Vol.

15 · July
IAMURE 2015
International Journal of Ecology and Conservation
Print ISSN 2244-1573 · Online ISSN 2244-1581
International Peer Reviewed Journal
doi: [Link]
Journal Metrics: H Index = 1 from Publish or Perish
This journal is included in Thomson Reuters Journal Masterlist
Published by IAMURE Multidisciplinary Research,
an ISO 9001:2008 certified by the AJA Registrars Inc.

Mangrove Ecosystems and Associated

Fauna with Special Reference to Mangrove
Crabs in the Philippines: A Review
GINALYN C. CUENCA
[Link]
djinaleene8@[Link]
Davao Medical School Foundation
Davao City, Philippines

EDISON D. MACUSI
[Link]
edmacusi@[Link]
Aquaculture and Fisheries Group, Wageningen Institute of Animal Sciences,
Wageningen University of Life,
Wageningen, The Netherlands

NEIL ANGELO S. ABREO

[Link]
[Link]@[Link]
Davao Medical School Foundation
Davao City, Philippines

CYRIL TOM B. RANARA

[Link]
tomranara74@[Link]
Davao Del Norte State College
Panabo City, Philippines

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 Volume 15 · July 2015

MICHAEL B. ANDAM
[Link]
michael_andam@[Link]
Davao Del Norte State College
Panabo City, Philippines

LEMUEL C. CARDONA
[Link]
cardona_lemuel@[Link]
Davao Del Norte State College
Panabo City, Philippines

GRACE C. GUANZON
[Link]
graceconservaguanzon@[Link]
Davao Del Norte State College
Panabo City, Philippines

Gunning Fog Index: 11.79 Originality: 100% Grammar Check: 92%

Flesch Reading Ease: 41.39 Plagiarism: 0%

ABSTRACT

Mangrove ecosystems are very important because of its many ecological and
economic functions. It provides food, shelter, and protection to a myriad of
organisms and maintains connectivity with other coastal ecosystems. It holds
diverse forms of both floral and faunal associations, which are used by humans
for different purposes. This review discusses the physiognomy, ethnobiology,
composition of mangrove ecosystems with emphasis on associated fauna and
mangrove crabs, fish production, and the human threats that can alter the
stability of mangrove ecosystems. Different journal articles, books, manuals,
and other online sources from both local and international sources were used to
discuss the review. Mangrove ecosystems were found to be occurring in tropical
and subtropical areas and are distributed horizontally in estuarine and vertically
in intertidal zones due to a number of environmental and biotic factors. The
ecosystem is composed of diverse associations of communities of both floral and

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faunal groups. Many of these associated fauna are benthic macroinvertebrates

and fishes, a number of which are sourced as fish products. Fisheries are the
topmost economic use of mangrove ecosystems. Fish products harvested include
mud crabs, shrimps, mollusks, and fishes. The continued growth of human
population and the industrialization of fishing exceeded our sustainable use of
waters leading to the declining fish landings. Such event gave rise to aquaculture,
which has become the main cause of diminishing mangrove ecosystems. Other
human threats to mangrove ecosystems include mining, overexploitation for
economic services, and urban development. Along with mangrove decline is the
ecosystem’s susceptibility to climate change particularly on the rising sea level.
This indicates a higher risk to flooding, tsunamis, cyclones, and storm surges.
Hence, protection, conservation, rehabilitation and restoration of mangrove
ecosystems, and its sustainable use are essential. Eco-friendly aquaculture must
also be implemented.

KEYWORDS

Mangrove, fauna, flora, mud crab, aquaculture, fisheries, literature review,

Philippines

INTRODUCTION

Mangrove ecosystem is composed of mangrove forest community and

its associated abiotic factors (Kathiresan & Bingham, 2001). Such factors are
salinity (Long & Giri, 2011), tidal inundation (Naidoo, 2006; Rezende, Pinto,
Gonçalves Jr, & Petrucio, 2013), and soil characteristics, which are waterlogged
and oxygen deficient (Duke, Ball & Ellison, 1998; Reef, Feller & Lovelock, 2010).
Other than mangrove trees, the forest community includes microbes, fungi,
plants, and animals (Kathiresan & Bingham, 2001). Generally, the ecosystem is
situated in the tropics and subtropics (Long & Giri, 2011) traversing between
30°N and 30°S latitudes (Morrisey et al., 2010). Many of the mangrove trees
display adaptive features to fit in its ecosystem. These include modified roots for
breathing and support (Duke et al., 1998; Giesen, Wulffraat, Zieren & Scholten,
2007), low water potentials and high salt dilution within cells, salt excretion and
water storing mechanisms in leaf tissues, and buoyant and viviparous propagules
for wider range of distribution and successful initiation in mangrove forests,
respectively (Duke et al., 1998).

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Mangrove ecosystems are known for their productivity because of their large
reservoir of carbon from roots and leaves with mean net primary productivity
approximates of 2 to 5 mg C ha−1 year−1 (Alongi, 2009) making them the primary
nutrient source for many organisms from different trophic levels (Bouillon,
Connolly & Lee, 2008). Mangrove plants are projected to distribute 10% of its
carbon into the coastal waters (Allan & Castillo, 2007) through its leaf litter fall
(Bouillon et al., 2008). Carbon may as well reach offshore communities (Nerot,
Meziane, Provost-Govrich, Rybarczyk & Lee, 2009) by means of mangrove
outwellings as influenced by tidal inundation (Lee, 1995).
Although the productivity of mangroves is high, nutrient availability is low
due to its soil characteristics. On account of this, preservation of nutrients in such
ecosystem is vital to achieving efficient nutrient cycling and maintaining high
productivity. One important way is through the feeding activity of invertebrates
in which mangrove leaf litter is consumed and is returned to the nutrient-poor
mangrove soil as fecal matter (Reef et al., 2010). Such invertebrates include
various crabs (Nordhaus, Wolff & Diele, 2006; Nordhaus & Wolff, 2007) like
sesarmids (Sheaves & Molony, 2000), brachyurans (Nordhaus, Diele, & Wolff,
2009), grapsid crabs (Nerot et al., 2009), and fiddler crabs (Nicholson, 2009).;
gastropods; and worms. Moreover, detritivores feed on other forms of detritus
materials (Sheaves & Molony, 2000; Nerot et al., 2009).
High productivity in mangrove habitats needs to be sustained as it is the
most common type of ecosystems used as nurseries next to coral reefs (El-Regal
& Ibrahim, 2014). It provides temporary or permanent dwelling ( Rönnbäck,
Troell, Kautsky & Primavera, 1999) for numerous reef fishes and invertebrates
not only for its copious food supply, but also for its protective root structures
(Honda, Nakamura, Nakaoka, Uy, & Fortes, 2013; El-Regal & Ibrahim, 2014),
and for growth (Jones, Walter, Brooks & Serafy, 2010). This signifies the role
of mangrove ecosystems in providing habitat connectivity with other important
marine ecosystems. This was already observed in New Caledonia, Zanzibar
(Paillon, Wantiez, Kulbicki, Labonne & Vigliola, 2014), Biscayne Bay mangrove
shore (Jones et al., 2010), and in Western Pacific (Solomon Islands, Great Barrier
Reef, and Moreton Bay, Australia) (Olds, Albert, Maxwell, Pitt & Connolly,
2013). In Egyptian Red Sea, nearly 72% of juvenile fishes found in mangrove
habitats are coral reef fishes and practically 85% are of economic value such as
the snappers, lizardfish, milkfish, goatfish, barracudas, and rabbit fish ( El-Regal
& Ibrahim, 2014). This indicates the importance of interconnectivity between
mangrove and reef ecosystems in furthering efficient reef reserves (Olds et al.,
2013). Moreover, mangroves also revealed connectivity with seagrass beds as
observed in Indo-Pacific. The study revealed that seagrass beds in proximity with

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mangroves have more numerous fishes and a number of which are economically
important fishes (Unsworth et al., 2008).
Moreover, mangroves have a lot of economic uses to humans (Rezende et
al., 2013, Mukherjee et al., 2014; Paillon et al., 2014). In fisheries alone, various
seafood products are harvested in mangrove areas such as mud crabs, shrimps,
mollusks, and fishes (Primavera, 2005; Giesen et al., 2007). In Southeast Asia,
77% of all mangrove species are known for its many uses, from which 41% is for
medicinal use, 25% for construction materials, 22% for food and consumption,
17% for ornamental use, and at least 12% for fuel (Giesen et al., 2007). Moreover,
mangrove ecosystems can also be utilized for ecotourism and education (Giesen
et al., 2007). Among other countries, Philippines utilize mangrove forests for fish
production (Brander et al., 2012), fuel (J.H. Primavera, 2000; J.H Primavera,
2000), and for responsible tourism (Miranda et al., 2013).
Other important functions of mangrove trees are coastal protection
(Primavera, 2000) and serving as bio-indicators (Anouti, 2014). Mangrove forests
act as barriers to waves, wind, and cyclones by means of trapping sediments,
and reducing and breaking waves (Primavera, 2000; 2005; Giesen et al., 2007).
They also filter the bodies of water serving as sinks for sequestering pollutants
and contaminants carried by tidal inundations from different sources of waters
(Anouti, 2014).
Taken together, mangrove ecosystems are vastly important for their ecological
and economic functions (Figure 1). Thus, it is important that such ecosystems
are protected and preserved.

OBJECTIVES OF THE STUDY

The objectives of this study are to discuss the physiognomy and ethnobiology
of mangrove ecosystems, the composition of its ecosystem particularly on its
associated communities with emphasis on mangrove crabs that will be related
to the utilization of mangrove ecosystem for fish production and for other
anthropogenic activities that can alter the stability of mangrove ecosystems, and
so affecting the organisms in it and humans.

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Figure 1. Important Functions of Mangrove

METHODOLOGY

Different journal articles from local and international sources, books, manuals,
and other online sources ([Link]; Google Scholar; Pubmed Central; Academia.
edu; Elsevier; Sciencedirect; Plos One; Springer; Researchgate; and Philippine
e-journals) were used to accomplish the objectives of this review paper. The words
“mangrove ecosystem”, “mangrove associated fauna and flora”, “crab production”,
“mangrove crabs”, and “status of mangroves” were used as search terms in the
online sources. This literature review is presented in a thematic approach, which
is started with a discussion on the significant functions of mangrove ecosystem,
followed by its distribution pattern. The different associated communities were
examined and crabs were given emphasis since they are considered to be important
mangrove fisheries that are harvested and sold to the market and at the same
time they are keystone species in mangrove forests. The review also discussed the
threats of human activities that will affect the function and structure of mangrove
ecosystems, which affects the associated communities.

RESULTS AND DISCUSSION

Mangrove Physiognomy
The distribution of mangroves is evaluated based on its global scale, regional
pattern, the locality within estuaries, and on intertidal setting (Duke et al., 1998).

A. Scale
Mangroves are dispersed worldwide in two hemispheres, the Indo-West
Pacific and Atlantic East Pacific, from which the former propagated more species

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of mangroves in a sum of 58 taxa than the latter with only 13 species. In regional
scale, mangroves occur in six locales including Western Americas and Eastern
Pacific, Eastern Americas and Caribbean, Western Africa, Eastern Africa and
Madagascar, Indo-Malesia and Asia, and Australia and the Western Pacific (Duke
et al., 1998). Global distribution of mangroves has extended to approximately
137, 760 km2 in 2000 with Asia holding the largest cover of mangroves that
equals to 42% and South America having the smallest reaching only 11%. Next
to Asia is Africa with 20% mangroves, followed by North and Central America
with 15% (Giri et al., 2011).

B. Latitude
Mangrove species dispersal in both global and regional scales is limited by
latitude (Morrisey et al., 2010). This declining distribution of mangroves at higher
latitudes is in parallel with the latitudinal diversity gradient pattern of many
plant and animal species in both global hemispheres, and marine and terrestrial
environments (Mittelbach et al., 2007). The latitudinal diversity gradient pattern
is described by many ecologists as the increase of species diversity towards the
equator and the decrease in the direction of increasing latitudes due to some
factors, one of which is climate (Rohde, 1992; Rosenzweig, 1992; Mittelbach
et al., 2007) as observed in terrestrial environment or solar energy in marine
environment (Roy, Jablonski, Valentine & Rosenberg, 1998). Thus, mangrove
species predominantly cultivate in the tropics and subtropics residing in latitudes
between 30°N and 30°S (Morrisey et al., 2010; Giri et al. 2011).

C. Climatic Factors

1. Temperature
Mangrove species dispersal in both global and regional scales is also limited
by climatic factors such as low and intense temperatures, patterns of rainfall,
and aridity (Morrisey et al., 2010) with temperature as the main determinant of
patterns of distribution (Smith & Duke, 1987; Duke et al., 1998). Mangroves
are situated in regions with average annual temperature of about 18 oC or higher
or with an estimated absolute temperature of greater than 15oC (Giesen et al.,
2007). Geographical regions with more than 20°C of average air temperature
during months of coldest temperatures and with seasonal extent of 10oC and
less sets the limit to mangrove distribution, indicating that global and regional
distribution of mangroves is constrained in low and intense temperatures (Duke
et al., 1998). High temperatures were observed to have unfavorable effects on
two mangrove species in Duyen Hai District in Ho Chi Minh City Vietnam

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such as curtailed physiologic events in R. apiculata affecting its photosynthetic

and transpiration activities at a temperature as high as 40 oC (air) or 42 oC (soil),
and even death to Avicennia propagules when exposed to a temperature of 39 oC
to 40 oC (Hong & San, 1993).
Nevertheless, some species were observed to have grown in temperate regions
with dry summer, and without dry season climates, in which the summer
temperature can be hot with a temperature higher than 22 °C during the hottest
summer, or can be warm with a temperature within 4 oC to 10 °C; and a few
located in areas of higher latitudes like in Bermuda, southern Kyushu in Japan,
Southern South Africa, Southern Australia, and North New Zealand, which only
documented one to three species of mangroves. Such species are the Aegiceras
corniculatum, A. marina, Bruguiera gymnorrhiza, Kandelia candel, R. mucronata,
A. germinans, and R. mangle (Morrisey et al., 2010). Heritiera littoralis has low
tolerance on rising temperature of about 26 oC, whereas A. marina can traverse
up to 38 oS latitude (Smith & Duke, 1987) as in Southern Australia, which is
positioned at 38 °45′S latitude (Morrisey et al., 2010); however its growth is
restricted in areas with a temperature of 37 oC (Hastuti, Anggoro & Pribadi,
2012). Among all mangrove species, A. marina is the most widespread (Morrisey
et al., 2010). In the Philippines, A. marina is dominant in several provinces
including Calatagan, Batangas (Lunar & Laguardia, 2013), Looc, Romblon
(Sabigan, Garcia, & Race, 2013), Palompon, Leyte (Picardal, Avila, Tano, &
Marababol, 2011), and in Badian, Cebu (Valenzuela, Bacalso, Gano, Pilones, &
Picardal, 2013).

