrove species distri-
Species totals 8 39 13 34 24 24
bution. Similar limitations apply to northern dis-
Pak Pakistan, Ind India, Mal Maldives, ANI Andaman and
Nicobar Islands, Sri Sri Lanka, Ban Bangladesh tributional limits (Quisthoudt et al. 2012; Chen
et al. 2017).
386 P. Saenger et al.

Table 23.6 Mangrove species distribution of East Asia

Species Mya Mal Sin Indo Tha Cam Vie Phi Chi Jap
Acanthus ebracteatus * * * * * * * * *
Acanthus ilicifolius * * * * * * * * *
Acanthus volubilis * * * * *
Acanthus xiamenensisa *
Acrostichum aureum * * * * * * * * * *
Acrostichum speciosum * * * * * * *
Aegialitis annulata *
Aegialitis rotundifolia * * *
Aegiceras corniculatum * * * * * * *
Aegiceras floridum * * * *
Aglaia cucullata * * *
Avicennia alba * * * * * * * *
Avicennia marina * * * * * * * * * *
Avicennia officinalis * * * * * * *
Avicennia rumphiana * * * *
Brownlowia argentata * * * *
Brownlowia tersa * * * * * * *
Bruguiera cylindrica * * * * * * * *
Bruguiera exaristata *
Bruguiera gymnorhiza * * * * * * * * * *
Bruguiera hainesii * * * * * *
Bruguiera parviflora * * * * * * *
Bruguiera sexangula * * * * * * * * *
Camptostemon philippinense * *
Camptostemon schultzii * *
Ceriops australis *
Ceriops decandra * * * * * * *
Ceriops tagal * * * * * * * * *
Ceriops pseudodecandra *
Ceriops zippeliana * *
Cynometra iripa * * *
Dolichandrone spathacea * * * *
Exceocaria agallocha * * * * * * * * * *
Excoecaria indica * * * * *
Heritiera fomes * *
Heritiera littoralis * * * * * * * * * *
Kandelia candel * * * * * * *
Kandelia obovata * * *
Lumnitzera littorea * * * * * * * * *
Lumnitzera racemosa * * * * * * * * * *
Nypa fruticans * * * * * * * * * *
Osbornia octodonta * * *
Pemphis acidula * * * * * * * * *
Phoenix paludosa * * *
Rhizophora apiculata * * * * * * * * * *
Rhizophora mucronata * * * * * * * *
Rhizophora stylosa * * * * * * *
Scyphiphora hydrophylacea * * * * * * * *
Sonneratia alba * * * * * * * * * *
(continued)
23 Mangrove Biogeography of the Indo-Pacific 387

Table 23.6 (continued)

Species Mya Mal Sin Indo Tha Cam Vie Phi Chi Jap
Sonneratia apetala * **
Sonneratia caseolaris * * * * * * * * *
Sonneratia griffithii * * *
Sonneratia lanceolata *
Sonneratia ovata * * * * * * * * *
Xylocarpus granatum * * * * * * * * *
Xylocarpus moluccensis * * * * * *
Species totals 36 43 36 50 37 25 34 38 25 11
Mya Myanmar, Mal Malaysia, Sin Singapore, Indo Indonesia, Tha Thailand, Cam Cambodia, Vie Vietnam, Phi The
Philippines, Chi China, Jap Japan
**Introduced
a
It has been suggested that Acanthus xiamenensis is synonymous with A. ilicifolius, but it is retained here until further
data becomes available