2. Rainfall
In the western coasts of Africa and South America, and probably Western
Asia and Middle East, another characteristic of climate called aridity limits the
distribution of mangrove species, which is why there is no record that shows
occurrence of these species in the two former regions mentioned. An arid
biogeographic region has a precipitation greater than 30 mm on any month of the
year with an expected evaporation proportion of less than 0.03 (Morrisey et al.,
2010). Similar climatic factor, aridity (seasonal), was proposed to have caused the
absence of Bruguiera parviflora, Camptostemon schultzii, and Lumnitzera littorea
in northern and western portions of Australia, but this has remained inconclusive
yet for lacking proofs on quantitative data (Smith & Duke, 1987).
The absence of mangrove species in the western coasts of Africa and South
America can likewise be attributed to the patterns of rainfall (Morrisey et al.,
2010). In terms of temperature and degree of rainfall, mangroves are typically
restricted in regions of lower temperatures, and typically more species are located

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in regions with higher rainfalls. With these, species richness is highest in regions
with bigger size of estuaries with moderate to higher rainfall and regulated
salinity. Rainfall additionally influences the suitability of the species that can lead
to distinctions of attributes like plant height and density (Duke et al., 1998).
The most diverse mangrove ecosystem in the Philippines is found in Bohol with
26 to 34 species. One of its study sites was recorded to have an average annual
temperature of 28oC and an average annual rainfall of 1 500 mm to 2 000 mm
(Middeljans, 2014). This clearly shows that temperature and the degree of rainfall
are important factors that influence mangrove distribution.
According to Hong and San (1993), freshwater is important for mangroves
to reach its prime growth thru the regulation of salt concentration both in the
plant’s body and in the soil, and one source of freshwater other than the water
from the rivers and runoffs, is rainfall. This is the reason the densest mangrove
forests are positioned in Southeast Asia since it is showered with heavy rains in
summer periods (Hong & San, 1993). As denoted in the hypothesis of Wells
(1985), brackish water, which is a mixture of fresh and salt water, grows more
mangrove species than freshwater. As an example, Ceriops tagal does not occur
in mangrove areas in Northwestern Australia where mangroves are exposed to
freshwater the whole year. While, in the northern coast of Queensland higher
species richness is observed in estuaries with more freshwater, which oppose the
observations in northern Australia. In Western Australia, mangroves that collect
groundwater have higher species richness (Smith & Duke, 1987).
Both temperature and precipitation preceded the classification of four areas
based on the occurrence of mangroves. These are the warm, humid, sub-humid,
semi-arid, and arid areas. From these four areas, the warm, humid areas contain
most of the mangrove species in the global and regional scale reaching up to 90%
like in areas of South Mexico to Colombia, in Caribbean, North Brazil, and
Southeast Asia to North Queensland Australia, while arid areas have none with
the exclusion of the Red Sea, Persian Gulf, Gulf of California, and Ethiopian
coasts. In sub-humid and semi-arid areas, mangroves are sporadic and rare,
respectively. Sub-humid areas include East Africa, India, South Queensland
Australia, Mexico, and Venezuela; on the other hand, semi-arid areas are the
Indus Delta in Pakistan, Gujarat in India, Ecuador, and Western and Northern
Australia (Duke et al., 1998).

D. Propagule and Sapling Viability

Global and regional distribution of mangroves is reliant on the ability of
propagules (Duke et al., 1998) and sapling to institute in the areas (Hastuti et al.,
2012), which considers the characteristics of propagules, its environment, and

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of its geographic features. Several factors influence the dispersal of propagules

through the bodies of water and between land masses (Duke et al., 1998), and
the successful establishment of saplings would expand geographic extent of
mangroves (Hastuti et al., 2012). One factor is the buoyancy and sustainability
of propagules in the water, which varies among mangrove species (Duke et al.,
1998). Propagules need to be established during their floating time since they
sink when floating time ends, and they can successfully establish in the absence
of tidal disturbances (Rabinowitz, 1978). Floating time of A. marina is shorter,
lasting up to few days only similar to Laguncularia racemosa that can be viable
in the water for less than 8 days (Duke et al., 1998) with the emergence of its
embryonic roots within 5 to 8 days, followed by its positive institution. In a
Costa Rican estuary, the establishment of Avicennia propagule was restricted by
the presence of recurrent tidal inundation (Delgado, Hensel, Jiménez & Day,
2001).
On the other hand, Rhizophora sp. and A. germinans can be viable for
months, ranging from 14 days to 110 days in the case of A. germinans (Duke et
al., 1998). In a study conducted by Delgado et al. (2001), A. germinans instituted
effectively by 97% in his “no flooding by tides” treatment than in the treatment
flooded by tides with only 67% success level in propagule institution. Bruguiera
gymnorrhiza seems to be the most efficacious in terms of dispersal since it spans
all through Indo-West Pacific (Duke et al., 1998). However, based on the study
of Duke et al. (1998), influences of tidal properties on mangrove distribution
is not yet known but in terms of species richness it has been observed to show
some effects like in tidal amplitudes (Smith & Duke, 1987), which has reduced
propagule establishment in Quang Ninh and northern Hai Pong Vietnam when
tidal amplitude has reached three to four meters.
In central Vietnam and northwestern Ca Mau, propagules have grown well
as they are exposed only to minor sizes of tides (Hong & San, 1993). The size
of propagules also varies, and the smaller the propagule like in Avicennia and
Laguncularia, the more that it requires an absence of tidal disturbances. This
would mean that mangroves with smaller propagules establish in areas with
lesser tidal inundation, which is landward, and those with larger propagules
like in Rhizophora and Pelliciera establish in lower ground or seaward that is
more disturbed by tides, indicating the relationship between propagule dispersal
properties and spatial distribution of mangroves (Rabinowitz, 1978). The
physical characteristic of the environment is another factor that affects propagule
establishment. The buoyancy of propagules increases with lower temperature and
salinity of the water, while cooler temperatures and the absence of appropriate
locales hinder the successful cultivation of mangrove propagules (Duke et al.,

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1998). One example is Avicennia, which is predisposed to abortive growth and

development of sapling with unsuitable environments (Hastuti et al., 2012).
Surface water currents influence propagule dispersal thru its degree and direction
as caused by climatic factors (Duke et al., 1998) or as directed by the winds
(Hong & San, 1993). Geographic barriers such as land masses and extensive
stretch of water impede effective dispersal of propagules (Duke et al., 1998).
Similar to global and regional scale, propagule properties influence dispersal
in intertidal zones. In the study of Delgado et al. (2001) in Costa Rica estuarine,
Avicennia and Laguncularis propagules effectively established by 66% and
73%, respectively, in lower intertidal zone, whereas in upper intertidal zones,
success rate is 51% for Avicennia and 18% for Laguncularis. Avicennia propagule
establishment was attributed to the degree of tidal flooding present, while in
Laguncularis, it was attributed to selective feeding of crabs on the species
propagules that resulted to its higher mortality in different intertidal zones than
in Avicennia spp. (Delgado et al., 2001).

E. Salinity
Mangroves are distributed horizontally and vertically in estuarine and
intertidal zones, respectively (Primavera et al., 2012) as influenced by several
factors including salinity, tidal features, substrate characteristics, (Giesen et
al., 2007, Primavera et al., 2012) and waterlogging (Duke et al., 1998). The
horizontal pattern of distribution separates mangrove species into downstream,
intermediate, and upstream (Duke et al., 1998; Primavera et al., 2012) that vary
in their degree of salinity with 27% to 28% salinity in the downstream, 1% to
17% salinity in intermediates, and with 0% to 4% in the upstream (Duke et al.,
1998).
Salinity influence the distribution of mangroves by affecting the plant’s growth
rates, which occur higher in areas with lower salinity level. Optimal growth of
mangroves is achieved in low salinity zones since carbon is apportioned more on
the leaf tissues than in the roots generating a higher shoot to root ratio (Duke et
al., 1998). Root biomass increases as more thin roots form, which is necessary to
improve nutrient uptake in saline soils (Reef et al., 2010). High levels of Na+ in
soils impair the procurement of other ions like K+, Mg2+, and Ca2+ (Naidoo, 2006;
Chen & Ye, 2014) and hinder nitrate uptake. When an ionic imbalance occurs,
other internal processes are affected like protein synthesis, conduction of stomata
(Chen & Ye, 2014), hydraulic conductivity, and carbon dioxide exchange. In A.
marina, the photosystem II of photosynthesis was damaged because of exposure
to excessive salinity and light (Naidoo, 2006). Hence, in general, growth rate
is slower in higher salinity conditions (Duke et al., 1998), which can lead to

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dwarfing of some species like A. marina as observed by Naidoo (2006).

Other than decreasing growth rates, high salinity also causes several adverse
effects to mangrove species. In a study conducted on Excoecaria agallocha, a
salinity above 15 psu resulted to decrease or inhibition of seed germination,
increase leaf senescence, ionic imbalance, ion toxicity, altered metabolic process
especially photosynthesis, and oxidation damage; whereas salinity of 25 psu
caused weakened leaf production, failure on seedling establishment, and mortality
of seedling (Chen & Ye, 2014). Seedling initiation and growth are crucial factors
since it stretches the extent of mangrove species in its geographic distribution
(Hastuti et al., 2012).
Mangrove species respond differently on high salinity soils and some species
are salt tolerant due to the presence of adaptive mechanisms like in the case of
Aegiceras corniculatum, which contains leaf glands that excrete salt making its leaf
petiole and surface appear whitish. This species also requires higher salinity of
2 – 4% to attain its prime growth, while R. mucronata requires 1 to 2% salinity
for growth (Giesen et al., 2007). Avicennia seedling attain its optimal growth in 5
to 30% salinity (Hastuti et al., 2012). Other species like Sonneratia caseolaris can
tolerate both high and low salinity soil (Giesen et al., 2007). In relation to this,
mangroves that grow in the downstream such as A. alba, A. marina R. mucronata,
R. stylosa (Hong & San, 1993), R. mangle, and A. germinans, and Sonneratia
alba are the most salt-tolerant, while upstream species such as S. caseolaris, S.
lanceolata, and Mora are the least salt-tolerant. Species in intermediates are A.
bicolor, Pelliciera, and R. racemosa (Duke et al., 1998). In the Philippines, A.
avicennia, A. alba, and S. alba are the downstream species while R. mucronata,
S. caseolaris, Xylocarpus granatum, and Nypa fruticans are the intermediate to
upstream species (J. Primavera et al., 2012). Root structures also vary between
species and different zones. Normally pneumatophores are found in mangroves
situated at low intertidal zone, while prop and knee roots, and buttress or plank
roots are frequently found in mid-intertidal and high intertidal zones, respectively
(Samson & Rollon, 2010).

F. Tidal Inundation
Vertical distribution of mangroves in intertidal zones partitions mangrove
species into three zones, the low, mid, and high intertidal zones (Duke et al.,
1998, Giesen et al., 2007) or into five zones in Southeast Asia (Giesen et al.,
2007) based on the degree of tidal inundation that cause variations in salinity,
waterlogging, and faunal features (Duke et al., 1998). In the three zonation
pattern, the low intertidal zone is the most frequently inundated and flooded
with intermediate high currents for more than 45 times per month. Next to

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this is the mid-intertidal zone, being inundated and flooded with normal high
currents for more than 20 times but less than 45 times per month, while high
intertidal zone is the least flooded zone being drenched in water for less than
20 times per month only (Duke et al., 1998). Conversely, in the five zonation
pattern; zone 1 is highly subjected to seawater and tidal regime during every high
tide; zone 2 is subjected to seawater and flooded by medium high tides; zone
3 is the mid zone; zone 4 is a landward zone that is only flooded with spring
tides; and lastly, zone 5 is exposed to brackish water to nearly freshwater with
occasional flooding (Primavera et al., 2012).
Flooding causes mangrove soil to become anoxic or have a low level of oxygen,
and so root biomass is decreased or in other species, root tips may even die (Reef
et al., 2010). In terms of substrate, nearly all mangrove species grow best on
muddy soils, while a number establish well on sandy and coral islands containing
shells, coral and seaweed remains (Giesen et al., 2007), and some propagate
on sandy to muddy substrates (Primavera et al., 2012). In Southeast Asia, R.
mucronata, and Bruguiera grow well on muddy soils (Giesen et al., 2007) and in
the Philippines too with the addition of Ceriops decandra, Xylocarpus (Primavera
et al., 2012), S. alba, A. marina (Lunar & Laguardia, 2013), and Nypa (Juario &
Ontoy, 2005). In Indonesia, R. stylosa and S. alba commonly establish on sandy
and rocky substrates as well (Giesen et al., 2007). A. marina occurs in various
types of substrates, whereas A. alba together with S. alba and R. stylosa grow on
sandy to muddy substrate (Primavera et al., 2012). S. alba, R. stylosa, and C. tagal
dominated in Badian, Cebu with sandy to muddy substrate (Valenzuela et al.,
2013).

G. Biotic Factors
Associated faunal activities is another factor that influence the zonal
distribution of mangroves. Examples of these are sesarmid crabs and mollusks
that were observed to inhibit propagule establishment of Avicennia by damaging
its propagules, thereby displacing this mangrove genus in areas dominated by
such associated fauna (Duke et al., 1998). A similar effect to Avicennia was also
observed in Australia caused by the feeding behavior of grapsid crabs (Osborne
& Smith, 1990; Clarke & Myerscough, 1993) and wallabies (Clarke &
Myerscough, 1993). Insects, on the other hand, are detrimental to the seeds and
propagules of other mangrove species, particularly those species with hard seed
capsules like Heritiera and Xylocarpus. Weevils that form burrows in Rhizophora
zone show comparable effects (Duke et al., 1998). Macroalgae like Hormosira
banksii also revealed detrimental effects, lessening the initiation of mangrove
seedling on mudflats shrouded with the said macroalgae (Clarke & Myerscough,

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1993). On the other hand, burrowing activities of associated fauna may limit
or enhance mangrove growth. As an example, crustaceans like crabs improve
substrate oxygenation by means of such activity and its burrows may serve as
establishment area for the propagules of R. mangle that are hindered to initiate
in its densely arranged root networks (Duke et al., 1998). Moreover, wind was
also observed to influence the distribution of mangrove propagules depending
on the buoyancy and morphological characteristics of propagules. In accordance
to this, propagules with lower density and with a floatable shape like in the case
of H. littoralis, which floats on the surface of water and displays a sailboat-like
structure, can be dispersed more by wind and water currents (Stocken et al.,
2013). R. mucronata has denser propagules, and so it is not distributed well by
water currents resulting to its institution in the front rows (Perera, Amarasinghe
& Somaratna, 2013).
In addition, some mangrove species situated in the front rows like Rhizophora,
enhance initiation of other mangroves by providing a stable coastline conducive
for growth (Duke et al., 1998).

Mangrove Ethnobiology

A. Fisheries Production
The topmost economic use of mangrove forests is fisheries (Mukherjee et al.,
2014). Fishing has become one of the major income-generating activities like
in West Africa (Satyanarayana et al., 2012). Fisheries products are harvested in
mangrove forests for food in numerous countries (Brander et al., 2012) including
those in Southeast Asia (Giesen et al., 2007). Such fisheries products are mud
crabs, shrimps, mollusks, and fishes (Primavera, 2005; Giesen et al., 2007). In
the ASEAN region, annual coastal fish catches of mangrove associated fishes,
crustaceans, and mollusks comprise 21% of the total catch. From this, the
associated fishes make-up 30% of the yearly catches of finfish, while associated
prawns constitute nearly 100% of the entire prawn resources in ASEAN
(Primavera, 2005).

B. Crab Production in Mangroves

Mud crabs (Scylla) are widely collected in Asia-Pacific both as local food and
for export, which has led to overfishing in some countries. In Matang Mangrove
Forest Reserve in Perak, Malaysia, S. serrata and S. olivacea are harvested and
distributed in the market with S. serrata as the main harvest. These mangrove
crabs are sold in the market in different sizes from 7.5 cm to more than 10 cm
in carapace width, indicating that even small crabs are bought by consumers,

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which is attributed to its cheaper price (Kosuge, 2001). In Mtwapa Kenya, S.

serrata crabs were stocked in a pen culture system that is situated in the intertidal
area. The crabs were enclosed by a fish trap and were collected after four months
(Mwaluma, 2002). In the Philippines, mud crab production has already become
prevalent in numerous provinces (Bureau of Fisheries and Aquatic Resources,
2014). In 1996, production has reached 2 440.00 metric tons and increased
further to 4 945.00 metric tons in 2000 (Figure 2), 6 836.50 metric tons in 2005
(CountrySTAT Philippines, 2006-2015), and 14 436.45 metric tons in 2010
(Bureau of Fisheries and Aquatic Resources, 2010; CountrySTAT Philippines,
2006-2015). The highest production by volume occurred in 2012 with 16
359.63 metric tons, which had a value of 4, 822, 360.34 pesos (CountrySTAT
Philippines, 2006-2015).

Figure 2. Mud crab production in brackish fishpond from 1996 to 2014

(Source: CountrySTAT Philippines 2006-2015)

The Associated Flora and Fauna in Mangrove Ecosystems

Mangrove ecosystem serves as habitat for various associated communities of
flora (Kathiresan & Bingham, 2001) and fauna (Superales & Zafaralla, 2008),

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which are composed of diverse organisms that dwell in various mangrove niches
such as on the surface of substrate, underneath the substrate, on fallen leaves, on
trunks, and in the water (Giesen et al., 2007; Superales & Zafaralla, 2008; Reef
et al., 2010; Andreote et al., 2012).
Substrate communities hold abundant amounts of benthic organisms that are
classified based on their substrate location and size. Benthos are either epibenthic
or infaunal invertebrates (Dittmann, 2001; Kathiresan & Rajendran, 2005) that
occur at the soil surface or at the surface of bottom entities, and within the
substrate, respectively (Encyclopædia Britannica, Inc. 2015). Benthos are also
divided into meiofauna, which are less than .5 to 1 mm in size and macrofauna,
which are greater than 1mm body size (Dittmann, 2001).
Benthic macroinvertebrate community is predominated by epibenthic
invertebrates such as gastropods, crustaceans (Dittmann, 2001) particularly
mangrove crabs, bivalves, and other arthropods. This community is significant
as it is involved in evaluating water quality and pollution levels, functioning as
a bioindicator in rivers and mangrove ecosystems while in other areas it depicts
the condition of water currents. The ability of macroinvertebrates to assess health
of water can be attributed to its several features, which include biodiversity that
allows broader span of reactions to environmental changes, reduced movement,
sensitivity to water and habitat fluctuations, prolonged life cycles, and to their ease
of access (Superales & Zafaralla, 2008). On the other hand, meiofauna is present
in fewer representatives in mangrove ecosystems as compared to epibenthos and
they serve as food to macrofauna like shrimps and juvenile fishes (Dittmann,
2001).
Other associated communities of mangrove ecosystems are involved in
nutrient cycling and decomposition process. Microbial community is responsible
for nutrient recycling and preservation in the soil making it available for plants’
continued growth and development in an unfavorable environment, as well as for
animal utilization (Holguin, Vazquez & Bashan, 2001; Rajendran & Kathiresan,
2007). Microbes convert plant nutrients into materials that can be used by plants
like nitrogen and phosphorus through the process of decomposition that can
be completed in at least 2 months to more than 6 months in exchange of food
that they get from the roots in a form of exudates (Holguin et al., 2001). The
decomposition process is comprised of three periods, namely, the leaching phase,
followed by the oxidation phase, and lastly the disintegration of physical and
biological components of the plant (Kathiresan et al., 2011). The end products
of decayed leaves also include proteins, recalcitrant humic substance, and energy
from microbial biomass. The decomposition of A. marina leaves and Rhizophora
spp. trunks improved nitrogen concentrations by 0.5% and 500%, respectively

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while phosphorus concentration from degraded red mangrove leaves increased

by 0.09% (Holguin et al., 2001). The presence of microbial community is
predisposed to mangrove properties like soil characteristic particularly having
low oxygen level (Reef et al., 2010), nutrient cycling, availability of organic and
inorganic compounds, as well as to biogeographical, and anthropogenic factors
(Andreote et al., 2012), while climatic factors can escalate the seasonality of
bacterial extent (Lee, 1999).
Nitrogen fixation process that occur in different areas such as in decayed
leaves, pneumatophores, tree bark, in rhizosphere soil, and in the surface and
within sediments are also performed by various bacteria (Holguin et al., 2001).
This process is considered to be the foremost mechanism to make nitrogen, a vital
nutrient limiting factor, available for plant uptake (Potts, 1979) by converting
gaseous nitrogen into nitrate and ammonium forms that can be utilized by
plants (Reef et al., 2010; Giblin et al., 2013). Ammonium is generated through
denitrification process, from which nitrite and nitrate are converted to ammonium
(Reef et al., 2010), however, in Goa, India, this process is only accounted for 0 to
72% of nitrate reduction (Fernandes, Bonin, Michotey, Garcia, & LokaBharathi
et al., 2012). The dissimilatory nitrate reduction to ammonium (DNRA) is
another nitrogen pathway, which revealed a higher reduction rate of nitrate in
some areas, reaching up to 99% of nitrate reduction, indicating an effective
recycling and conservation of ammonium to outweigh nitrogen limitation as
compared to denitrification (Fernandes et al., 2012; Giblin et al., 2013), though
it is still lower from the ammonium that can be yielded in decomposition process.
The highest rate of DNRA takes place in zones with a rich source of organic
matter such as those closest to the swamp surface. The process starts with the
reduction of nitrate to nitrite in the presence of periplasmic nitrate reductase
complex (NapAB), followed by the reduction of nitrite to ammonium with the
action of pentaheme cytochrome C nitrite reductase (NrfA) (Giblin et al., 2013).
More so, nitrogen fixation is dependent on the presence of light (Potts, 1979),
temperature, and occur periodically, displaying higher rates during summer
periods and lower during winter season (Kathiresan & Bingham, 2001). In the
absence of light, reductants provide alternative means (Potts, 1979). Microbial
community also prefers to perform nitrogen fixation in anoxic soil coupled with
high organic matter in mangrove soils (Reef et al., 2010). Nitrification is the
reverse process of denitrification, which transforms ammonium to nitrate. Unlike
in denitrification and DNRA pathways, nitrification occurs at the surface layer
of the soil, which is aerobic, yet still mangroves utilize ammonium as the main
source of nitrogen (Reef et al., 2010).
Likewise, fungi community participate in the formation of detritus material

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with the bacteria (Holguin et al., 2001) promoting more phosphorus availability
to mangrove plants (Reef et al., 2010). Both bacteria and fungi comprise 91% of
the whole microbial biomass in mangrove sediments (Holguin et al., 2001), but
fungi were found to generate more decomposing enzymes than bacteria that can
break down cellulose, starch, lipid, protein, and lignin (Kathiresan et al., 2011).
Besides fungi and bacteria, 2% protozoa and 7% microalgae are involved in the
decomposition process (Holguin et al., 2001).
Microalgae (phytoplankton and benthic) and macroalgae communities add
on to the net production of mangrove ecosystems. Phytoplankton provides
additional 20 to 50% to the total net production, whereas in Northeastern
Australia algal communities and benthic cyanobacteria collectively donate only
6% to the production of mangrove habitat. Hong Kong had 4% more production
as compared to Australia (Kathiresan and Bingham, 2001). However, for benthos
community, microalgae serve as the principal source of carbon with carbon stable
isotope proportion of –17.3±1.7‰ (Bouillon et al., 2002).
Moreover, phytoplankton productivity is higher in lagoons or the borders of
mangrove than in estuarine locations. In South India, phytoplankton showed
higher production yields in mangrove areas than in nearby waters as influenced
by the refractive materials from mangroves that encourage plankton growth.
The lower productivity of microalgal communities can be due to salinity fluxes,
low light penetration, intense temperature during summer, and due to the
tannins discharged by roots and by the decomposition process of leaves and
woods (Kathiresan & Bingham, 2001). In Vietnam, algal species observed
belong in Cyanophyta, Chlorophyta, Phaeophyta, and Rhodophyta that grow in
various types of mangrove substrates (sandy, muddy, rocky), or they are affixed
to the roots and trunks or to the seagrasses, which are also associated flora of
mangrove ecosystems (Hong & San, 1993). Other associated floral communities
include trees, epiphytes, salt marsh grasses, ferns, mosses, and other plants that
can withstand the physicochemical characteristics of mangrove environment
particularly salinity (Kathiresan & Rajendran, 2005).
Furthermore, communities in the water column include the fishes and
zooplankton (Kathiresan & Bingham, 2001). Several studies have revealed the
massive use of fishes on mangrove ecosystems for food, protection (Honda
et al., 2013; El-Regal & Ibrahim, 2014), and growth (Bouchereau, Chaves &
Monti, 2008; Jones et al., 2010). Fishes are present in mangrove habitats as either
sedentary organisms like the Apogon conklini and Bairdiella ronchus, or migrants
like Atherinomorus stipes and Sphyræna barracuda that frequently and occasionally
move around, respectively (Bouchereau et al., 2008). They are also observed to
occur at peak abundance and biomass in the furthermost inland microhabitats

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with the larger fishes of more than 10 cm in length occurring in areas with
pneumatophores than in areas with nearby prop roots. This dispersal pattern
can be attributed to the presence of more food and lesser structural barriers from
roots in spaces with pneumatophores. On the other hand, large carnivorous fishes
(more than 10 cm in length) are found in greater density in the seaward zones
(Rönnbäck et al., 1999). Fish assemblies in mangrove habitats are dictated by the
properties of water in terms of temperature, pH, and level of dissolved oxygen,
while salinity is found to be a weak factor (Bouchereau et al., 2008).
Zooplankton communities are important components of a food web,
serving as food to many juvenile fishes due to their small size, accessibility, and
abundant supply (Krumme & Liang, 2004). They are also an essential factor
in governing the distribution of adult epifauna. Zooplankton can enormously
occur in large amounts in mangrove ecosystems, attaining up to 105 species
per cubic meter with a biomass equal to 623 mg per cubic meter, though there
are other mangrove ecosystems with lower abundance like in the west coast of
India (Kathiresan & Bingham, 2001). The difference of its extent is linked to
tidal variations and cycles. In Northern Brazil, the number of zooplankton that
is predominated by Copepods occurs higher in spring tide than in neap tide
(Krumme & Liang, 2004). Diverse forms of zooplankton are categorized into
three according to size. Microplankton is composed of organisms with a size
range of 20 to 199 micrometers like the rotifers and ciliates. Organisms with a
body size of 200 micrometers to 2 millimeters are categorized as mesoplankton.
An example of this is the copepod. Lastly, those that grow larger than 2 mm
are members of the macrozooplankton like the jellyfishes. This group feed on
the smaller mesoplankton. The planktonic larvae of many benthos are termed
as meroplankton such as the larvae of decapods and the zoeae (Kathiresan &
Bingham, 2001).