The distribution of mangrove species richness nating higher and lower salinities (Voris 2000). In
is shown in Fig. 23.1, based on data from Spalding turn, these processes have led to bottleneck and
et al. (2010) as contained at the portal site called founder effects on the abundant mangroves of the
TroCEP (Tropical Coastal Ecosystem Portal): region and led to high rates of speciation.
http://www.nies.go.jp/TroCEP/index.html. While The second identifiable trend in Fig. 23.1 is
the nature of the data has some limitations (e.g. the attenuation of species numbers with increas-
Western Australia has 20 mangrove species listed ing latitude generally related to limiting tempera-
for the entire ~21,000 km of coastline, where tures. Thus, the rapid loss of species on the east
mangrove species numbers range from 20 in the and west coast of Australia, in southern Africa
north to 1 in the south) and the algorithm has dif- and along the shores of the East China Sea can be
ficulties modelling mid-ocean rather than near- appreciated. The rapid decline in species on the
shore species numbers, Fig. 23.1 nevertheless northern shores of East Africa and into the Red
shows two important biogeographical trends. Sea, Gulf of Oman and Arabian Gulf is almost
First, the distribution of mangrove species certainly due to limiting aridity rather than tem-
richness across the IWP clearly shows that maxi- perature. An attenuation of species numbers from
mal richness occurs along the shorelines of continental land masses to offshore archipela-
Makassar Strait, between Borneo and Sulawesi in goes is also discernible although less marked.
Indonesia. This area has the greatest interdigita- Other contemporary biogeographical pro-
tion of land and sea in the world, a condition cesses result from human activities such as pollu-
stretching back to the Holocene. While some sea- tion, water diversion, selective clearing and, most
ways may have closed due to tectonic or sea-level recently, major mangrove afforestation or reha-
changes, the area has always maintained some bilitation programmes, with or without novel
direct connections between the tropical Pacific species introductions. These are briefly reviewed
and tropical Indian oceans as Makassar Strait lies below.
more or less seaward of the Sunda Shelf. High
precipitation and relatively low salinities in this
area also provided favourable conditions for the Historical Biogeographical
development and maintenance of the mangrove Processes Affecting Mangrove
niche. Furthermore, cycles in sea level during the Distributions
numerous late Pliocene and Pleistocene glacial-­
interglacial intervals have led to the appearance Historical biogeographic processes involve
and disappearance of islands, the openings and changing the genetic diversity of populations
closing of marine corridors and bridges and alter- directly or via altered or redirected gene flow
388 P. Saenger et al.

Table 23.7 Mangrove species distribution in the Pacific and Australasian region
Species Mar MIs Kir Sam Fij PNG NCa NZ Au CI
Acanthus ebracteatus * *
Acanthus ilicifolius * * *
Acrostichum aureum * *
Acrostichum speciosum * * *
Aegialitis annulata * *
Aegiceras corniculatum * *
Avicennia alba *
Avicennia integra *
Avicennia marina * * * *
Avicennia officinalis *
Avicennia rumphiana *
Brownlowia argentata * *
Bruguiera cylindrica * *
Bruguiera exaristata * *
Bruguiera gymnorhiza * * * * * * * * *
Bruguiera hainesii * *
Bruguiera parviflora * *
Bruguiera sexangula * * *
Camptostemon schultzii * *
Ceriops australis *
Ceriops pseudodecandra * *
Ceriops tagal * * *
Cynometra iripa * *
Dolichandrone spathacea * * *
Diospyros littorea * *
Exceocaria agallocha * * * * *
Excoecaria ovalis *
Heritiera littoralis * * * * *
Lumnitzera littorea * * * * * *
Lumnitzera racemosa * * *
Nypa fruticans * *
Osbornia octodonta * *
Pemphis acidula * * * * *
Rhizophora apiculata * * *
Rhizophora mucronata * * *
Rhizophora stylosa * * * *
Scyphiphora hydrophylacea * * *
Sonneratia alba * * * * *
Sonneratia caseolaris * * *
Sonneratia lanceolata * *
Sonneratia ovata *
Xylocarpus granatum * * * * * *
Xylocarpus moluccensis * *
Species totals 3 5 4 3 5 39 18 1 38 4
Mar Marianas Islands, MIs Marshall Islands, Kir Kiribati, Sam Samoa, Fij Fiji, PNG Papua New Guinea, NCa New
Caledonia, NZ New Zealand, Au Australia, CI Christmas Island

between populations. Thus, for example, loss accompanied by a reduction in genetic diver-
or fragmentation of habitat is usually associ- sity: Parani et al. (1997) found that Avicennia
ated with a reduction in population size that is marina in India, under stress from intense
23 Mangrove Biogeography of the Indo-Pacific 389