Flora
1. Plant Groups
Mangrove forests are normally abundant in associated flora (Figure 3), which
are more distributed in humid and wet localities than in arid areas (Kathiresan &
Rajendran, 2005). Diverse forms of associated flora perform significant roles and
functions in mangrove ecosystems (Table 1).
Mangrove plants are grouped into true mangroves and mangrove associates
depending on their zone of occurrence. True mangroves are plants that cultivate
only in zones with saline or brackish water, while mangrove associates are plants
that grow in coastline zone or mainland flora, although they can often be
observed growing together with the back mangroves (Hong & San, 1993). In

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Southeast Asia, the associated mangrove plants include 22 species of legumes,

17 species of sedges, 11 species of orchids, ten species of milk weeds (climbers
and epiphytes), 10 species of ferns, nine species of true grasses, seven species of
palms, seven species of coffee plants, six species of Terminalia family, six species
of spurges, and six species of mistletoe family (Giesen et al., 2007). Vietnam is
one of those countries in Southeast Asia with records on associated mangrove
plants like palms, grasses, legumes, and flowering plants (Hong & San, 1993). In
the Philippines, 14 species were documented in Catanduanes Island, Luzon from
10 plant families (Masagca, 2008), and five species of seagrasses were found in
Malhiao, Badian, Cebu (Valenzuela et al., 2013). Seagrasses were also observed
associated in mangrove areas in Gazi Bay, Kenya (Kathiresan & Bingham, 2001).
Grasses and about 100 epiphytes were noted in Yucatan Peninsula (Kathiresan
& Bingham, 2001). In Sundarban mangroves (between Bangladesh and India),
approximately 350 species of vascular plants were observed (Gopal & Chauhan,
2006).

2. Associated Microbiota, Fungi, and Algae

a. Bacteria
Bacterial groups are found in different parts of the mangroves dwelling on
surfaces as epiphytes, on leaves like the Flavobacterium, on the stem where
plentiful Vibrio spp. are situated, on pneumatophores like the cyanobacteria,
which also grow in any immersed surfaces (Kathiresan & Bingham, 2001),
and in the rhizosphere that was observed to have an impact on the activity of
microorganisms. Microbial population that grow in the rhizosphere can settle
in the roots of a mangrove sapling, and live on for a prolonged time after the
saplings are relocated (Gomes et al., 2010).
Bacteria are involved in controlling the availability of nutrients in an
immensely nutrient deficient mangrove soil (Reef et al., 2010). Bacterial mass in
mangrove sediments can reach an extent of 91%, making them essential in the
flow of nutrients in mangrove soils (Vazquez Holguin, Puente, Lopez-Cortes, &
Bashan, 2000). Like, for example, iron, phosphorus, and sulfur components of
mangrove soil are manipulated by the sulfate-reducing bacteria (Kathiresan &
Bingham, 2001) in an anoxic substrate. These bacteria become the prevailing
microbial component of mangrove soil with a redox potential of -75 to -150 mV,
performing an alternate respiratory pathway thru sulfate reduction (Reef et al.,
2010). Redox potential is a measure of electron availability in a system, which is
essential since electrons play a fundamental role in maintaining the stability of
soil nutrients (DeLaune & Reddy, 2005). The sulfate-reducing bacteria are also

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significant in processing pollutants like hydrocarbons (Gomes et al., 2010), while

some bacteria are found to treat industrial wastes, and others weaken the growth
rates of Staphylococcus and Pseudomonas species affecting the decomposition of
organic substances (Kathiresan & Bingham, 2001).
Inorganic nutrients in the bodies of water are made available to mangroves
using acid production in bacteria. Organic acids are presumed to solubilize
inorganic phosphate into forms that can be utilized by mangrove plants like the
orthophosphate. Such solubilizing bacteria dwell in the water, sediments, and in
the roots of associated seagrasses. Examples of such sulfate-solubilizing bacteria
are the Bacillus spp. and Vibrio spp. (Vazquez et al., 2000). Unbound phosphorus
levels is heightened by a sulphate-reducing bacteria through the reduction of iron
into another form that decreases phosphorus affinity to it; as a result, phosphorus
becomes available for plant uptake (Reef et al., 2010). Phosphorus is essential
to mangrove plants as it is needed for growth (Naidoo, 2006; Lovelock, Ball,
Martin & Feller, 2009). In a soil that is deficient with oxygen, phosphorus binds
with iron that leads to a decrease of phosphorus mobility to the plant system;
hence the activity of sulphate-reducing bacteria is very important. Phosphate
– solubilizing bacteria, likewise, contribute to phosphorus uptake of mangrove
plants in the form of phosphate. However, it is only limited to aerobic sections
(Reef et al., 2010). High levels of phosphate-solubilizing bacteria in mangrove
soils corresponded w/ the occurrence of roots forming a symbiotic relationship
(Holguin et al., 2001).
Heterotrophic and chemoolithoautotrophic microbes (Giblin et al., 2013),
and denitrifying bacteria generate ammonium from nitrate as the dominant form
of nitrogen in mangrove soils that can be absorbed by plants. Additional nitrogen
availability is produced predominantly by nitrogen-fixing cyanobacteria, which
are found associated with roots and decomposing leaves of mangroves (Reef et
al., 2010). Usually, cyanobacteria are more abundant adhering on mangrove tree
parts than on fresh leaf litter, which can be attributed to higher tannins released in
leaf litter (Kathiresan & Bingham, 2001). In a Brazilian mangrove, 1.2 to 3.5%
of cyanobacteria was observed from sediments, while others that participate both
in methane and nitrogen cycles are from Rhodobacteraceae, Planctomycetaceae,
and Burkholderiaceae (Andreote et al., 2012). Photosynthetic bacteria include
members of purple sulfur bacteria, purple non-sulfur bacteria, and green sulfur
bacteria (Kathiresan & Bingham, 2001).
The metagenomic study of Andreote et al. (2012) in four mangrove
sites uncovered eight different groups under Domain Bacteria, which
include proteobacteria, firmicutes, actinobacteria, bacteriodetes, chloroflexi,
plantomycetes, cyanobacteria, acidobacteria; and some from Domain Archea

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(1.2%) that are mainly from methanogenic Euryarchaeota. Among the different
bacteria observed, proteobacteria is the most numerous group observed
constituting 47.1 to 56.3% with representatives that are typically found in
classes Gammaproteobacteria, Deltaproteobacteria, Alphaproteobacteria,
Betaproteobacteria, and Epsilonproteobacteria (Andreote et al., 2012).
Proteobacterial associations of Desulfobacterales, Chromatiales, and
Campylobacterales are involved in processing nutrients and treating contaminants
of organic matter in anoxic environment, in oxidizing hydrogen sulfide to sulfur,
and in denitrification and other sulfide-oxidation processes, respectively (Gomes
et al., 2010). Azotobacters were found in decomposing senescent leaves and
decomposing activity is highest within 30 to 50 days. Such azotobacters were
classified as Vibrio, Flavobacterium, Pseudomonas, Acinetobacter, Corynebacterium
(Rajendran & Kathiresan, 2007).
Moreover, some bacteria live symbiotically with other organisms existing as
pathogens or parasites. Examples are the rod and sulfur-oxidizing bacteria that
take place as endosymbionts within the hindguts of detritivores and within
Lucinacea bivalves, respectively while other sulfur-oxidizing bacteria occur as
obligate ectosymbionts on ciliates. Bdellovibrios and Bacillus thuringiensis are
examples of pathogenic bacteria that affect Vibrio spp., and larvae of mosquitoes
like in Anopheles maculatus and Aedes aegypti, correspondingly. On the other
hand, actinomycetes can display antifungal activity (Kathiresan & Bingham,
2001).

b. Fungi
In the same way, fungi are essential in nutrient cycling (Kathiresan &
Bingham, 2001), as well as in preconditioning of mangrove wood, and in the
degradation of mangrove leaves (Alias & Jones, 2000), which in effect bring
protein and lipid-filled detritus material to mangrove substrate (Alias, Zainuddin
& Jones, 2010). Factors that influence dispersal and occurrence of marine fungi
include environmental vicinities, direction of water currents (Vittal & Sarma,
2006), temperature, salinity, pH, breadth of tide, presence and absence of bark
in mangrove wood, specificity of host, competition with other organisms on
nutrient sources (Alias et al., 2010), substratum richness and preference, and
vertical dispersal (Vittal & Sarma, 2006; Alias et al., 2010). Those that inhabit
the wood are influenced by the chemical component of wood, occurrence or
lack of wood, physico-chemical factors, and its various substrata (Alias & Jones,
2000). Fungi demonstrated an inclination on various substrata of an individual
mangrove tree. Such substrata are the decomposing wood, saplings, and aerial
roots. Verruculinata enalia is hugely recurrent on mangroves in the Bay of Bengal

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region, India, existing in various plant substrata, with greater occurrence on wood
and least on pneumatophores (Vittal & Sarma, 2006). Fungi were also found
to be more numerous during the early phase of decomposition when tannin is
in its higher level, which can be attributed to the presence of tannase enzyme
responsible for disintegrating tannin like in the case of Aspergilli (Rajendran &
Kathiresan, 2007). Other enzymes present in fungi that degrade mangroves are
the pectinase, protease, and amylase (Holguin et al., 2001).
Documented species of marine fungi in mangrove areas include ascomycetes,
deuteromycetes, and basidiomycetes, with the ascomycetes as the dominant
species, while basidiomycetes as the minor group (Alias & Jones, 2000;
Rajendran & Kathiresan, 2007). Ascomycetes with fruit bodies that are fully or
half submerged in the water exist in flooded dwellings (Vittal & Sarma, 2006).
One hundred thirty-nine species of marine fungi in mangroves were identified in
Malaysia (Alias et al., 2010). In Kuala Selangor, Malaysia one hundred percent
of marine fungi occupied the mangrove woods from 6 weeks to 96 weeks (Alias
& Jones, 2000). In Bay of Bengal region, India, one hundred thirty-one fungi
species under 77 genera were recorded that displayed vertical zonation, taking
place mostly in intertidal zones (Vittal & Sarma, 2006). Mitosporic fungi are also
found in mangroves (Alias a& Jones, 2000).
Arbuscular mycorrhizal fungi display symbiotic relationship with mangrove
roots, which aid in the mobilization of minerals like phosphorus, but such fungi
occur only in low salinity zones of mangrove stands tolerating up to 20 PSU of
salinity, and around the roots due to the presence of oxygen. In higher salinity
zones, mangroves become more vulnerable to the toxicity of metals (Reef et al.,
2010).

c. Fungus-like Protists
Other associated floras that participate in the decomposition process
of mangroves include the fungus-like protists such as thraustochytrids,
chytridiomycetes, and marine oomycetes. Thraustochytrids and chytridiomycetes
were both observed in the mangroves of Costa Rica and Goa, India, while
oomycetes particularly Halophytophthora were observed in Singapore, which
usually competes with the true fungi (Kathiresan & Bingham, 2001).

d. Microalgae
Microalgae contribute to less than 50% of the net production of mangrove
ecosystems making them essential food items to higher trophic organisms due
to their high nutritional content comparable to detritus materials (Kathiresan
& Bingham, 2001). In Pichavaram mangroves of Southern India, 94 species of

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phytoplankton were recognized with diatoms as the main microalgae present

(Rajkumar, Perumal, Prabu, Perumal & Rajasekar, 2009). According to
Kathiresan and Bingham (2001), microalgae productivity is limited in mangrove
areas with low availability of light and with high temperature during summer
months. This is in contrast with the study of phytoplankton in Pichavaram from
which phytoplankton is observed at its peak during summer months, concurring
with the constancy of water features (Rajkumar et al., 2009). On the other hand,
phytoplankton in Oman did not display seasonal variation. The 25 groups of
phytoplankton identified in Oman was primed by Peridinium quinquecorne Abe,
1927, reaching up to more than 90% in its occurrence (Al-Hashmi, Al-Azri,
Claereboudt, Piontkovski & Amin, 2013). In Visakhapatnam, East coast of India,
the dispersal of 55 phytoplankton genera recognized was probably influenced by
light requirement and hydrographical circumstances particularly temperature,
salinity, and nutrient. This phytoplankton community includes Chlorophyceae,
Bacillariophyceae, Cyanophyceae, and Euglenophyceae with eight, 17, seven,
and two species, respectively (Rao, Kaparapu & Rao, 2015).

e. Macroalgae
Together with the microalgae, macroalgae contribution less than 50% in
the net production of mangrove ecosystems (Kathiresan & Bingham, 2001).
The pattern of distribution for macroalgae is influenced by a number of factors
including salinity, temperature, aridity, tidal inundation, wave activity, rate of
wetting occurrence, and amount of light (Kathiresan & Bingham, 2001). In the
mangrove area of Sandspit, Karachi, Ulvales (Chlorophyta) species were observed
floating and settling on mud with the tidal cycle. Its abundance in growth was
influenced by low temperature and low intensity of light (Saifullah & Rasool,
1998). In Papua New Guinea, 25 species were documented with Rhodophyta
as the dominant group (15 species) and Chlorophyta and Phaeophyta as the
least groups with five species recognized from each (King, 1990). In Vietnam,
seven species of algae in mangrove habitats were documented in Nha Tarang
coast, and 68 species of thallophytic forms were recorded in Khanh Hoa province
belonging from four algal groups. The dominant group is the Cyanophyta (17
species), followed by Rhodophyta (19 species), then Chlorophyta (11 species),
and lastly Phaeophyta (three species). Rhodophyta is largely found as epiphytes
on mangrove roots, pneumatophores, and trunks, and on seagrasses, while the
Cyanophyta, Chlorophyta, and Phaeophyta dwell in various substrates (Hong
& San, 1993). Seaweeds (22 spp.) and seagrasses (three spp.) were also noted
in Pichavaram, Southeast India (Kathiresan, 2000). In the Philippines, three
macroalgae species were recognized (Valenzuela et al., 2013).

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Macroalgae do not only serve as food to several invertebrates and fishes

but also as habitat (Kathiresan & Bingham, 2001). In addition, Catanella
nipae (Rhodophyta) was found sensitive to copper concentrations, reducing its
productivity rate and accelerating its respiration rate; hence such species can be
utilized in the assessment of metal contamination of water (Melville & Pulkownik,
2006). Mangrove-associated microbiota, therefore, perform significant functions
to maintain mangrove stability, growth, and survival of species.