Table 23.8 Northern and southern limits of mangrove From their study of three populations of
vegetation on major land masses of the Indo-Pacific
Ceriops zippeliana (as C. decandra) on the west,
Continental land mass Northern limit Southern limit east and southern coasts of Malaysia, Tan et al.
Pacific America 30° 15′ 5° 30′ (2005) found that the three populations showed a
Red Sea/Southern 30° 31′ 32° 59′ high level of genetic differentiation and, when
Africa
Western Australia – 33° 16′
grouped per geographic regions, there was a clear
Eastern Australia – 38° 45′ differentiation between populations from the
Pacific Asia 31° 21′ – eastern Indian Ocean (West Malaysia) from those
of the western Pacific Ocean (East and Southern
Malaysia). They suggested that this split resulted
Table 23.9 Southern latitudinal limits of shared man- from the historical lowering of sea level near the
grove species on the eastern coasts of Australia and Africa Strait of Malacca during the recent Pleistocene
E. African E. Australian glaciations. Similar findings were also reported
Species Southern limit Southern limit for Ceriops tagal (Huang et al. 2012).
Sonneratia alba 23° 55′ 22° 30′ Similarly, Wee et al. (2017) and Yang et al.
Pemphis 24° 18′ 24° 01′ (2017) have shown that the widely distributed
acidula mangrove Sonneratia alba has very restricted
Xylocarpus 26° 01′ 25° 30′ gene flow due to various genetic barriers and geo-
granatum
Heritiera 26° 33′ 22° 15′
graphic distance. Thus the northern Australian
littoralis population is genetically differentiated from pop-
Ceriops tagal 26° 50′ 28° 11′ ulations on New Caledonia and in South East
Lumnitzera 26° 50′ 27° 30′ Asia, while the populations of Mozambique, the
racemosa Bay of Bengal and southern Japan are also dis-
Acrostichum 28° 58′ 26° 05′ tinctive. They concluded that the effects of oce-
aureum
anic barriers due to sea-level change were
Rhizophora 31° 42′ 18° 25′
mucronata compounded by the fact that Sonneratia alba
Bruguiera 32° 14′ 29° 25′ propagules have a limited dispersal distance.
gymnorhiza Minobe et al. (2010) examined the genetic
Avicennia 32° 59′ 38° 45′ diversity of Bruguiera gymnorhiza and found
marina that genetic diversity was low within any local
population and that differentiation between the
Pacific Ocean, Bay of Bengal and the Arabian
grazing and pollution, showed reduced levels Sea was the result of very low gene flow between
of polymorphism. each of these regional seas, coupled with frequent
Restricted gene flow between populations can fluctuation of population size due to changes in
be caused by such factors as changing sea levels, sea level. In the West Central Pacific, the genetic
closing of marine seaways, alterations in current diversity of Bruguiera gymnorhiza was signifi-
patterns or distance. Given that eight major cantly higher in specimens from Iriomote Island,
glacial-­interglacial intervals have occurred dur- Okinawa, than on the more remote Pohnpei
ing the last 3 million years (Late Pliocene and Island, Micronesia (Sugaya et al. 2003). Dispersal
Pleistocene) with sea-level changes ranging from limitations related to propagule viability over
−70 to 90 m (Rossi 1981), changing configura- extended distances appear to be the major cause.
tion of land masses has been significant (Voris Fujimoto et al. (1996) have investigated what
2000). The emergence during Pleistocene times constitutes the critical rise of sea level affecting
of the Sunda and Sahul shelves would have con- mangroves. They found that with gradual sea-­
stituted major barriers to gene flow both to the level rise of 1–2 mm/year over the last 2000 years,
north-west and to the south-east. extensive mangrove forests developed on Kosrae
390 P. Saenger et al.

Fig. 23.1 Approximate distribution of mangrove species richness in the Indo-West Pacific. (Based on data from
Spalding et al. (2010) in http://www.nies.go.jp/TroCEP/index.html. Species size classes are based on a geometrical
progression)