Figure 3. Mangroves and associated flora and microbiota

Table 1. Mangrove associated flora and their role in mangrove ecosystem.

Large Group Subgroups Functions References
Bacteria Denitrifying bacteria Nitrogen fixer: produce Reef et al., 2010
ammonium from nitrate
Cyanobacteria Participate in methane Andreote et al., 2012
and nitrogen cycles

Proteobacteria Nutrient processor; treats Gomes et al., 2010

contaminants

Azotobacters Decomposer Rajendran & Kathiresan,

2007
Rod and sulfur-oxidizing Endosymbionts Rajendran & Kathiresan,
bacteria 2007

Bdellovibrios and Pathogens Rajendran & Kathiresan,

Bacillus thuringensis 2007

Sulfate-solubilizing Contribute to phosphorus Reef et al., 2010; Gomes et

bacteria uptake; Process al., 2010
pollutants

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Fungus-like Thraustochytrids, Decomposers Kathiresan & Bingham,

protists Chytridiomycetes, and 2001
marine Oomycetes

Microalgae Phytoplankton Primary producers Kathiresan & Bingham,

2001; Bouillon et al., 2002
Macroalgae Rhodophyta, Primary producers as Kathiresan & Bingham,
Chlorophyta, and habitat to invertebrates 2001; King, 1990; Hong &
Phaeophyta and fishes for San, 1993
bioindicator of water
quality Kathiresan & Bingham,
2001

Melville & Pulkownik,

2006

Vascular Seagrass; Saltmarsh Nutrient source /primary de Boer 2000

Plants (grasses), palms, producer
legumes, epiphytes,
shrubs, vines, and trees

Fauna
Mangroves have already been proven by several studies to provide food and
habitat to diverse groups of animals in different trophic levels (Figure 4) (Hong
& San 1993; Giesen et al., 2007; Valenzuela, Bacalso, Gano, Pilones & Picardal,
2013) that either live temporarily or permanently in mangrove ecosystems
(Rönnbäck et al., 1999).

1. Zooplankton
Zooplankton species are one of the food items in mangrove ecosystems. These
species are small, abundant in supply, and with the ease of access especially to
numerous juvenile fishes. Examples of such species found in Furo do Meio of
Northern Brazil are the Copepods, appendicularia, chaetognaths, ostracods,
and the larvae of benthos and fishes. Among these, Copepods are the most
abundant species occurring up to 85% in the community. Factors shaping
zooplankton community include spring tide/neap tide cycles and high water/
low water variations (Krumme & Liang, 2004). Copepods and coelenterates
composed 95% of the macrozooplankton in Pichavaram (India) while tintinnids
(Tintinopsis spp.) compose 90% of the microzooplankton in the same area. Other
microzooplankton recorded were rotifers, and larvae of Copepods (nauplius) and
mollusks (veliger) (Kathiresan, 2000).

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2. Sponges
Sponges found in mangroves feed on the organic matter as their key source of
carbon. Other sponge species not associated with mangroves such as reef sponges
do not feed on mangrove-derived carbons since it will cause them necrosis
(Hunting et al., 2013). Other species form a mutualistic association with the
prop roots of R. mangle (Lee, 1999). In Badian Cebu Island, four species of
sponges were documented (Valenzuela et al., 2013).

3. Crustaceans
The dispersal and diversity of benthic invertebrates in mangrove habitats in
Pondicherry is influenced by environmental factors namely salinity, dissolved
oxygen levels, the composition of organic matter, and concentration of sulfide
nutrient. There were 76 taxa of invertebrates recorded in the area (Kumar & Khan,
2013). The most frequently occurring benthos in mangroves are the epibenthic
species like gastropods and crustaceans (Dittmann, 2001). Several literatures have
already indicated the existence of such species in different mangrove habitats in
various countries including the Philippines (Valenzuela et al., 2013), Sundarban
(Gopal & Chauhan, 2006), India (Kathiresan, 2000; Kumar & Khan, 2013),
Indonesia (Sukardjo, 2004), Vietnam (Hong & San, 1993), and New Zealand
(Stokes, Healy & Mason, 2009). Examples of crustaceans observed are the
isopods, amphipods (Kathiresan, 2000; Thilagavathi et al., 2013) tanaids,
cirripedes, crabs, hermit crabs, shrimps (Kathiresan, 2000), and cumacea
(Thilagavathi et al., 2013). In the Philippines, four species of crustaceans were
documented in Cebu (Valenzuela et al., 2013) and 5 in Leyte (Picardal, Avila,
Tano & Marababol, 2011), and tree-climbing crabs were recorded in Southern
Luzon (Masagca, 2011).

4. Mollusks
Mollusks that thrive in the mangrove ecosystems are either dwelling in the
soil or are adhered on plant parts like on the stem, leaves, and roots. Due to
their varied habitats, they feed on different food types that categorize them
into 6 groups: the suspension feeders, litter feeders, plant and deposit feeders,
animal feeders (carnivorous), deposit feeders and mud raspers, and algae and
detritus feeders (Hong & San, 1993). Frequently documented mollusks in
mangroves are the bivalves (Giesen et al., 2007; Superales & Zafaralla, 2008) and
gastropods (Dittmann, 2001). In Pondicherry, the important species Cassidula
nucleus, Melampus ceylonicus, and Sphaerassiminea minuta that can be utilized
as pollution indicators were recorded (Kumar& Khan, 2013). In Malubog Bay,
Western Mindanao, mangrove mollusks comprise 56.4% and 30.76% of which

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are bivalves while the remaining 25.64 % are gastropods (Superales & Zafaralla,
2008).

5. Other Invertebrates
Other associated invertebrates in mangrove areas include cnidarians, spiders,
ants, crickets, moths (Valenzuela et al., 2013), insects, echinoderms, (Picardal et
al., 2011; Valenzuela et al., 2013), polychaetes (Hong & San, 1993; Kathiresan,
2000; Thilagavathi et al., 2013), and annelids (Superales and Zafaralla, 2008).
Nematodes comprise 50 to 70% in Pichavaram mangroves (Kathiresan, 2000).
In the Indo-Malaysian region, insects predominate the mangrove forests with
500 species recorded. Ctenophores, byrozoans, and other invertebrates are also
found in the region (Kathiresan & Rajendran, 2005).

6. Fishes
Mangroves are also home to many vertebrate species especially to fishes,
providing them nurseries for the juveniles, area for growth, food, and shelter
(Jones et al., 2010; Honda et al., 2013; El-Regal & Ibrahim, 2014). Both the
juvenile and adult stages of small gobioid fishes were observed in Southern
Thailand, which was predominated by Pandaka pygmaea (75.9%), Acentrogobius
kranjiensis (15.0%) and Acentrogobius malayanus (3.8%) (Yokoo et al., 2012).
In Pichavaram, 177 fish species were documented (Kathiresan, 2000). In the
Philippines, Belonidae, Gobiidae, and Pomacentridae species were recovered in
Cebu (Valenzuela et al., 2013), and 4 fish genera were attained in Leyte (Picardal
et al., 2011). In Florida, USA, fishers catch fishes that are found in prop roots
such as mangrove snapper, seatrout, redfish, tarpon, and snook (Lorenz, 2013).
Other fishes spend only their juvenile stage in mangrove ecosystems like several
species of coral reef fishes, and some of those are commercially important
(Jones et al., 2010; Honda et al., 2013; El-Regal & Ibrahim, 2014). Fishes
are positioned intermediary in their trophic level in both coastal and estuarine
ecosystems, predating on enormous density of benthos, and being predated upon
by shorebirds as top predators (Fujii, 2012).

7. Other Vertebrates
Other mangrove associated vertebrates include tunicates and golden crown
flying fox recorded in Leyte (Picardal et al., 2011); Egretta (migratory bird),
white-collard kingfisher, olive-backed sunbird, and mangrove skink (reptile)
observed in the mangroves of Cebu (Valenzuela et al., 2013); Gerygone warbler in
Australia (Nyári & Joseph, 2012); and snake species (aquatic, arboreal, and some
poisonous), monkeys, wild pigs, crocodiles, monitor lizards, and numerous birds

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in Vietnam (Hong & San, 1993). Birds are usually considered as the highest
trophic level in coastal and estuarine ecosystems that feed on fishes and benthic
macrofauna (Fujii, 2012).

8. Endangered Vertebrates
The Sundarban is a dwelling to a some rare and endangered vertebrates such
as Batagur baska, Pelochelys bibroni, Chelonia mydas, and Panthera tigris (Gopal &
Chauhan, 2006). Florida, USA also have protected species in mangrove forests,
which include manatee, small-toothed sawfish, goliath grouper, bottlenose
dolphin, white-crowned pigeon, reddish egret, striped mud turtle, key deer,
American crocodile, bald eagle, osprey brown pelican, and mangrove cuckoo
(Lorenz, 2013).

9. Associated Animal-like Protists (Protozoa)

The protozoa species participate in the decomposition process of mangroves
similar to bacteria and fungi (Tam, Vrijmoed & Wong, 1990; Holguin et al.,
2001). Protozoans like flagellates, amoebae, and ciliates were observed in sheets
in the prop roots of R. mangle in Florida, USA. These protozoans preyed on
bacteria and phytoplankton like diatoms (Maybruck & Rogerson, 2004).

Figure 4. Mangrove food web of associated fauna

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The Crabs Associated with Mangroves

Mangrove crabs display a vital role in the maintenance of ecosystem (Arya,
Trivedi & Vachhrajani, 2014) contributing 10% to the system function (Wolff,
Koch, & Isaac, 2000). Such crabs are known for their role in returning nutrients
to the soil by feeding on leaf litter and cutting them into small bits and pieces
(Reef et al., 2010). Nutrients are recycled by way of fecal matter excretion to
the soil. Fecal matter is comprised of more nutrients and much fewer tannins
compared to the leaves, which improves detrital matter cycle (Micheli, Gherardi
& Vannini, 1991). In can also create a basis of food chain for small invertebrates
feeding on animal excrements or it can be re-exported in the form of particulate
matter (Lee, 1998). Examples of crabs that feed on mangrove leaf litters are the
sesarmids (Sheaves & Molony, 2000), brachyurans (Nordhaus et al., 2009),
grapsid crabs (Lee, 1998; Nerot et al., 2009), and fiddler crabs (Nicholson,
2009).
Records have shown that mangrove crabs are involved in soil turnover through
their burrowing activities, carrying the organic material back to the upper areas
of the soil (Micheli et al., 1991). Bioturbation is another outcome of burrowing
that causes changes in the soil in terms of surface features, dispersal of particle size,
and level of aerification, which consequently alters phytotoxin concentrations in
the substratum (Lee, 1998). This, in return, aids in facilitating mangrove growth
and recovery (Micheli et al., 1991) that influence production (Lee, 1998). Soil
aerification also affects reproductive output as observed in Rhizophora (Smith,
Boto, Frusher, & Giddins, 1991). In an experiment conducted, soil sulfide and
ammonium concentrations accelerated when grapsid crabs were removed from
the mangrove forest. This can affect the growth, dispersal, and reproductive
patterns of mangroves (Smith et al., 1991). Mud crabs have chelipeds containing
dactylus that serves as a flexible digit involved in walking and burrowing together
with its other three pairs of appendages (Paital & Chainy, 2012). Burrows are
important to crabs as it lessens the effect of high temperatures, and functions
as a pool of water that moistens the gill chambers (Wolff et al., 2000). Ucides
cordatus (Linnaeus, 1763) is one of those mangrove crabs that retain burrows in
the sediment with length ranging from 10 to 144 cm (Araujo & Calado, 2012).
Since crabs have a large impact on ecosystem structure and function, they are
very essential in mangrove forests as keystone species (Smith et al., 1991).
Efficient nutrient cycling in mangroves is essential since its soils are very
poor in nutrient availability as they are low in oxygen, saline, acidic, and
often waterlogged. Such nutrient-poor soil is a key factor that limits mangrove
productivity (Reef et al., 2010). Hence, leaf litter processing and bioturbation
activities of mangrove crabs play a significant function in promoting nutrient

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supply in mangrove ecosystems.

Some mangrove crab species feed on mangrove propagules (Smith et al.
,1991; Cannicci et al., 2012), consuming 100% of the propagules in Australian
mangroves particularly on Avicennia (Smith et al., 1991). This activity influences
the distribution patterns of saplings in mangrove forests, lessening their
abundance in areas where they are predated by crabs (Cannicci et al., 2012) like
the grapsid crabs (Smith et al., 1991).
Crabs also exhibit the prey-predator relationship with other organisms,
which is important in maintaining the ecosystem (Arya et al., 2014). The tree-
climbing grapsid crabs observed in Southern Luzon consume algae, mollusks,
and crustaceans other than the leaves (Masagca, 2011). Tree-climbing crabs were
also recorded in East African Mangroves (Fratini, Vannini, Cannicci & Schubart,
2005).