Island, Micronesia, by accumulating mangrove 2000; Arnaud-Haond et al. 2006; De Ryck et al.
peat (Fujimoto et al. 1996). However, during the 2016), Rhizophora stylosa (Islam et al. 2014),
period of rapid sea-level rise of around 10 ­mm/ Kandelia obovata (Chen et al. 1996), Bruguiera
year that occurred between 4100 and 3700 year gymnorhiza (Islam et al. 2006) and Sonneratia
BP, the mangrove forest ceased peat accumula- alba (Wee et al. 2017). Thus, the findings of De
tion and retreated landward, suggesting that the Ryck et al. (2016) on the east African coast high-
critical rate of sea-level rise enabling mangrove light the genetically impoverished situation in
peat accretion on islands was between 2 and peripheral populations due to the rarity of long
10 mm/year. distance dispersal, followed by inbreeding and
The distributions of the two species of dispersal limitation due to the coastal geomor-
Kandelia offer a good example of vicariance, phology and the availability of suitable habitats.
where two populations develop more or less allo- Interestingly, in their study of Sonneratia alba,
patrically with no or very limited gene flow Wee et al. (2017) clearly demonstrated the range
between them (Kado et al. 2004). However, edge effect by regressing Ho (observed heterozy-
because of the recent divergence times of many gosity) and FIS (inbreeding coefficient) against
pantropical taxa (including mangroves, e.g. latitude: Ho decreased with increasing latitude,
Avicennia; see Li et al. 2016), it is increasingly and FIS increased with increasing latitude.
recognized that long distance dispersal might be A summary of where some of the processes
the more important process determining the pres- described above have been identified in man-
ent distribution of many pantropical taxa. grove species of the IWP is shown in Table 23.10
Several mangrove species show strong and Fig. 23.2. This summary indicates that the
inbreeding and self-fertilization, particularly at South China Sea and the Strait of Malacca, with
range edge, e.g. Avicennia marina (Maguire et al. their generally shallow depths and seasonally
23 Mangrove Biogeography of the Indo-Pacific 391

Table 23.10 Areas of restricted gene flow in the Indo-West Pacific identified from molecular studies of mangroves
Location Indicative lat/long Species Presumed cause References
1. SWIO 30°S/33°E A. marina Dispersal limitations and habitat De Ryck et al.
availability (2016)
2. Arabian 14°N/74°E B. Bottlenecks during glaciation Minobe et al.
Sea gymnorhiza (2010)
3. WCIO 10°S/65°E S. alba Dispersal limitations Wee et al. (2017)
4. Malacca 1°N/103°E B. Bottlenecks during glaciation Minobe et al.
Str. gymnorhiza (2010)
C. tagal Dispersal limitations Huang et al. (2012)
C. Glaciation Tan et al. (2005)
zippeliana
S. alba Glaciation Yang et al. (2017)
S. alba Dispersal limitations Wee et al. (2017)
5. SCS 10°N/110°E C. tagal Bottlenecks during glaciation Ge and Sun (2001)

S. alba Glaciation Yang et al. (2017)

A. marina Dispersal limitations Kado et al. (2004)
K. candel Dispersal limitations Kado et al. (2004)
6. WCPO 20°N/140°E B. Dispersal limitations Sugaya et al.
gymnorhiza (2003)
7. Timor Sea 10°S/127°E S. alba Land barrier during glacial periods Wee et al. (2017)
C. tagal Dispersal limitations Huang et al. (2012)
8. Coral Sea 20°S/154°E S. alba Dispersal limitations Wee et al. (2017)
C. tagal Dispersal limitations Huang et al. (2012)
SCS South China Sea, WCIO Western Central Indian Ocean, SWIO South-West Indian Ocean, WCPO Western Central
Pacific Ocean

reversing ocean currents, are significant areas of Friess 2016). For example, in eastern Africa,
demonstrated restricted gene flow for several uncontrolled wood extraction, particularly of
mangrove species and are worthy of further Rhizophora, Bruguiera and Ceriops – which,
investigation for other IWP mangroves. together with Heritiera, are used for construc-
tion – has adversely affected many mangrove
areas. Avicennia marina, which is widely used for
Contemporary Anthropogenic firewood and charcoal, has also been affected sig-
Factors Affecting Mangrove nificantly (Ngoile and Shunula 1992; Rasolofo
Distributions 1993). Along the Arabian peninsula, mangrove
wood has been used for building huts and boats,
From the brief descriptions below, it is obvious for making bird traps and for firewood while the
that the major anthropogenic factors affecting foliage has been grazed by camels and been used
mangroves and their distribution are overexploi- for fodder. Over-exploitation as camel fodder
tation, land use changes, hydrological alteration around the town of Djibouti has been documented
and pollution. (Faye 1993).
Giri et al. (2010) have shown that the total
mangrove extent has in recent years declined to
Overexploitation 137,760 km2, some 12% below the latest esti-
mates by the Food and Agriculture Organization.
Around the world, mangroves have been overex- Of even more concern is that of much of the
ploited (Blasco et al. 2001; Zorini et al. 2004; remaining areas of mangroves, a large proportion
Hauff et al. 2006; Mangora 2011; Richards and is degraded and that undisturbed, dense
392 P. Saenger et al.