Issues and Concerns in Mangrove Ecosystems

Mangrove ecosystems have already decreased in number since the last 50
years, diminishing to about one-third of mangrove forests worldwide (Alongi,
2002). Mangrove areas in the world are found mainly (75%) in 15 countries
and from which only 6.9% are protected. In the year 2000, mangroves occupy
a sum area of 137,760 km2, but this has declined approximately by 12.3% after
several years (Giri et al., 2011). The Philippines have lost more than half of the
mangroves with 240, 800 hectares remaining as of 2010 (Primavera et al., 2014).
The density of human population is frequently regarded as a factor clearly
associated with the continued degradation of mangrove forests. Human
activities attributed to the main causes of mangrove degradation (Figure 5)
include aquaculture, mining, overexploitation for mangrove economic services
(timber and fisheries products), and urban development (Alongi, 2002) such as
conversion to salt ponds, settlements, agriculture like rice and coconut field, and
tourist resorts (Primavera, 2005; Primavera et al., 2014). From these, aquaculture
remains the topmost basis of declining mangrove forests (Primavera et al., 2014).
In Sundarban, mangrove destruction was instigated by the conversion of mangrove
stands into paddy areas and shrimp farms, and by prevalent overexploitation of
timber, fishes, prawns, and fodder (Gopal & Chauhan, 2006). In the Philippines,
mangrove decline is correlated to the offshore fish production (Primavera, 2000)
and brackish culture ponds became the greatest threat to mangrove degradation
as well as its ecological and economic services (Primavera, 1993).
The continued growth of population especially in poor countries and the
industrialization of fishing have exceeded the sustainable production of our
waters that lead to declining fish landings. This event gave rise to aquaculture

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(Tidwell & Allan, 2001). Nowadays, aquaculture supplies one-third of the overall
fisheries production from 16% in 1991, and globally, it reaches up to a sum of
51.4 million, more than 90% of which is from Asia. However, aquaculture leaves
several environmental effects, which are the accumulation of salts both in soil
and water, introduction of invasive species, dispersal of parasites and diseases,
deficiency of bycatch in wild seed and broodstock collection, dislocation of wild
populations, coastal pollution from aquaculture wastes, negative effects on genetic
trait, negative effects of chemicals and antibiotic, and degradation of habitat.
Environmental impact with prime effects is the loss of habitat (Primavera, 2005).
In Vietnam, associated mammals in mangroves declined because of hunting and
degradation of forest (Hong & San, 1993).
More so, the growth of population brings a continual threat on the mangrove
habitats especially on the effects of freshwater influx from the upstream zones
and the exploration of oil in coastal areas as another peril to mangroves (Gopal
& Chauhan, 2006).
The continued degradation of mangrove ecosystems indicates greater
susceptibility to the impacts of climate change primarily on the sea level rise
(Ellison & Zouh, 2012; Gilman, Ellison, Duke, & Field, 2008).This event greatly
affects the seaward mangroves like Avicennia (Ellison & Zouh, 2012) with low
net sedimentation rates that limit landward migration (Ellison & Zouh, 2012;
Gilman et al., 2008). In Cameroon, Central Africa, seaward mangroves already
experienced a loss of 89% with the effects of sea level rise (Ellison & Zouh,
2012). Mangroves can trap suspended sediments carried by tidal inundation to
approximately 80%, which is comparable to about 10–12 kg of sediment m−1creek
length/spring tides. As an effect, substrate elevates at a rate of approximately 0·1
cm year−1 (Furukawa, Wolanski, & Mueller, 1997). Sedimentation, therefore,
keeps mangrove at pace with the rising sea level (Ellison & Zouh, 2012).
Climate change can result to a global mangrove loss at an optimal rate of
10 to 15% (Alongi, 2008). This further signifies higher risks to flooding, storm
surges, cyclones, and tsunamis related to loss or weakened coastal protection and
increased threat to mangrove associated communities and human settlement.

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Source: Valiela, Bowen & York (2001)

Figure 5. Human use of mangrove ecosystems leading to its degradation: (SC)

shrimp culture; (FU) forest use; (FC) fish culture; (DF) diversion of freshwater;
(LR) land reclamation; (H) herbicides; (A) agriculture

SYNTHESIS

Mangrove distribution is influenced by climatic, environmental, and

physiochemical factors, propagule properties, and biotic interactions. Mangrove
shrubs and trees are dispersed differently in mangrove ecosystems based on the
tidal level (low, mid, high) they are exposed to, on the elevation (downstream,
intermediate, and upstream), and on the type of substrate from which they grow
(Primavera et al. 2012, 2014). Factors may overlap and in order for mangrove
species to successfully establish, such influencing factors must be suitable for the
plant’s characteristics and adaptabilities. This indicates the importance of site
selection in mangrove reforestation programs. However, many of such programs
had mismatching of mangrove species and its environment that led to poor
survival rate just like what happened in Visayas, Mindanao (Melana, Melana,
& Mapalo, 2005), and in Indonesia (Brown, Fadillah, Nurdin, Soulsby, &
Ahmad, 2014). Hence, mangrove reforestation programs must ascertain that the
right mangrove species are planted in their suitable location and environmental
characteristics to enhance growth and dispersal, thereby successfully reforesting
mangrove areas.

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Mangrove ecosystems are increasingly utilized for fisheries production.

However, such fisheries products are unsustainably used leading to altered
mangrove ecosystem affecting food web and mangrove stability. This indicates the
need for sustainable use of mangrove ecosystems to maintain both its ecological
and economic services.
Mangrove ecosystem is also rich in various associated communities, which
are essential in maintaining the stability of the ecosystem. This indicates that
the associated fauna and flora including the microbiota should be included in
rehabilitation and reforestation activities of degraded mangrove ecosystems.
The microbiota is essential in the site locations of mangroves. However, this is
not observed in rehabilitation and reforestation programs. Studies conducted
on the characteristics of the different species of mangrove associated flora, the
factors that affect their growth and dispersal in mangrove ecosystems, their role
and interaction with other biotic factors that would highlight their significance
in mangrove ecosystems, and their biodiversity index is also very limited. The
presence of diverse forms of animals in different trophic levels interacting with
each other validates the importance of protection and conservation of mangrove
ecosystems worldwide. Nevertheless, similar to associated flora, studies on
mangrove associated fauna and their biodiversity index is very limited both
locally and globally. The sources used in the methodology are also limited to
open access journals only and very few studies related to mangrove associated
fauna and flora are conducted in the Philippines. Thus, mangrove rehabilitation
and reforestation programs must include its associated communities to promote
balance in mangrove ecosystems and enhance species diversity.
One of the essential fauna in mangrove ecosystems are the crabs. They serve
as key species in the ecology of mangrove ecosystems as they are crucial in
sustaining nutrient availability in a nutrient-poor environment, in maintaining
high ecosystem productivity, and in sustaining the food web that supports a
myriad of organisms in different trophic levels. This designates the importance
of sustainable use of mangrove crabs since this group of animals are increasingly
utilized by humans as food and income generating product. In the Philippines,
mud crabs are utilized as food. Studies conducted on such species are mainly
focused on its economic use and propagation. Local and global literature on its
ecological role particularly in mangrove ecosystems are very limited. Hence, its
ecological importance is often neglected.
Humans have become the main culprit in the continued degradation
of mangrove ecosystems worldwide. This can be attributed to the lack of
understanding on the characteristics of mangrove ecosystems and on the
importance of ecosystem stability, as well as the factors involved to maintain its

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stability. The characteristics of mangrove ecosystems must be well understood

by humans to achieve successful reforestation, rehabilitation, and conservation
programs. Understanding mangrove ecosystem stability promotes biodiversity
and sustainable use of its products. The regulation of anthropogenic activities
may help in preventing further degradation of mangrove ecosystems, however,
with the continued increase of human population mangrove ecosystems remain
threatened due to its food resources. Thus, the sustainable use of mangrove
ecosystems must be actively encouraged for the continuity of mangrove’s
ecological and economic functions.

CONCLUSIONS

Mangroves generally occur in the tropical and subtropical areas with more
species found in the Indo-West Pacific hemisphere. They are distributed in
different regions, with Asia holding the largest cover of mangroves, as influenced
by climatic factors (temperature and rainfall). They are distributed horizontally
and vertically in estuarine and intertidal zones as directed by abiotic and biotic
factors.
Mangrove ecosystem holds a number of organisms that are sourced as fisheries
products that are utilized both for food and export. Common products are
mud crabs, shrimps, mollusks, and fishes. Mangrove ecosystems are comprised
of diverse forms of associated communities of flora, fauna, and microbiota. Its
associated flora is composed of true mangroves and associates that are distributed
in zonation pattern. Its fauna is composed mainly of benthic macroinvertebrates
and fishes. The growth of human population is related to the unsustainable use
of fish products like mud crabs, which resulted to the decrease of fish landings
that started aquaculture. The greatest human threat to mangrove ecosystems is
aquaculture and its prime effect is the loss of habitat. The continued degradation
of mangrove ecosystems increases its vulnerability to the impacts of climate
change particularly on the rising of sea level. The seaward mangroves with low
net sedimentation rate are at higher risk for degradation from the rising of sea
level due to limited landward migration. Mangrove degradation increases the
risk of tsunamis, storm surges, cyclones, and floods due to the loss or weakened
coastal protection that can affect both mangrove associated communities and
human settlements.
Hence, the distribution of mangrove ecosystems is only limited to particular
geographical location. Its growth is dictated by both abiotic and biotic factors
particularly climatic factors. The ecosystem is used by humans for its food
resources since it is composed of diverse associated fauna. The associated flora,

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fauna, and microbiota are essential in maintaining the stability of mangrove

ecosystem. Anthropogenic activities, especially aquaculture, altered this stability.

RECOMMENDATIONS

Aquaculture is parallel to the declining mangrove ecosystems. In line with this,

mangrove protection, conservation, rehabilitation and reforestation programs
must be properly implemented. Proper matching of mangrove plants and site
location, sustainable use of mangrove ecosystems, and eco-friendly aquaculture
must be enforced.

TRANSLATIONAL RESEARCH

The outcome of this study entitled “Mangrove Ecosystems and Associated

Fauna with Special Reference to Mangrove Crabs in the Philippines: A Review”
will be translated into brochures that will be distributed to fishermen and coastal
settlers. The brochure entitled “Ang Importansiya sa Bakawan ug ang Iyang mga
Kahayopan” will improve the knowledge of fishermen and coastal settlers from
Samal Island, Davao City, and Panabo City on the sustainable use of mangrove
ecosystems.

LITERATURE CITED

Al-Hashmi, K., Al-Azri, A., Claereboudt, M. R., Piontkovski, S., & Amin,
S. M. N. (2013). Phytoplankton Community Structure of a Mangrove
Habitat in the Arid Environment of Oman: The Dominance of Peridinium
[Link] of Fisheries & Aquatic Science, 8(5). Retrieved on April
28, 2015 from [Link]

Alias, S. A., & Jones, E. B. G. (2000). Colonization of mangrove wood by marine

fungi at Kuala Selangor mangrove stand, Malaysia. Fungal Diversity, 5, 9-21.
Retrieved on March 19, 2015 from [Link]

Alias, S. A., Zainuddin, N., & Jones, E. B. (2010). Biodiversity of marine fungi
in Malaysian mangroves.  Botanica Marina,  53(6), 545-554. Retrieved on
February 14, 2015 from [Link]

95
IAMURE International Journal of Ecology and Conservation

Allan, J. D., & Castillo, M. M. (2007). Stream ecology: structure and function of

running waters. Springer Science & Business Media. Retrieved on April 27,
2015 from [Link]

Alongi, D. M. (2009).  The energetics of mangrove forests. Springer Science &

Business Media. Retrieved on March 13, 2015 from [Link]

Alongi, D. M. (2008). Mangrove forests: Resilience, protection from tsunamis,

and responses to global climate change. Estuarine, Coastal and Shelf Science,
76(1), 1–13. doi:10.1016/[Link].2007.08.024

Alongi, D. M. (2002). Present state and future of the world’s mangrove forests.
Environmental conservation, 29(03), 331-349. Retrieved on March 17, 2015
from [Link]

Andreote, F. D., Jiménez, D. J., Chaves, D., Dias, A. C. F., Luvizotto, D. M.,
Dini-Andreote, F., ... & de Melo, I. S. (2012). The microbiome of Brazilian
mangrove sediments as revealed by metagenomics. PLoS One, 7(6), e38600.
Retrieved on February 19, 2015 from [Link]

Anouti, F. A. (2014). Bioaccumulation of Heavy Metals within Mangrove

Ecosystems. J Biodivers Endanger Species 2: e113. doi: 10.4172/2332-
2543.1000 e113 Page 2 of 2 Volume 2• Issue 2• 1000e113 J Biodivers
Endanger Species ISSN: 2332-2543 JBES an open access journal distribution
in mangrove sediments along the mobile coastline of Frenc h Guiana. Mar
Chem, 98, 1-17. Retrieved on February 20, 2015 from [Link]

Araujo, M. de S. L. C., & T. C. dos S. Calado (2012). Burrows Architecture of

the Crab Ucides Ccrdatus (Linnaeus, 1763) (Crustacea, Decapoda, Ucididae)
in a Mangrove Swamp of Brazil. Tropical Oceanograpgy Online, 155–165.
Retrieved on December 29, 2014 from [Link]

Arya, S., Trivedi, J. N., & Vachhrajani, K. D. (2014) Brachyuran Crabs as a

Biomonitoring tool: A Conceptual Framework for Chemical Pollution
Assessment. Retrieved on April 1, 2015 from [Link]

Bouchereau, J. L., Chaves, P. D. T., & Monti, D. (2008). Factors structuring the
ichthyofauna assemblage in a mangrove lagoon (Guadeloupe, French West
Indies). Journal of Coastal Research, 969-982. Retrieved on March 24, 2015

96
 Volume 15 · July 2015

from [Link]

Bouillon, S., Connolly, R. M., & Lee, S. Y. (2008). Organic matter exchange
and cycling in mangrove ecosystems: recent insights from stable isotope
studies. Journal of Sea Research, 59(1), 44-58. Retrieved on February 20, 2015
from [Link]

Bouillon, S., N. Koedam, A. Raman, and F. Dehairs. 2002. Primary producers

sustaining macro-invertebrate communities in intertidal mangrove forests.
Oecologia 130:441–448.