Fig. 23.2 Approximate distribution of mangrove species richness in the Indo-West Pacific with the numbered locations
(as per Table 23.10) of those areas where restricted gene flow has been demonstrated for several mangrove species

­ angroves have disappeared from many parts of

m Saltworks commenced in the Rufiji Delta,
the world (Blasco et al. 2001). Tanzania, in 1930. However, had spread in the
1970–80 period and by 1989, there were 3100 ha
of saltponds in the mangroves, many of which
Land Use Changes were poorly constructed. The mangroves have
been most affected in Bagamoyo, Dar es Salaam,
Conversion of mangroves for agricultural pur- Tanga, Mtwara and Lindi.
poses is widespread in the Indo-West Pacific. For In parts of the Middle East, the principal con-
example, in the Rufiji Delta in Tanzania, man- cern has been coastal infill and construction work
grove conversion to rice commenced in the in and around mangroves, sometimes on a very
1960s, which was ultimately prohibited by law in large scale (Vousden and Price 1985). Infilling
1987. While initially good harvests were and reclamation works are still occurring in many
obtained, ultimately rice fields decreased produc- parts of the world (e.g. Tarut Bay, Saudi Arabia).
tivity and were abandoned. Rice cultivation on Possibly the largest effect on mangroves has
the landward edge of the mangroves in arisen from the construction of aquaculture facil-
Madagascar has also been attempted on a large ities in the mangroves. In countries like Indonesia,
scale (e.g. in the delta of the Tsiribihina), but it is India, Thailand, Vietnam and China, massive
now largely abandoned because of the saliniza- destruction of mangroves occurred in the 1980s
tion of the soil. Salt ponds also exist in the delta and beyond. In 1978 the first shrimp mariculture
as well as shrimp culture, causing the loss of attempts were made in Kenya supported by UN
extensive mangrove areas (Rasowo 1992; agencies. With average production levels of
Rasolofo 1993). 525 kg/ha/year, the Ngomeni aquaculture pilot
23 Mangrove Biogeography of the Indo-Pacific 393

project caused the destruction of 60 ha of man- Quwain Bay, United Arab Emirates, in January
groves in Ngomeni Creek (Rasowo 1992). 1998, can still affect considerable mangrove
Similarly, shrimp farming in the Mekong Delta areas – in this case some 20 km of coastline
of Vietnam increased massively in the 1990s and (Youssef et al. 2000).
became unsustainable due to the unplanned
development, the self-pollution of farms, the out-
break of viral diseases and the destruction of Reversing the Anthropogenic
mangroves (de Graaf and Xuan 1998). Changes

Reducing Habitat Loss

Hydrological Alteration
With the current annual loss of mangrove area at
Mangrove losses caused by changed hydrologi- 0.4% around the world (Hamilton and Casey
cal conditions are widespread in the IWP (Ghosh 2016), there is increasing interest in implement-
et al. 2015; Haque and Reza 2017). On the other ing better management of existing healthy man-
hand, localized losses have also been reported. grove areas. Rogers et al. (2016) suggested
For example, near the port on Khor Farasan, several actions that could improve protection of
Saudi Arabia, a causeway has adversely affected coastal wetlands in Australia and the ecosystem
a large area of Avicennia marina, and it appears services they provide. As these actions seem
that initially insufficient culverts were provided. appropriate more widely, they are briefly listed
As a result, some mangrove mortality resulted. and discussed. The suggested initiatives include
More culverts were subsequently inserted, and benchmarking and improving coastal wetland
the surviving mangroves now appear to be doing extent and health through restoration and/or reha-
well. Mandura and Khafajii (1993) concluded bilitation, reducing complexity and inconsistency
that 63.3% of the trees died upstream of the in governance arrangements and facilitating wet-
causeway. land adaptation and ecosystem service delivery
using a range of relevant mechanisms. They sug-
gested that actions that build on the momentum
Pollution to mitigate climate change by sequestering car-
bon could achieve multiple desirable objectives
Adverse effects of pollution on mangroves have such as climate-change mitigation and adapta-
been widely reported throughout the IWP (Vane tion, floodplain rehabilitation and habitat protec-
et al. 2009; Chowdhury et al. 2017). Of least con- tion. Clearly for mangroves, such an approach
cern is pollution by sewage, which the mangroves should include the establishment or extension of
appear to tolerate reasonably well. On the other mangrove green belts to provide a natural, self-­
hand, heavy metal pollution has been found to repairing barrier for coastal protection, as well as
reduce genetic diversity in Avicennia marina the afforestation of suitable coastal areas with
(Manurung et al. 2017). They found that genetic mangroves (Saenger 2011).
diversity in this species was inversely propor-
tional to pollution levels in three estuaries, rang-
ing from non-polluted to moderately and highly  everse Trend of Mangrove Loss
R
polluted. by Rehabilitation
In contrast to the Red Sea, the mangroves of
the Arabian Gulf coast are extremely limited in López-Portillo et al. (2017) have recently reviewed
their distribution and were adversely affected by mangrove restoration or rehabilitation projects
the large oil spill from the Gulf War (Pashaei around the world and concluded that, with detailed
et al. 2015). Smaller oil spills, e.g. at Umm-Al-­ and site-specific project design, mangrove restora-
394 P. Saenger et al.

tion/rehabilitation schemes can offset some man- formance was better with plants in mixed stands
grove losses and replace some of the ecosystem than in monospecific ones (Das et al. 1997).
services that may have been reduced or lost during The deltaic Chokoria Sundarbans, situated
mangrove deterioration. They provided a frame- south of Chittagong on the Bangladesh east coast,
work for planning and implementation of restora- were badly degraded due to a proliferation of
tion projects, as well as monitoring and reporting shrimp cultivation and salt farms. With the
protocols. A similar framework, together with a removal of mangroves, shrimp cultivation
case study of the restoration of the Chokoria became unsustainable, and most farms closed,
Sundarbans of Bangladesh, was described by leaving the area vulnerable to frequent cyclones
Biswas et al. (2009). and tidal surges. Considering the run-down con-
To date, insufficient research has focused on ditions of the site, an area-specific restoration and
the establishment and maintenance of restoration community plan was prepared by IUCN
projects, and accumulated practical experience Bangladesh, with active participation of the local
has not been widely disseminated. In addition, community. While some initial success was
innovative research in this area has only just reported, the necessary database to assess pre-
commenced. For example, El-Tarabily and and post-restoration conditions was not realized
Youssef (2011) have recently shown that bacteria due to funding shortages, and it was likely com-
capable of producing high levels of 1-amino-­ munity interest would wane (Biswas et al. 2009).
cyclopropane-1-carboxylic acid deaminase, such The project nevertheless assisted with developing
as Pseudoalteromonas maricaloris, have the a useful planning framework.
potential as biological inoculants to promote Das (2017) assessed the contribution of
growth of mangrove seedlings in rehabilitation Avicennia marina plantations in the Indian state
projects in nutrient-poor sediments in hypersa- of Gujarat to the inshore and offshore fisheries’
line coastal areas. sectors and found that the mangrove plantations
had significantly increased the catch of mangrove-­
dependent fish in both sectors. Despite these
Some Examples of Success Stories plantations being monospecific and sparse, they
nevertheless contributed around US$ 0.57 billion
Many mangrove restoration/rehabilitation projects to the fishery sector of the state.
have been undertaken, but there are a few studies
that have evaluated the flow of ecosystem services
from such regenerated ecosystems (see Field Importance of Mangroves,
1996; Primavera and Esteban 2008). It is only Seagrasses and Saltmarshes
recently that attempts have been made to under-
stand and quantify restoration trajectories by With annual mangrove losses of around 0.4% and
which success or otherwise of restoration/rehabili- other halophytes declining at similar rates, it seems
tation projects can be usefully compared (Twilley timely to reconsider the importance of halophytes
and Rivera-Monroy 2005; Aung et al. 2011; Salmo in general and mangroves in particular.
et al. 2013; López-Portillo et al. 2017).
Mangrove restoration around Paradip port of
Orissa, India, was undertaken using ten man- Trophic Significance
grove species that had been nursery-raised.
Planted over a 10 ha degraded wetland area in Early studies suggested that mangroves provided
pure and mixed stands, survival rates greater than the primary carbon source fuelling detrital-based
70% were found for S. apetala, A. officinalis, R. food webs, and therefore they represented the
mucronata, K. candel and H. fomes. Sonneratia major route of energy flow in such systems.
apetala recorded maximum growth in height However, mangrove litter dynamics, however,
(3.0 m after 2 years), and generally, growth per- have shown that all mangrove systems are highly
23 Mangrove Biogeography of the Indo-Pacific 395