Brander, L. M., Wagtendonk, A. J., Hussain, S. S., McVittie, A., Verburg, P. H.,
de Groot, R. S., & van der Ploeg, S. (2012). Ecosystem service values for
mangroves in Southeast Asia: A meta-analysis and value transfer application.
Ecosystem Services, 1(1), 62-69. Retrieved on March 4, 2015 from [Link]
gl/DYOOMq

Brown, B., Fadillah, R., Nurdin, Y., Soulsby, I., & Ahmad, R. (2014). CASE
STUDY: Community Based Ecological Mangrove Rehabilitation (CBEMR)
in Indonesia. From small (12-33 ha) to medium scales (400 ha) with pathways
for adoption at larger scales (> 5000 ha). SAPI EN. S. Surveys and Perspectives
Integrating Environment and Society, (7.2). Retrieved on March 24, 2015
from [Link]

Cannicci, S., Bartolini, F., Penha-Lopes, G., Fratini, S., Fusi, M., & Dahdouh-
Guebas, F. (2012). Functions of macrobenthos in mangrove forests.  VLIZ
Special Publication, 57, 1-9. Retrieved on March 28, 2015 from [Link]
gl/pU1w8k

Chen, Y., & Ye, Y. (2014). Effects of salinity and nutrient addition on mangrove
Excoecaria agallocha.  PloS one,  9(4), e93337. Retrieved on April 4, 2015
from [Link]

Clarke, P. J., & Myerscough, P. J. (1993). The intertidal distribution of the

grey mangrove (Avicennia marina) in southeastern Australia: the effects of
physical conditions, interspecific competition, and predation on propagule
establishment and survival.  Australian Journal of Ecology,  18(3), 307-315.
Retrieved on April 25, 2015 from [Link]

97
IAMURE International Journal of Ecology and Conservation

de Boer, W. F. (2000). Biomass dynamics of seagrasses and the role of mangrove

and seagrass vegetation as different nutrient sources for an intertidal
ecosystem.  Aquatic Botany,  66(3), 225-239. Retrieved on January 9, 2015
from [Link]

DeLaune, R. D., & Reddy, K. R. (2005). Redox potential. Encyclopedia of Soils in

the Environment, 3. Retrieved on January 10, 2015 from [Link]

Delgado, P., Hensel, P. F., Jiménez, J. A., & Day, J. W. (2001). The importance
of propagule establishment and physical factors in mangrove distributional
patterns in a Costa Rican estuary. Aquatic Botany, 71(3), 157-178. Retrieved
on January 9, 2015 from [Link]

Dittmann, S. (2001). Abundance and distribution of small infauna in mangroves

of Missionary Bay, North Queensland, Australia.  Revista de biología
tropical,49(2), 535-544. Retrieved on February 14, 2015 from [Link]
gl/ThdxrZ

Duke, N., Ball, M., & Ellison, J. (1998). Factors influencing biodiversity
and distributional gradients in mangroves.  Global Ecology & Biogeography
Letters,  7(1), 27-47. Retrieved on December 27, 2014 from [Link]
rPxxFg

Ellison, J. C., & Zouh, I. (2012). Vulnerability to climate change of mangroves:

assessment from cameroon, central Africa. Biology, 1(3), 617–38. doi:10.3390/
biology1030617

El-Regal, M. A. A., & Ibrahim, N. K. (2014). Role of mangroves as a nursery

ground for juvenile reef fishes in the southern Egyptian Red Sea. The Egyptian
Journal of Aquatic Research, 40(1), 71-78. Retrieved on December 20, 2014
from [Link]

Fernandes, S. O., Bonin, P. C., Michotey, V. D., Garcia, N., & LokaBharathi,
P. A. (2012). Nitrogen-limited mangrove ecosystems conserve N through
dissimilatory nitrate reduction to ammonium. Scientific reports, 2. Retrieved
on February 8, 2015 from [Link]

Fratini, S., Vannini, M., Cannicci, S., & Schubart, C. D. (2005). Tree-climbing
mangrove crabs: a case of convergent evolution.  Evolutionary Ecology

98
 Volume 15 · July 2015

Research, 7(2), 219-233. Retrieved on February 5, 2015 from [Link]

sYQbR5

Fujii, T. (2012). Climate change, sea-level rise and implications for coastal and
estuarine shoreline management with particular reference to the ecology
of intertidal benthic macrofauna in NW Europe.  Biology,  1(3), 597-616.
Retrieved on April 28, 2015 from [Link]

Furukawa, K., Wolanski, E., & Mueller, H. (1997). Currents and Sediment
Transport in Mangrove Forests. Estuarine, Coastal and Shelf Science, 44(3),
301–310. doi:10.1006/ecss.1996.0120

Giblin, A., C. Tobias, B. Song, N. Weston, G. Banta, and V. Rivera-Montoy.

2013. The Importance of Dissimilatory Nitrate Reduction to Ammonium
(DNRA) in the Nitrogen Cycle of Coastal Ecosystems. Oceanography 26(3).

Giesen, W., Wulffraat, S., Zieren, M., & Scholten, L. (2007). Mangrove guidebook
for Southeast Asia. FAO Regional Office for Asia and the Pacific. Retrieved on
January 15, 2015 from [Link]

Gilman, E. L., Ellison, J., Duke, N. C., & Field, C. (2008). Threats to mangroves
from climate change and adaptation options: A review. Aquatic Botany, 89(2),
237–250. doi:10.1016/[Link].2007.12.009

Giri, C., Ochieng, E., Tieszen, L. L., Zhu, Z., Singh, A., Loveland, T., ... &
Duke, N. (2011). Status and distribution of mangrove forests of the world
using earth observation satellite data. Global Ecology and Biogeography, 20(1),
154-159. Retrieved on January 15, 2015 from [Link]

Gomes, N. C., Cleary, D. F., Pinto, F. N., Egas, C., Almeida, A., Cunha, A., ...
& Smalla, K. (2010). Taking root: enduring effect of rhizosphere bacterial
colonization in mangroves. PLoS One, 5(11), e14065. Retrieved on February
8, 2015 from [Link]

Gopal, B., & Chauhan, M. (2006). Biodiversity and its conservation in the
Sundarban Mangrove Ecosystem. Aquatic Sciences, 68(3), 338-354. Retrieved
on March 17, 2015 from [Link]

99
IAMURE International Journal of Ecology and Conservation

Hastuti, E. D., Anggoro, S., & Pribadi, R. (2013). The Effects of Environmental
Factors on the Dynamic Growth Pattern of Mangrove Avicennia
marina. JOURNAL OF COASTAL DEVELOPMENT,16(1), 57-61. Retrieved
on March 28, 2015 from [Link]

Holguin, G., Vazquez, P., & Bashan, Y. (2001). The role of sediment
microorganisms in the productivity, conservation, and rehabilitation of
mangrove ecosystems: an overview. Biology and fertility of soils, 33(4), 265-
278. Retrieved on February 20, 2015 from [Link]

Honda, K., Nakamura, Y., Nakaoka, M., Uy, W. H., & Fortes, M. D. (2013).
Habitat use by fishes in coral reefs, seagrass beds and mangrove habitats in
the Philippines. Plos one, 8(8), e65735. Retrieved on January 7, 2015 from
[Link]

Hong, P. N., & San, H. T. (1993). Mangroves of Vietnam-IUCN.  The World

Conservation Union, Bangkok, Thailand. Retrieved on January 17, 2015 from
[Link]

Hunting, E. R., Ubels, S. M., Kraak, M. H., & van der Geest, H. G. (2013).
Root-derived organic matter confines sponge community composition in
mangrove ecosystems. Ecological Processes, 2(1), 1-4. Retrieved on March 24,
2015 from [Link]

Jones, D. L., Walter, J. F., Brooks, E. N., & Serafy, J. E. (2010). Connectivity
through ontogeny: fish population linkages among mangrove and coral reef
habitats. Marine Ecology Progress Series, 401, 245-258. Retrieved on March
12, 2015 from [Link]

Juario, J., & Ontoy, D. (2005). Reassessment of the Mangrove Forest in Ormoc
Bay, Philippines. UPV Journal of Natural Sciences, 10, 181–196. Retrieved
April 28, 2015 from [Link]
the_Mangrove_Forest_in_Ormoc_Bay_Philippines

Kathiresan, K. (2000). A review of studies on Pichavaram mangrove, southeast

India. Hydrobiologia, 430(1-3), 185-205. Retrieved on March 13, 2015 from
[Link]

100
 Volume 15 · July 2015

Kathiresan, K., & Bingham, B. L. (2001). Biology of mangroves and mangrove

ecosystems. Advances in marine biology, 40, 81-251. Retrieved on March 13,
2015 from [Link]

Kathiresan, K., & Rajendran, N. (2005). Mangrove ecosystems of the Indian

Ocean region. Indian Journal of marine sciences, 34(1), 104-113. Retrieved on
March 13, 2015 from [Link]

Kathiresan, K., Saravanakumar, K., Anburaj, R., Gomathi, V., Abirami, G., Sahu,
S. K., & Anandhan, S. (2011). Microbial enzyme activity in decomposing
leaves of mangroves. Int J Adv Biotechnol Res, 2(3), 382-389. Retrieved on
March 13, 2015 from [Link]

King, R. J. (1990). Macroalgae associated with the mangrove vegetation of Papua

New Guinea. Botanica Marina, 33(1), 55-62. Retrieved on March 14, 2015
from [Link]

Kosuge, T. (2001). Brief assessment of stock of mud crabs Scylla spp. in Matang
Mangrove Forest, Malaysia and proposal for resources [Link]
Agricultural Research Quarterly, 35(2), 145-148. Retrieved on February 24,
2015 from [Link]

Krumme, U., & Liang, T. H. (2004). Tidal-induced changes in a copepod-

dominated zooplankton community in a macrotidal mangrove channel in
northern Brazil.  Zoological Studies,  43(2), 404-414. Retrieved on February
15, 2015 from [Link]

Kumar, P. S., & Khan, A. B. (2013). The distribution and diversity of benthic
macroinvertebrate fauna in Pondicherry mangroves, India.  Aquatic
biosystems,9(1), 15. Retrieved on February 15, 2015 from [Link]
Z2SkGD

Lee, S. Y. (1995). Mangrove outwelling: a review. Hydrobiologia, 295(1-3), 203-

212. Retrieved on January 10, 2015 from [Link]

Lee, S. Y. 1998. Ecological role of grapsid crabs in mangrove ecosystems: a review.

Marine and Freshwater Research 49(4):335–343.

101
IAMURE International Journal of Ecology and Conservation

Lee, S. Y. 1999. Tropical mangrove ecology: Physical and biotic factors influencing
ecosystem structure and function. Austral Ecology 24(4):355–366.

Long, J. B., & Giri, C. (2011). Mapping the Philippines’ mangrove forests using
Landsat imagery. Sensors, 11(3), 2972-2981. Retrieved on March 15, 2015
from [Link]

Lorenz, J. J. (2013). Benthic Habitat: Mangroves. Retrieved on February 10,

2015 from [Link]

Lovelock, C. E., Ball, M. C., Martin, K. C., & Feller, I. C. (2009). Nutrient
enrichment increases mortality of mangroves.  PLoS One,  4(5), e5600.
Retrieved on December 27, 2014 from [Link]

Lunar, B. C., & Laguardia, M. A. (2013, January 17). Comparative Study of

Diversity of Mangroves in Two Conservation Sites of Calatagan, Batangas,
Philippines. IAMURE International Journal of Marine Ecology. Retrieved
April 28, 2015 from [Link]
view/337

Masagca, J. T. (2008). Occurrence and Distributional Range of Mangrove

Vascular Flora of Catanduanes Island, Luzon, Philippines. Retrieved on
March 29, 2015 from [Link]

Masagca, J. T. (2011). Occurrence of arboreal-climbing grapsids and other

brachyurans in two mangrove areas of Southern Luzon, Philippines.
BIOTROPIA,  18(2). Retrieved on March 29, 2015 from [Link]
d9zHwW

Maybruck, B. T., & Rogerson, A. (2004). Protozoan Epibionts on the Prop Roots
of the Red Mangrove Tree, Rhizaphora mangle. Retrieved on March 20, 2015
from [Link]

Melana, D. M., Melana, E. E., & Mapalo, A. M. (2005). Mangroves management

and development in the Philippines. Retrieved on March 29, 2015 from
[Link]

Melville, F., & Pulkownik, A. (2006). Mangrove macroalgae as potential estuarine

test species in phytotoxicity tests using physiological endpoints. Australasian

102
 Volume 15 · July 2015

Journal of Ecotoxicology, 12(1), 21-28. Retrieved on February 18, 2015 from

[Link]

Micheli, F., Gherardi, F., & Vannini, M. (1991). Feeding and burrowing ecology
of two East African mangrove crabs.  Marine Biology,  111(2), 247-254.
Retrieved on March 29, 2015 from [Link]

Middeljans, M. (2014). The mangrove species composition and forest structure

between managed and unmanaged Nipa palm. Retrieved April 28, 2015 from
[Link]

Miranda, S. J. M., Reyes, J. O., Tan, A., Villanueva, L. C., Zara, S. M. F., Banaag,
A. G., & Rosales, R. (2013). Mangrove Plantation as a Tourist Attraction in
San Juan Batangas, Philippines. International Journal of Academic Research in
Business and Social Sciences, 3(8), 369–375. Retrieved April 28, 2015 from
[Link]

Mittelbach, G. G., Schemske, D. W., Cornell, H. V., Allen, A. P., Brown, J. M.,
Bush, M. B., ... & Turelli, M. (2007). Evolution and the latitudinal diversity
gradient: speciation, extinction and biogeography. Ecology letters, 10(4), 315-
331. Retrieved on December 19, 2014 from [Link]

Morrisey, D. J., Swales, A., Dittmann, S., Morrison, M. A., Lovelock, C. E., &
Beard, C. M. (2010). The ecology and management of temperate mangroves(Vol.
48, pp. 43-160). R. N. Gibson (Ed.). CRC Press, Boca Raton(USA). Retrieved
on April 4, 2015 from [Link]

Mukherjee, N., Sutherland, W. J., Dicks, L., Hugé, J., Koedam, N., & Dahdouh-
Guebas, F. (2014). Ecosystem Service Valuations of Mangrove Ecosystems
to Inform Decision Making and Future Valuation Exercises. PloS one, 9(9),
e107706. Retrieved on April 5, 2015 from [Link]

Mwaluma, J. (2002). Pen culture of the mud crab Scylla serrata in Mtwapa
mangrove system, Kenya. Western Indian Ocean Journal of Marine Science,1(2),
127-133. Retrieved on March 6, 2015 from [Link]

Naidoo, G. (2006). Factors contributing to dwarfing in the mangrove Avicennia

marina. Annals of Botany, 97(6), 1095-1101. Retrieved on February 10, 2015
from [Link]

103
IAMURE International Journal of Ecology and Conservation

Nerot, C., Meziane, T., Provost-Govrich, A., Rybarczyk, H., & Lee, S. Y. (2009).
Role of grapsid crabs, Parasesarma erythrodactyla, in entry of mangrove leaves
into an estuarine food web: a mesocosm study.  Marine biology,  156(11),
2343-2352. Retrieved on April 6, 2015 from [Link]

Nicholson, C. (2009). Mangroves and Crabs as Ecosystem Engineers in Zanzibar.