variable and that the export of organic matter is at local communities. Such activities have even
best about one-third of the total organic matter higher benefit-cost ratios with the inclusion of
production. More importantly, stable isotope the indirect benefits resulting from the avoided
analyses have investigated the real pathways of maintenance cost for the sea dike system that the
energy flow in mangrove ecosystems and have mangrove stands protect from coastal storm
progressively shown that the contribution of surges. Thus, Tri et al. (1998) concluded that a
mangroves to many species in the estuarine food strong case for mangrove rehabilitation could be
web was highly variable, but not as significant as made as an important component of a sustainable
previously believed (Loneragan et al. 1997; coastal management strategy. It should also be
Thimdee et al. 2004, 2008; Taylor et al. 2018). noted that the correlation between those coastal
Mangrove foliage is nutritious and has been communities not protected by mangrove forests
used as fodder. For example, mangrove propa- and the resulting high loss of life and property as
gules of Avicennia marina have been used as a a result of the 2004 tsunami was significant and
substitute for corn in commercial feed for finger- has led to enhanced restoration attempts of man-
lings of bluespot mullet (Valamugil seheli) at grove habitat along several Indian Ocean shore-
rates of up to 300 g kg−1 (Belal 2004). Use of lines (Erwin 2009; Saenger 2011).
mangrove propagules rather than dietary corn led
to increased growth, body moisture, ash and pro-
tein and a reduction in body fat. Biotechnology Potential
of Halophytes

Ecosystem Services Mangroves have large potential as sources of

novel agrochemicals and compounds of medici-
Mangroves provide a range of ecosystem ser- nal value (Bandareanayake 2002). Also, man-
vices of considerable economic value (Saenger groves help with the immobilization of heavy
2002; Hussain and Badola 2008; Iftekhar and metals in mangrove sediments (Chowdhury et al.
Takama 2008; Walters et al. 2008; Van 2017). Because of the ability of mangroves to tol-
Oudenhoven et al. 2014; Friess 2016). In the erate and bioremediate the high levels of nutri-
Sundarban mangroves of Bangladesh, Rahman ents and pollutants found in sewage water, they
et al. (2018) found that mangrove fodder have been proposed as a simple and effective
accounted for 2% of the provisioning services alternative to conventional sewage treatment,
obtained by 100 households in 3 villages and that particularly given their low cost and low mainte-
fishery capture, fuel energy and honey accounted nance (Lambs et al. 2011). Mangroves have also
for 86%, 7% and 4.7%, respectively, of an annual been found to be tolerant of, and to accumulate,
total of US$1135 per ha. high concentrations of trace metals and to thus
The significant role of mangroves in carbon minimize metal movement to adjacent ecosys-
sequestration has recently been highlighted as tems (Vane et al. 2009; Chowdhury et al. 2017).
part of the climate-change discussions. Mangrove Finally, mangroves and halophytes in general
ecosystems contain some of the largest carbon carry the genetic material that has given them
stocks quantified to date (Rahman et al. 2015). their salt-tolerance capability. While the physio-
Using an economic cost-benefit model, Tri logical and biochemical basis of salt tolerance is
et al. (1998) showed that mangrove rehabilitation only gradually being elucidated, there is a grow-
undertaken, inter alia, to enhance sea defence ing appreciation of some of the genetic complexi-
systems in three coastal districts of northern ties involved. The possibility of using mangrove
Vietnam, is desirable from an economic perspec- proteins or genes to enhance the salt resistance of
tive based solely on the direct use benefits by our crop plants is increasingly promising.
396 P. Saenger et al.

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