Retrieved on April 6, 2015 from [Link]

Nordhaus, I., Diele, K., & Wolff, M. (2009). Activity patterns, feeding and
burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high
intertidal mangrove forest in North Brazil.  Journal of Experimental Marine
Biology and Ecology, 374(2), 104-112. Retrieved on February 9, 2015 from
[Link]

Nordhaus, I., & Wolff, M. (2007). Feeding ecology of the mangrove crab Ucides
cordatus (Ocypodidae): food choice, food quality and assimilation efficiency.
Marine Biology,  151(5), 1665-1681. Retrieved on February 9, 2015 from
[Link]

Nordhaus, I., Wolff, M., & Diele, K. (2006). Litter processing and population
food intake of the mangrove crab Ucides cordatus in a high intertidal forest
in northern Brazil.  Estuarine, Coastal and Shelf Science,  67(1), 239-250.
Retrieved on February 9, 2015 from [Link]

Nyári, Á. S., & Joseph, L. (2012). Evolution in Australasian mangrove

forests: multilocus phylogenetic analysis of the Gerygone warblers (Aves:
Acanthizidae). PloS one, 7(2), e31840. Retrieved on March 24, 2015 from
[Link]

Olds, A. D., Albert, S., Maxwell, P. S., Pitt, K. A., & Connolly, R. M. (2013).
Mangrove‐reef connectivity promotes the effectiveness of marine reserves
across the western Pacific.  Global Ecology and Biogeography,  22(9), 1040-
1049. Retrieved on February 10, 2015 from [Link]

Osborne, K., & Smith Iii, T. J. (1990). Differential predation on mangrove

propagules in open and closed canopy forest habitats. Vegetatio, 89(1), 1-6.
Retrieved on February 10, 2015 from [Link]

104
 Volume 15 · July 2015

Paillon, C., Wantiez, L., Kulbicki, M., Labonne, M., & Vigliola, L. (2014). Extent
of Mangrove Nursery Habitats Determines the Geographic Distribution of
a Coral Reef Fish in a South-Pacific Archipelago. PloS one, 9(8), e105158.
Retrieved on February 10, 2015 from [Link]

Paital, B., & Chainy, G. B. (2012). Biology and conservation of the genus
Scylla in India subcontinent.  Journal of environmental biology/Academy of
Environmental Biology, India, 33(5), 871-879. Retrieved on April 23, 2015
from [Link]

Perera, K. S., Amarasinghe, M. D., & Somaratna, S. (2013). Vegetation Structure

and Species Distribution of Mangroves along a Soil Salinity Gradient in a
Micro Tidal Estuary on the North-western Coast of Sri Lanka.  American
Journal of Marine Science,  1(1), 7-15. Retrieved on April 23, 2015 from
[Link]

Picardal, J. P., Avila, S. T. R., Tano, M. F., & Marababol, M. S. (2012). The
Species Composition and Associated Fauna of the Mangrove Forest in
Tabukand Cabgan Islets, Palompon, Leyte, Philippines.  CNU Journal of
Higher Education,  5(1). Retrieved on April 23, 2015 from [Link]
gqFKN1

Potts, M. (1979). Nitrogen fixation (acetylene reduction) associated with

communities of heterocystous and non-heterocystous blue-green algae in
mangrove forests of Sinai.  Oecologia,  39(3), 359-373. Retrieved on March
20, 2015 from [Link]

Primavera, J. H. (1993). A critical review of shrimp pond culture in the Philippines.

Reviews in Fisheries Science, 1(2), 151–201. doi:10.1080/10641269309388539

Primavera, J. H. (2000). Integrated mangrove-aquaculture systems in Asia.

Integrated coastal zone management, 121-130. Retrieved on April 28, 2015
from [Link]

Primavera, J. . (2000). Development and conservation of Philippine mangroves:

institutional issues. Ecological Economics, 35(1), 91–106. doi:10.1016/S0921-
8009(00)00170-1

105
IAMURE International Journal of Ecology and Conservation

Primavera, J. H. (2005). Mangroves and aquaculture in Southeast Asia. Retrieved

on April 28, 2015 from [Link]

Primavera, J. H., Savaris, J. P., Bajoyo, B. E., Coching, J. D., Curnick, D. J.,
Golbeque, R. L., ... & Koldewey, H. J. (2012). Manual for community-based
mangrove rehabilitation.  London, Zoological Society of London, Mangrove
Manual Series, (1), 240. Retrieved on April 28, 2015 from [Link]
b6dQxt

Primavera, J., W. Yap, J. Savaris, R. Loma, A. Moscoso, J. Coching, C.

Montilijao, R. Poingan, and I. Tayo. 2014. Manual on Mangrove Reversion
of Abandoned and Illegal Brackishwater Fishponds. Zoological Society of
London, Mangrove Manual Series, (2), 108. Retrieved on April 28, 2015 from
[Link]

Rabinowitz, D. (1978). Dispersal properties of mangrove propagules. Biotropica,

47-57. Retrieved on March 29, 2015 from [Link]

Rajendran, N., & Kathiresan, K. (2007). Microbial flora associated with

submerged mangrove leaf litter in India.  Revista de biología tropical,  55(2),
393-400. Retrieved on March 29, 2015 from [Link]

Rajkumar, M., Perumal, P., Prabu, V. A., Perumal, N. V., & Rajasekar, K. T.
(2009). Phytoplankton diversity in Pichavaram mangrove waters from south-
east coast of India.  J Environ Biol,  30, 489-498. Retrieved on February 2,
2015 from [Link]

Rao, M. D. S., Kaparapu, J., & Rao, M.N. (2015). Micro Algal Population in
Mangrove Habitats of the Visakhapatnam, East Coast of India. Journal of
Algal Biomass Utilization, 6(2),5–10. Retrieved on February 2, 2015 from
[Link]

Reef, R., Feller, I. C., & Lovelock, C. E. (2010). Nutrition of mangroves. Tree

Physiology, 30(9), 1148-1160. Retrieved on December 20, 2014 from http://
[Link]/QXVy1L

Rezende, R. D. S., Pinto, M. D. O., Gonçalves Jr, J. F., & Petrucio, M. M.

(2013). The effects of abiotic variables on detritus decomposition in Brazilian
subtropical mangroves.  Acta Limnologica Brasiliensia,  25(2), 158-168.

106
 Volume 15 · July 2015

Retrieved on April 1, 2015 from [Link]

Rohde, K. (1992). Latitudinal gradients in species diversity: the search for the
primary cause.  Oikos, 514-527. Retrieved on December 21, 2015 from
[Link]

Rönnbäck, P., Troell, M., Kautsky, N., & Primavera, J. H. (1999). Distribution
Pattern of Shrimps and Fish Among Avicennia andRhizophora Microhabitats
in the Pagbilao Mangroves, Philippines.  Estuarine, Coastal and Shelf
Science,48(2), 223-234. Retrieved on April 4, 2015 from [Link]
bVD8og

Rosenzweig, M. L. (1992). Species diversity gradients: we know more and less

than we thought.  Journal of Mammalogy, 715-730. Retrieved on April 1,
2015 from [Link]

Roy, K., Jablonski, D., Valentine, J. W., & Rosenberg, G. (1998). Marine
latitudinal diversity gradients: tests of causal hypotheses.  Proceedings of the
national Academy of Sciences,  95(7), 3699-3702. Retrieved on January 6,
2015 from [Link]

Sabigan, G. B., Garcia, R. G., & Race, K. N. B. (2013). The Status of Mangrove
Ecosystem at Buenavista , Looc , Romblon. International Journal of Marine
Ecology, 1(January), 52–60. doi:[Link]
v1i1.340

Saifullah, S. M., & Rasool, F. (1998). Standing crop of macroalgae in mangrove

habitat of Sandspit area, Karachi. Pakistan Journal of Marine Sciences, 7, 61-
64. Retrieved on April 2, 2015 from [Link]

Samson, M. S., & Rollon, R. N. (2011).  Mangrove revegetation potentials of

brackish-water pond areas in the Philippines. INTECH Open Access Publisher.
Retrieved on April 6, 2015 from [Link]

Satyanarayana, B., Bhanderi, P., Debry, M., Maniatis, D., Foré, F., Badgie, D.,
... & Dahdouh-Guebas, F. (2012). A socio-ecological assessment aiming
at improved forest resource management and sustainable ecotourism
development in the mangroves of Tanbi Wetland National Park, The Gambia,
West Africa. Ambio, 41(5), 513-526. Retrieved on April 1, 2015 from http://

107
IAMURE International Journal of Ecology and Conservation

[Link]/hmSbCF

Sheaves, M., & Molony, B. (2000). Short-circuit in the mangrove food chain.
Marine Ecology Progress Series,  199, 97-109. Retrieved on March 17, 2015
from [Link]

Smith, T. J., Boto, K. G., Frusher, S. D., & Giddins, R. L. (1991). Keystone
species and mangrove forest dynamics: the influence of burrowing by crabs
on soil nutrient status and forest productivity.  Estuarine, coastal and shelf
science,  33(5), 419-432. Retrieved on April 5, 2015 from [Link]
RVxeHQ

Smith III, T. J., & Duke, N. C. (1987). Physical determinants of inter-estuary

variation in mangrove species richness around the tropical coastline of
Australia.  Journal of Biogeography, 9-19. Retrieved on April 28, 2015 from
[Link]

Stocken, T., De Ryck, D. J. R., Balke, T., Bouma, T. J., Dahdouh-Guebas, F., &
Koedam, N. (2013). The role of wind in hydrochorous mangrove propagule
dispersal. Biogeosciences, 10(6), 3635-3647. Retrieved on April 3, 2015 from
[Link]

Stokes, D., Healy, T., & Mason, N. (2009). The benthic ecology of expanding
mangrove habitat, Tauranga Harbour, New Zealand. Retrieved on March 27,
2015 from [Link]

Sukardjo, S. (2004). Fisheries associated with mangrove ecosystem in Indonesia:

a view from a mangrove ecologist. Retrieved on April 5, 2015 from http://
[Link]/HBTFEu

Superales, J. B., & Zafaralla, M. T. (2008). Distribution of benthic

macroinvertebrates at Malubog Bay of Moro Gulf, Western Mindanao,
Philippines. Journal of Environmental Research and Development, 2(3), 303-
313. Retrieved on March 10, 2015 from [Link]

Tam, N. F., Vrijmoed, L. L. P., & Wong, Y. S. (1990). Nutrient dynamics associated
with leaf decomposition in a small subtropical mangrove community in Hong
Kong. Bulletin of Marine Science, 47(1), 68-78. Retrieved on March 10, 2015
from [Link]

108
 Volume 15 · July 2015

Thilagavathi, B., Varadharajan, D., Babu, A., Manoharan, J., Vijayalakshmi, S.,
& Balasubramanian, T. (2013). Distribution and diversity of macrobenthos
in different mangrove ecosystems of Tamil Nadu coast, India.  Journal of
Aquaculture Research and Development, 4(6), 12. Retrieved on Mrch 12, 2015
from [Link]

Tidwell, J. H., & Allan, G. L. (2001). Fish as food: aquaculture’s contribution.

EMBO reports, 2(11), 958-963. Retrieved on April 23, 2015 from [Link]
gl/es1fOp

Unsworth, R. K., Salinas De Leon, P., Garrard, S. L., Jompa, J., Smith, D. J., &
Bell, J. J. (2008). High connectivity of Indo-Pacific seagrass fish assemblages
with mangrove and coral reef habitats.  MARINE ECOLOGY-PROGRESS
SERIES-,  353, 213. Retrieved on February 16, 2015 from [Link]
gl/9TmX6u

Valenzuela, H. Y., Bacalso, A. D., Gano, C. B., Pilones, K. D., & Picardal, J.
P. (2013). The Species Composition and Associated Flora and Fauna of the
Mangrove Forest in Badian, Cebu Island, Philippines. IAMURE International
Journal of Marine Ecology, 1(1). Retrieved on April 28, 2015 from [Link]
gl/ctdlGe

Valiela, I., Bowen, J. L., & York, J. K. (2001). Mangrove Forests: One of the
World’s Threatened Major Tropical Environments At least 35% of the area
of mangrove forests has been lost in the past two decades, losses that exceed
those for tropical rain forests and coral reefs, two other well-known threatened
environments. Bioscience, 51(10), 807-815. Retrieved on April 4, 2015 from
[Link]

Vazquez, P., Holguin, G., Puente, M. E., Lopez-Cortes, A., & Bashan, Y. (2000).
Phosphate-solubilizing microorganisms associated with the rhizosphere of
mangroves in a semiarid coastal lagoon. Biology and Fertility of Soils, 30(5-6),
460-468. Retrieved on March 28, 2015 from [Link]

Vittal, B. P. R., & Sarma, V. V. (2006). Diversity and ecology of fungi on

mangroves of Bay of Bengal region-an overview.  Indian Journal of Marine
Sciences, 35(4), 308-317. Retrieved on March 27, 2015 from [Link]
uGaMZk

109
IAMURE International Journal of Ecology and Conservation

Wolff, M., Koch, V., & Isaac, V. (2000). A trophic flow model of the Caeté
mangrove estuary (North Brazil) with considerations for the sustainable use of
its resources. Estuarine, Coastal and Shelf Science, 50(6), 789-803. Retrieved
on April 1, 2015 from [Link]

Yokoo, T., Kanou, K., Moteki, M., Kohno, H., Tongnunui, P., & Kurokura, H.
(2012). Assemblage structures and spatial distributions of small gobioid fishes
in a mangrove estuary, southern Thailand. Fisheries science, 78(2), 237-247.
Retrieved on February 13, 2015 from [Link]

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